Review on a particular Ascomycete that

affects cucurbits

Monosporascus cannonballus
By

Marcel Barbier
University of Florida, Plant Medicine Program
Monosporascus cannonballus, Pathogen profile

INTRODUCTION

Monosporascus cannonballus is considered the main responsible agent of causing the

disease known as sudden wilt, sudden death, root rot and/or vine decline which is a
disease known in arid and semi-arid regions worldwide (4, 7, 16). This fungus has been
reported and investigated worldwide where melon and other cucurbits are commercially
grown in large scale Arizona, California, and Texas in the United States, Guatemala,
Honduras, Japan, Taiwan, Tunisia, Spain, Saudi Arabia, and Israel (3, 11, 14, 17, 19,
21, 24, and 30).

Pivonia, et al., in 2002 reported that fields in the Arava region of southern Israel, melon
(Cucumis melo L.) and watermelon (Citrullus lanatus) crops can be totally destroyed by
the disease in later summer, while disease incidence and severity in crops grown in the
same field during the following winter-spring season mostly are lower (21). Kim et al. in
1995 reported a similar phenomenon observed in Arizona (12). Barbier in 2008 reported
in Guatemala differences between soil temperature and relative humidity in the two
growing seasons (1). Wolff in 1996 also suggested that disease symptoms in melons
are affected by temperature stress in Texas. Pivonia, et al., suggested that soil
temperature differences between the two growing seasons could be an explanation for
these variations in incidence and severity.

Soil disinfestation by fumigation with methyl bromide has been a common and very
effective treatment for soil disinfection including soil pathogens. However, the United
Nations, through the Montreal Protocol, has signatures from over 120 countries banning
methyl bromide by the year 2015. Because methyl bromide depletes the stratospheric
ozone layer, the amount of methyl bromide produced and imported in the United States
was reduced incrementally through the Clean Air Act, and was definitely banned and
phased out in January 1, 2005. Alternatives to replace methyl bromide on controlling
Monosporascus cannonballus have not been as effective as methyl bromide was. In
1996 Martyn et al (16), reported that Metam-sodium, 1,3-dicloropropene, and a mixture
of ethylene dibromide and chloropicrin, were not effective on controlling M.
cannonballus when the products were applied alone without mixing. Since 1996,

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different products (chemical, biological and botanical) and products mixtures application
through the irrigation system, grafting, and solarization technique, has been evaluated,
without obtaining consisting results on controlling Monosporascus spp. in melon,
watermelon and other cucurbit species.

The risk that Monosporascus species continue spreading to other cucurbits cropping
areas is high.

Distribution:

Monosporascus cannonballus has been reported only members of Cucurbitaceae in

arid, hot areas. The most important hosts in the field are melon (Cucumis melo) and
watermelon (Citrullus lanatus) (18). According with the European and Mediterranean
Plant Protection Organization (EPPO), by 2010 Monosporascus cannonballus has been
reported in Europe (Italy, Norway, and Spain); Asia (India, Iran, Iraq, Israel, Japan,
Pakistan, Saudi Arabia, and Taiwan); Africa (Libya and Tunisia); and America
(Guatemala, Honduras, Mexico, United States, and Brazil). EPPO has Monosporascus
cannonballus in the Alert list formerly.

Map of areas in the world reporting Monosporascus spp. (2010)

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Monosporascus cannonballus, Pathogen profile

Monosporascus genus

During the sixty-second annual meeting of the American Phytopathological Society,

celebrated at Hot Springs, Arkansas, from the 4 th to the 8th of October, 1970 Troutman
and Matejka (28) presented the study: Three Fungi associated with cantaloupe roots in
Arizona. They reported that since 1967 they studied the causes of root rots of
cantaloupe and indentified Rhizoctonia solani and Verticillium albo-atrum together with
an unidentified fungus to be responsible of this disease. They reported that the
unidentified fungus did cause the decay of secondary roots that were typified by
numerous scattered, small, round, black bodies (today these bodies are known as
perithecium).

Was not until 1974, when Pollack and Uecker (29) described Monosporascus
cannonballus on secondary roots of Cucumis melo Lineus (cantaloupe) that was grown
in Yuma, Arizona. The report describes the development of the perithecium and the
nuclear cytology of this unusual Ascomycete and compares it with other previously
studied fungi. They found that the development and cytology of Monosporascus
cannonballus is a typical xylarious fungi in most respects, but they also reported that
there are important differences between this fungus and typical xylariaceous fungi. The
ostiole is formed by digestion of the tissues just below the apex of the perithecium,
followed by rupture of the outer wall layers of the apex. Peryphyses are not formed,
and the perithecial wall layers open by a rupture. Every ascus contains nuclei in the
extrasporic cytoplasm as well as in the spore itself, and the number of nuclei initially
included in the spore appears to vary. These characteristics were found to be atypical
to xylariaceous fungi type.

After Monosporascus cannonballus was reported, three other species have been
reported. The first one was Monosporascus eutypoides in 1976. Hawksworth and
Ciccarone (9) indicated that Uecker and Pollack (29) noted a strong resemblance to
Rechingeriella eutypoides Petrak, a species described from decayed roots of some
unidentified plant in Pakistan and authentic material of which they were able to study.
These authors considered that the two were distinct because of the larger ostiolar beaks

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Monosporascus cannonballus, Pathogen profile

and apparently bitunicate asci in R. eutypoides; Uecker and Pollack agreed with
Hawksworth and Booth (8) that R. eutypoides was not correctly placed in Rechingeriella
Petrak, a genus united with Zopfia Rabenh; by Hawksworth and Booth (8). But was von
Arx in 1976 who considered that R. eutypoides and Bitrimonospora indica belong to
Monosporascus and transferred both to this genus, and since then are synonyms of
Monosporascus eutypoides. Some years later, in 1978 Hawksworth and Booth (8)
conclude that no similar taxon is known apart from Anixiella monospora Malloch & Cain
and was certainly incorrectly placed in the genus Anixiella Saito & Minoura ex Cain, and
consider appropriate to transfer A. monospora to Monosporascus as Monosporascus
monosporus (Malloch & Cain); M. monosporus is only known from Iris rhizomes
originating from Iran. Last Monosporascus specie was reported in 2002 as
Monosporascus ibericus by Collado, González, Stchigel, Guarro and Peláez. They
reported that this fungus is a pyrenomycete that was isolated as an endophyte from
roots and stems of three plant species growing on sand flats and salt marshes in the
Ebro Delta in Spain (6). The main characteristic of this fungus is the presence of a
higher number of ascospores per ascus (up to six), compared to the other species of
the genus: M. cannonballus, M. eutypoides, and M. monosporus; M. ibericus produces
a cleistothecial ascomata with a tomentose peridium and lacks an anamorph. In addition
to the morphological data, the comparative analysis of the ITS-region sequences of
Monosporascus ibericus and the other Monosporascus spp., has supported the
recognition of the new species. A phylogenetic study based on the sequences of the
18S rDNA did not allow us to assess clearly the taxonomic position of the genus
Monosporascus, although the results indicated the genus might have affinities to the
Xylariales rather than to the Sordariales.

Monosporascus species reported up to date:

 Monosporascus cannonballus Pollack & Uecker 1974

 Monosporascus eutypoides (Petr.) v. Arx 1976
 Monosporascus monosporus (Malloch & Cain) D. Hawkswrth & Ciccarone. 1979
 Monosporascus ibericus Collado, González, Stchigel, Guarro & Peláez 2002

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Monosporascus cannonballus, Pathogen profile

Names:

Monosporascus cannonballus Pollack et Uecker

Classification:

Kingdom: Fungi; Phylum: Ascomycota; Class: Ascomycetes; Order: Sordariales; Genus:

Monosporascus; Species: Monosporascus cannonballus

Morphology:

Produces sexual spores The ascospores are The ascus has a layer of
called ascospores produced within an ascus. differentiated hyphae
around it, the perithecium
wall.

The perithecium is visible to Perithecium is imbedded in Small necrotic roots 1-3 mm

the naked eye as small or emergent on host roots. in diameter often support
black bulges in the root large numbers of
cortex. perithecium.

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Monosporascus cannonballus, Pathogen profile

Dimensions: Perithecium has a diameter between 222-568 µm; the base of the
perithecium is wide basally and measures between 74-148 µm, with a neck height that
could measure up to 148 µm. The ascus is between 50-110 by 35-50 µm. The
paraphysis is between 90-200 by 5-12.5 µm. The ascospores are between 32-47.5 µm
in diameter (31).

Main remarks between Monosporascus cannonballus and other species

Monosporascus species are: Monosporascus eutypoides as well as M. cannonballus
produced a perithecium, but M. eutypoides produced two ascospores per ascus instead
of one as happens in M. cannonballus. Monosporascus monospores and
Monosporascus ibericus produce a cleistothecium and not a perithecium, the ascospore
surface in M. monosporus has small pores on the ascospore surface.

Physiological symptoms

Aboveground

Field symptoms first reveal themselves The symptom may go disregarded if the
as underdeveloped plants. entire field is uniformly affected.

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Monosporascus cannonballus, Pathogen profile

Older crown leaves begin to turn chlorotic and then dry and necrotic within weeks
previous to harvest. The decaying of leaves advances very rapid to the end of the
vines, and end by causing the collapse of the vine. Dead of the canopy comes within 1
to 2 weeks after appearance of first foliar symptoms.

The bottom row in this

picture shows a group of
plants that received a
treatment with a botanical
fungicide that diminish
the ascospores
population. The front row
shows the control that did
not receive anything and
shows described
symptoms of
Monosporascus
cannonballus (1)
Fruit of diseased plants are smaller, may abscise from the pedicle before ripening and
have reduced sugar content. Fruit may also become sunburned due to lack of foliage
Number of fruits and size of it is directly correlated with fungus incidence and severity
(1).

Low Medium/Low Medium High

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Monosporascus cannonballus, Pathogen profile

Stem lesions are generally lacking and above ground symptoms may be confused with
other vine declines caused by Macrophomina phaseolina (charcoal rot), Didymella
bryoniae (gummy stem blight), Lasiodiplodia theobromae (Lasiodiplodia decline), and
Myrothecium roridum (Myrothecium canker) (16, 19).

Belowground Symptoms

Root lesions, root rot, loss of feeder roots and, in severely dry conditions, death of
taproot are results of Monosporascus root rot and vine decline.

Melon roots infected with Monosporascus

cannonballus showing lesions and loss of feeder
roots. Lesions are tan to red-brown. In severe cases
of Monosporascus cannonballus infection, most of
the root system may become necrotic and result in
death of the plant.
Lesions first develop as small areas of necrosis at
the joints between secondary and tertiary roots or at
the tips of young roots. Typically are dry, but with
excess of soil moisture they may appear as a wet
rot. Large, black perithecium form on dead roots and
are visible to the naked eye. The perithecium first
appear on smaller feeder roots in the first few
centimeters of soil and typically appear late in the
season.

Ruptured perithecium releasing ascospores. Shiny,

black, round ascospores are readily released from
the perithecium and are visible with a hand lens (16,
19).

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Monosporascus cannonballus, Pathogen profile

Disease Cycle and Epidemiology

Monosporascus cannonballus life cycle (1)

Infection of the roots occurs via germinating ascospores or active mycelium in the soil
(A). Barbier (1), suggest that initial infection can occur early in the season, from first
day after transplanting (cotyledons are completely unfolded and first soft not woody
roots are actively growing) up to the time inflorescence emergence (First flower initial
with elongated ovary visible on main stem). Tissue colonization (B, C) starts
immediately and when soil temperature rises is more aggressive. Tissue colonization
occurs during the flowering and production season. Perithecium formation in the roots
(D) occurs between flowering and fruit formation. The perithecium reaches it mature
stage when fruits are completely formed and start filling. The perithecium is ready to
release ascospores (E), when plant collapse happens. Ascospores are released (F, A)
after plant has collapsed and roots are dry.

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Ascospores are thought to be the primary inoculum, however their germination is rare
and the role in infection is unknown. Ascospores are believed to be the long-term
survival structures of the fungus. It is assumed that Monosporascus root rot and vine
decline is a monocyclic disease since no known asexual (anamorph) stage has been
identified (13, 16, 19).

Stanghellini et al (27) reported that ascospores of the fungus were recovered from soil
samples collected from native desert sites, and perithecia of the fungus was observed
on roots of a native plant, Lepidium lasiocarpum, growing in a native habitat.
Stanghellini concludes that Monosporascus sp. is an indigenous soilborne fungus.

Martyn (18) reports that Monosporascus cannonballus is homothallic (self-fertile) and

readily forms fertile perithecium in host root tissue and in vitro on artificial growth media.
Ascospores are thick, multi-walled spores and are extremely resistant to desiccation
and other factors. Germination of the ascospore is rare in vitro, however germination is
enhanced in situ when in the presence of root exudates from growing plants seedlings.
Soil microflora, most likely actinomycetes also are important in the germination of
ascospores in the field. Monosporascus cannonballus is adapted to hot, dry climates.
In vitro vegetative growth is optimal at 25 to 35°C (77 to 95°F), while perithecium are
formed most readily at 25 to 30°C (77 to 86°F). Monosporascus cannonballus may
survive for several days at temperatures up to 55 C°, but is killed within 90 min at 60°C.
Mycelial growth occurs over a pH range of 5 to 9, but is optimal from pH 6 to 7 and
inhibited completely at pH 4 and below. Monosporascus cannonballus appears also to
be adapted to slightly or moderately alkaline and saline soils. The fungus grows readily
on several standard laboratory growth media (e.g. potato dextrose agar, V-8 juice agar,
and water agar) and forms fertile black perithecium within 2 to 3 weeks. Perithecium are
readily visible against the light gray or dirty white mycelium.

Dissemination of Monosporascus cannonballus is unknown. It is likely that it is spread

by movement of infested soil or infected plant material. Ascospores may also be moved
via furrow water or heavy rains. Airborne spread is unlikely due to the large

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ascospores. Vegetative mycelium is effective at inhabiting decaying tissue; however

mycelium will not survive even moderate desiccation (13, 16, 19.)

Management

Management of Monosporascus cannonballus has proven to be difficult do to its heat

tolerance, thick-walled resting structures (ascospores), growing list of host plants and
the lack of genetic resistance in melons and common cultural practices that favor the
pathogen and disease development such as drip irrigation and black plastic mulch (15).

Combined practices that could help managing Monosporascus cannonballus:

 Soil treatment with pesticides: Biocides, Chemical fungicides, and/or Biological

fungicides.

 Cultural practices: Soil solarization, Grafting, Irrigation, and/or Fruit removal

Soil treatment with pesticides

Use of biocides in the soil: Soil disinfestation by fumigation with methyl bromide was a
common very effective treatment. Because methyl bromide depletes the stratospheric
ozone layer, was definitely banned and phased out in January 1, 2005. Metam-sodium,
1,3-dicloropropene, and a mixture of ethylene dibromide and chloropicrin, are been
used, but these biocides are not as effective as Methyl bromide was on controlling
Monosporascus cannonballus. Stanghellini et al, in 2003 reported that methyl iodide
injected as a hot gas is as effective as methyl bromide on controlling Monosporascus
cannonballus when using the same dose in Kg./Ha (26).

Chemical fungicides: Use of fungicides should be less expensive than fumigation.

Efficacy of different fungicides has been evaluated in vitro in laboratories, with good
results under those conditions (5, 23). Between 2000 and 2010, some selected
fungicides have been evaluated under field conditions (5, 23). Application timing,
frequency, and doses were evaluated. Cohen et al in 2000 evaluated 29 different
fungicides and out of these ones reported that fluazinam and kresoxim-methyl were the

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most effective. Pivonia et al in 2006 (22) reported that the strobilurines: azoxystrobyn,
pyraclostrobin, azoxystrobin + chlorothalonil and the imidazole prochloraz are effective
on controlling Monosporascus cannonballus. The issue regarding the use of these
products is regarding registration and cost. Fludioxonil applied at high rates is also
effective but is phytotoxic. Fluazinam, is less effective than others actually being
evaluated.

Biological fungicides: Sanz et al (25) reported that strains combination of Trichoderma

pseudokoniingii, Trichoderma viride and Trichoderma harzianum are effective on
controlling Monosporascus cannonballus in vitro. Barbier, evaluated a botanical
fungicide based on Melaleuca alternifolia and reported to be effective in controlling
Monosporascus cannonballus in vitro. Under field conditions Barbier evaluated
application timing, frequency, and doses indicating that the botanical fungicide is
effective when four applications were done during the first three weeks in the growing
stage of melon.

Cultural practices

Soil solarization

Melon collapse caused by the heat-tolerant soil-borne fungus Monosporascus

cannonballus is not controlled by current solarization technology applied to large soil
volumes because the temperatures achieved are not high enough to kill the pathogen's
ascospores (5).

Fruit removal

According to Hoon Lee study done in 2003, Monosporascus cannonballus severity on

cantaloupe roots were fruits are removed is less severe than on roots of plants were
fruits are not removed. Fruit removal results in increase root growth and carbohydrate
accumulation in the cantaloupe roots (10). Therefore, the practice of removing fruits
could be used as a cultural practice to retard the development of Monosporascus
cannonballus because a greater carbohydrate accumulation in the cantaloupe root will
occur and the symptoms will not occur as aggressive in plants without fruit removal.

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Pivonia et al (20) in 2001, confirm that plant collapse and death usually occurred during
the fruit maturation period. They reported that fruit removal from infected plants
prevented wilting, but did not prevent tylose formation and the associated increase in
the root resistance to water flow. Infected plants, showing reversible wilt symptoms
and from which fruits were removed, regained leaf turgor and remained alive. The
presence of fruits in Monosporascus cannonballus -infected plants apparently subjects
the plants to progressive water stress till they die. Fruit removal reduced leaf stomatal
conductance and increased root growth, thus enabling the plants to survive the
constraint to water uptake and translocation imposed by the pathogen, through root
destruction, tylose formation and root function.

Breeding and Grafting

Beltran et al (2), in 2008 suggest that disease control by grafting onto genus Cucurbita
seems to be related primarily by the increased resistance of its root system to infection
by M. cannonballus, and recommends the use of grafting as a disease management
measure for the disease.

Importance of Monosporascus cannonballus in the melon industry

According to Martyn (18) Monosporascus cannonballus the responsible agent causing

Sudden death, Root rot and/or Vine decline is an emerging disease worldwide, and was
not having enough attention among plant pathologists around the world.
Monosporascus cannonballus was described as a genus and as new specie by Pollack
and Uecker in 1974, but no pathogenicity trials were conducted at that time (29). The
first confirmed report of pathogenicity was from Israel in 1983 (24). Pathogenicity of
isolates from the United States was first reported in 1991 by Mertely et al. in Texas, and
the disease was named Monosporascus root rot and vine decline (19). Up to date
Monosporascus cannonballus has been reported in 18 countries, and it is very probable
that it will be reported in additional countries in the near future.

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Bibliography:

1. Barbier, M. (2008) Evaluation of BM-608 (Melaleuca alternifolia) oil extract on

controlling sudden wilt (Monosporascus cannonballus) in melon (Cucumis
melo). Zacapa, Guatemala. Unpublished research done for Biomor Israel Ltd.
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Comparative epidemiology of Monosporascus root rot and Vine decline in
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16. Martyn, R. D., & Miller, M. E. (1996). Monosporascus root rot and vine
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