BIOLOGICAL CONTROL OF CYPERUS SPECIES
FUNGI FOR BIOLOGICAL CONTROL OF WEEDY CYPERACEAE,
WITH EMPHASIS ON PURPLE AND YELLOW NUTSEDGES
(CYPERUS ROTUNDUS AND C. ESCULENTUS)
J. Pablo Morales-Payan,1,2 Raghavan Charudattan,1 and William M. Stall2 review the opportunities for the
control of nutsedges using mycoherbicides
1 Department of Plant Pathology and 2 Horticultural Sciences Department University of Florida, PO Box 110690, Gainesville, FL, USA.
Keywords
biocontrol, bioherbicide, biological control, Cyperaceae, integrated
weed management, mycoherbicide, nutsedge, organic agriculture,
weeds.
Purple and yellow nutsedges (Cyperus rotundus and C.
esculentus) are widely distributed weeds in many tropical
and subtropical regions of the world. Nutsedge management
is difficult partly because of their ability to produce
numerous tubers that sprout readily and produce more tuber
a few weeks after sprouting, thus perpetuating the weed.
These weeds are generally considered among the worst
weeds (Doll, 2000; Holm et al., 1991), particularly in
vegetable crops, for which the availability of selective
chemical herbicides is very limited or completely lacking.
Purple nutsedge is more commonly found in irrigated, warm
regions, whereas yellow nutsedge is predominant in wet
and/or cool environments. However, both species may
coexist in subtropical regions such as northern Mexico and
the southern USA.
Nutsedge
The effects of purple nutsedge interference with vegetable
crops can be devastating. When competing season-long with
eggplant (Solanum melongena), bell pepper (Capsicum
annuum) and tomato (Lycopersicon esculentum), purple
nutsedge reduced crop yield by 28, 65, and 70%, respec-
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© 2005. Research Information Ltd. All rights reserved
Nutsedge in tomatoes
tively (Morales-Payan et al., 1997; Morales-Payan & Stall
2001, 2003, and 2004). William and Warren (1975)
reported that purple nutsedge reduced cabbage (Brassica
oleracea var. capitata) and onion (Allium cepa) yield by 35
and 89%, respectively. Yellow nutsedge interference may
also reduce crop yields drastically. Yellow nutsedge interfer-
ence reduced the yield of asparagus (Asparagus officinalis)
(Agamalian, 1996) by 16%, eggplant by 20% (Morales-
Payan & Stall, 2004), onion by 45% (Keeling et al., 1990),
tomato by 65% (Morales-Payan et al., 2003c), and bell
pepper by 89% (Motis et al., 2001). In direct-seeded and
transplanted watermelon (Citrullus lanatus), season-long
interference from yellow nutsedge resulted in almost
complete yield loss (Buker et al., 2003).
In developing countries and in organic systems, nutsedges
are commonly managed by labor-intensive means such as
mulching, hoeing and cultivation, which may be expensive and
time consuming, may be insufficient to suppress nutsedge
and/or may need to be repeated during the season. In most
instances, shoot removal by mechanical and/or chemical
means may retard nutsedge growth and tuber production, but
repeated treatments are usually needed to suppress the weed to
a sufficient extent or to reduce its competitive ability with
crops (De la Cruz & Merayo, 1990; Keeling, 1990).
Efficacious chemical herbicides for nutsedge control may be
available for few crops (such as halosulfuron [3-chloro-5-
(4,6-dimethoxypyrimidin-2-ylcarbamoylsulfamoyl)-1-
methylpyrazole-4-carboxylic acid] for tomato), and/or
systems based on polyethylene mulching and methyl
bromide soil fumigation have been used as (so far) the only
DOI: 10.1564/16aug02
 BIOLOGICAL CONTROL OF CYPERUS SPECIES
practical and successful means for sufficient nutsedge
suppression (Chellemi, 2001). Environmental concerns
prompted the phase out of methyl bromide as an agricultural
soil fumigant in developed countries as of January 1, 2005.
Thus, the importance of nutsedges as problematic weeds is
likely to increase regardless of production system (conven-
tional or organic) in most horticultural crops infested with
those weeds. Efficacious bioherbicides for nutsedges may be a
valuable addition to current management strategies in conven-
tional production systems lacking chemical means for
nutsedge suppression, as well as in organic production
systems in which chemical herbicides are not permitted.
Biological enemies of nutsedge in nature
Many organisms have been documented as causing disease
or feeding on nutsedges in nature. Phatak et al. (1987)
published a partial list of previously reported natural
enemies of purple and yellow nutsedges, which contained
132 species of insects, 26 species of fungi, 10 species of
nematodes, 2 species of bacteria and one virus. Their list
also included farm animals such as geese (genera Anser,
Branta, and Chen), ducks (Anas spp.), and pigs (Suis spp.).
A number of organisms have been proposed as potential
agents for nutsedge control, such as Aleurocybotus sp.,
Antonina australis, Athesapeuta cyperi, Bactra minima
minima, B. verutana, B. truculenta, B. venosana, Chorizo-
coccus rostellum, Dercadothrips caespitis, Phenacoccus
solani, Rhizoecusi cacticans, Schoenabius sp., Sphenophorus
phoeniciensis (Mercado, 1979), and Puccinia canaliculata
(Phatak et al., 1987). Unfortunately, in most cases the results
of their evaluations were insufficient (partially due to a lack of
sustained research efforts), or nutsedges were never suffi-
ciently controlled to justify their commercial utilization. For
example, there has been extensive research on the use of the
moth Bactra verutana, which feeds on the shoots of nutsedge.
In numerous greenhouse and field experiments under different
environmental conditions, Bactra larvae were able to partially
destroy nutsedge shoots by feeding on them, but new shoots
sprouted from the tubers shortly afterwards, and the nutsedge
was able to produce new tubers (Frick, 1982; Frick &
Chandler, 1978; Frick & Wilson, 1981; Frick et al., 1978;
Frick et al., 1983; Porter, 1995). Similarly, in controlled
conditions, Bactra minima and B. venosana did not signifi-
cantly reduced nutsedge density (Phatak et al., 1987). In Italy,
B. furfurana, B. lancealana, B. robustana, B, venosana, and
B. bactrana were successful in partially defoliating purple
nutsedge, but tubers were undamaged and nutsedge recovered
(Trematerra & Ciampolini, 1989). Other insects feeding on
nutsedge were also pests of important crops or were crop
disease vectors (Charudattan & DeLoach, 1988) and/or were
cannibalistic and inflicted too little damage on nutsedges
(Story & Robinson, 1979), which disqualified them as useful
weed biocontrol agents.
Fungi associated with nutsedges and their
potential for biocontrol
Amongst nutsedge pathogens, fungi are the most abundant
(over 70 species) and the most studied as potential biological
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herbicides. The list of fungi genera associated with purple
and/or yellow nutsedge includes Alternaria, Ascochyta,
Balansia, Cercospora, Chaetophoma, Cintractia, Claviceps,
Cochliobulus, Corynespora, Curvularia, Dactylaria,
Dreshclera, Duosporium, Entyloma, Fusarium,
Macrophomina, Marasmius, Phaeotrichoconis, Pythium,
Phyllosticta, Phytophthora, Puccinia, Rhizopus, Sclerotinia,
Septoria, Tanatephorus, and Uredo. For some of these fungi,
there is documented research on their evaluation as potential
bioherbicides. For other fungi, only their occurrence as
nutsedge pathogens has been reported (Mello et al., 2003a;
Betria, 1973; Clay, 1986; Barreto & Evans, 1995, Evans,
1987; Kadir et al., 2000a; Prakash et al., 1996; Pomella &
Barreto, 1999; Upadhyay et. al., 1991; Welles, 1922). The
following summarizes the documented highlights,
advantages, and disadvantages of fungi in several genera to
be developed as bioherbicides.
Balansia
Inflorescence malformation and smut caused by Balansia
cyperi have been reported in purple nutsedge and green
flatsedge (C. virens) in the USA (Clay, 1986), in purple
nutsedge in the Dominican Republic (Morales-Payan et al.,
1998), and in purple nutsedge and royal flatsedge (Cyperus
elegans) in Mexico (Carrión & Chacón, 1993). Stovall and
Clay (1988) found that purple nutsedge plants inoculated
with B. cyperi produced fewer flowers and shoot biomass
than non-inoculated plants. However, inoculated plants also
produced more tubers (although smaller) than disease-free
plants. Moreover, nutsedge plants infected by B. cyperi were
less susceptible to Fusarium oxysporum, Rhizoctonia solani
and R. oryzae, and such fungal antagonism would be coun-
terproductive for weed suppression. Furthermore, fall
armyworms (Spodoptera frugiperda), which feed on
nutsedge, seemed to favor nutsedge plants not infected with
B. cyperi, as compared to nutsedge plants infected by this
fungus. Since infected nutsedge plants actually increase their
reproductive potential (more tubers) and repel other natural
enemies, there would seem to be little benefit from using B.
cyperi for purple nutsedge management.
Curvularia
De Luna et al. (1998, 2002a) reported the occurrence of
Curvularia tuberculata and C. oryzae on rice sedge
(Cyperus difformis), rice flatsedge (C. iria), and globe
fringerush (Fimbristylis miliacea) in the Philippines. When
provided with long dew period (24 hours) and high
temperature (28 C) conditions, inoculation on those three
weedy Cyperaceae resulted in rapid leaf blighting, followed
by seedling death within two weeks of inoculation. The two
fungal species were also pathogenic to purple nutsedge, but
did not cause plant death. Some rice cultivars were found to
be susceptible to some C. tuberculata isolates, but other rice
cultivars were tolerant to all the isolates tested (De Luna et
al., 2002b).
Shelby and Bewick (1991) conducted field experiments
spraying spores of Curvularia lunata for purple nutsedge
suppression in tomato. The pathogen produced typical
disease symptoms on nutsedge, but the extent of control was
not satisfactory. Moreover, C. lunata is known to cause a
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 BIOLOGICAL CONTROL OF CYPERUS SPECIES
stem disease in cassava (Manihot esculenta) (Msikita et al.,
1997), rice (Chu & Chen, 1973), and bentgrass (Agrostis
sp.) (Muchovej & Couch, 1987), which may be cause of
concern when the fungus is intended for use in those crops.
Whether the C. lunata isolate of Shelby and Bewick will
infect these crops has not been determined (the isolate could
have high specificity).
Based on the limited information available, potential
issues of host specificity and low efficacy may limit practical
applications for C. lunata. Because the humid environment
in which rice is commonly grown is bound to favor disease
progress, one of the main restraints for bioherbicide efficacy
(dew period, relative humidity) should not limit Curvularia
tuberculata as a biocontrol agent in rice. The susceptibility
of some rice cultivars may be an important issue, particu-
larly if the fungus is applied in areas where several rice
cultivars are grown in adjacent farms or fields.
Ascochyta
In India, Upadhyay et al. (1991) reported a purple nutsedge
leaf blight caused by Ascochyta cypericola. This fungus is
pathogenic only to plant species within the genus Cyperus,
and disease severity in purple nutsedge was reported to be
extensive. In addition, Stierle et al. (1991) reported that A.
cypericola produced cyperine, a phytotoxin that may have
potential as a natural herbicide (Dayan & Allen, 2000). A
related species, Ascochyta cyperiphthora sp. nov., was found
to cause leaf scorching in purple nutsedge in Brazil (Pomella
& Barreto, 1997), but its efficacy for biocontrol has not
been documented.
Information on Ascochyta spp. and/or its phytotoxin(s)
for Cyperaceae biocontrol is scarce, but what has been
reported may warrant further research. If efficacious,
selective, and economical, phytotoxins may prove more
practical than the fungi themselves, since the dew period,
relative humidity, and temperature requirements of the fungi
would be bypassed.
Puccinia
Several species in the genus Puccinia have been reported
attacking nutsedges, (predominantly yellow nutsedge) in the
USA (Phatak, 1984), Brazil (Barreto & Evans, 1995),
Panama (De la Cruz & Merayo, 1990; Esquivel, 1991),
India (Bedi & Sokhi, 1994), and Dominican Republic
(Morales-Payan et al., 1998). The species P. canaliculata and
P. romagnoliana seem to be most widespread. Researchers
such as Barreto and Evans (1995) have questioned the
taxonomic validity of several Puccinia species reported in
the literature. Barreto and Evans (1995) studied a complex
of six Puccinia rust fungi infecting purple nutsedge in Brazil.
From their taxonomic research, those authors concluded
that Puccinia conclusa, P. cypericola and P. philippensis
should be considered synonyms of P. canaliculata.
Okoli et al. (1997), using random amplified polymorphic
DNA analysis, found that previously observed differential
susceptibility of yellow nutsedge biotypes to P. canaliculata
were due to extensive genetic variability between yellow
nutsedge populations from different locations in the USA
and other countries. In contrast, there was less genetic
variability between purple nutsedge biotypes, which would
150 O u t l o o k s o n Pe s t M a n age m e n t – A u g u s t 2 0 0 5
make them more likely to respond to similar extents to
applications of P. canaliculata. Unfortunately, most purple
nutsedge biotypes exhibit a low level of susceptibility as
compared to yellow nutsedge.
Scheepens and Hoogerbrugge (1991) tested a P. canalicu-
lata strain from the USA on various Cyperaceae in the
Netherlands. Cyperus fuscus was infected by the fungus and
developed small pustules. C. esculentus leptostachyus from
several locations in the Netherlands were found to be differ-
entially susceptible to the fungus, from resistant to high
susceptibility (as interpreted from pustule number and size).
A C. esculentus selection grown as a crop and weedy C.
esculentus biotypes were resistant to the fungus. Other
Cyperaceae tested (Cyperus albostriatus, C. alternifolius, C.
flavescens, C. rotundus, Carex hirta, Eleocharis palustris,
and Scirpus maritimus) were also resistant to P. canaliculata.
When P. canaliculata spores were sprayed on yellow
nutsedge in Georgia (USA), tuber productivity was signifi-
cantly reduced, 90% of the shoots were killed within five
weeks of application, and the production of new shoots was
reduced by 66% as compared to control plants. When P.
canaliculata spores were applied with the chemical herbicide
paraquat [1,1’-dimethyl-4-4’-bypyridinium ion], yellow
nutsedge shoot production was 99% lower than in control
plants; in contrast, in plants sprayed with paraquat only,
shoot production was reduced by 10% as compared to
control plants (Phatak, 1984; Phatak et al., 1983, 1984, and
1987). Bruckart et al. (1988) combined P. canaliculata with
bentazon [3-isopropyl-1H-2,1,3-benzothiadiazin-4(3H)-one
2,2-dioxide] (0.3 or 0.6 kg a.i./ha), resulting in reduced
disease (50%) as compared to P. canaliculata without
bentazon. P. canaliculata reduced the number of tubers
(44%), but not the number of shoots produced per yellow
nutsedge plant. In yellow nutsedge infested-soybean
(Glycine max), P. canaliculata combined with the chemical
herbicides betanzon, acifluorfen [5-(2-chloro-a,a,a-trifluoro-
p-tolyloxy)-2-nitrobenzoic acid], or imazaquin [(RS)-2-(4-
isopropyl-4-methyl-5-oxo-2-imidazolin-2-yl)quinoline-3-
carboxylic acid], resulted in better yellow nutsedge control
than the rust or the chemical herbicides alone (Callaway et
al., 1986). Beste et al. (1992) applied P. canaliculata mixed
with the chemical herbicide pebulate [S-propyl
butyl(ethyl)thiocarbamate] for yellow nutsedge suppression
in tomato, and found the same extent of control with the
mixture as with pebulate alone.
Major efforts have been made to develop P. canaliculata
into a commercial bioherbicide (Phatak, 1992) and the Envi-
ronmental Protection Agency of the USA has granted P.
canaliculata a bioherbicide registration with the name of
‘Dr. BioSedge’. However, difficulties in large-scale
production of P. canaliculata spores (an obligate parasite),
its low efficacy against certain yellow nutsedge biotypes, and
its low or no pathogenicity against purple nutsedge, have
prevented wide acceptance of this fungus as a commercial
bioherbicide.
In India, Puccinia romagnoliana was found causing a rust
disease on purple nutsedge. In field trials in which this
potential bioherbicide was applied on purple nutsedge, the
rust reduced plant fresh and dry weights by 64% and 56%,
respectively, and tuber number and weight were reduced by