Crop Protection 55 (2014) 16e20

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Crop Protection
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A comparison of web blight epidemics on common bean cultivars with

different growth habits
Gesimria Ribeiro Costa-Coelho a, b, Adalberto Corra Caf Filho a, Murillo Lobo Jr. b, *
a
b

University of Braslia, Plant Pathology Department, Campus Universitrio Darcy Ribeiro, 70910-900 Braslia, DF, Brazil
Embrapa Rice and Beans, Rodovia GO-462 km 12, 75375-000 Santo Antnio de Gois, GO, Brazil

a r t i c l e i n f o

a b s t r a c t

Article history:
Received 25 March 2013
Received in revised form
4 October 2013
Accepted 4 October 2013

The progress of web blight caused by Thanatephorus cucumeris on common bean (Phaseolus vulgaris)
Carioca cultivars with different growth habits was compared in a three-season study. Field trials were
conducted in 2004/2005, 2005/2006 and 2006/2007 in a naturally infested area, with 10 susceptible
cultivars from group I (determinate bush, upright plants: cvs. Iapar 81, BRS Horizonte and FT Magnco);
group II (indeterminate bush plants: cvs. BRS Pontal, and Prola) and group III (plants of indeterminate
prostrate growth: cvs. BRSMG Talism, Apor, BRS Requinte, Carioca, Carioca Precoce). Severity of web
blight was assessed weekly and the resulting progress curves were t to the monomolecular model.
Overall disease impacts were estimated by the area under the disease progress curve (AUDPC), disease
progress rate and disease onset. No signicant interaction between AUDPC and planting season was
detected. Cultivars BRS Horizonte, FT Magnco and Iapar 81, with upright architecture, short guides,
high pods and closer branches presented the lowest AUDPCs. The highest AUDPCs were found among
indeterminate lodged plants with long guides and low pods (cvs Apor and Carioca Precoce), while
indeterminate bush cultivars were tted in an intermediate rank. Differences in disease epidemics
among the three groups were conrmed by intercept and disease progress rates of linearized models.
Such parameters were signicantly lower for upright genotypes, probably because they result in late
contact between neighbor plants and consequently delay the establishment of conducive microclimate
and mycelial bridges for web blight development. Yields were strongly reduced due to the extremely
high disease levels that prevailed at all three seasons. Nevertheless, a negative correlation between grain
yield and web blight severity was signicant in 2006/2007. Despite the fact that the number of cultivars
is too small to generalize responses on escape to web blight, results evidenced that choice of plant architecture is a useful strategy for web blight management.
2013 Elsevier Ltd. All rights reserved.

Keywords:
Thanatephorus cucumeris
Rhizoctonia solani
Epidemics
Disease escape
Tropical disease

1. Introduction
Web blight, caused by the soilborne fungus Thanatephorus
cucumeris Frank (Donk) (anamorph Rhizoctonia solani Kuhn), is a
major disease on common beans (Phaseolus vulgaris L.) grown in
several Brazilian warm and humid regions. The disease is also a
limiting factor in other regions of Latin America, the Caribbean and
Africa, as effective management is hardly achieved due to the
pathogens wide host range, long-lasting soilborne inoculum and
high aggressiveness (Allen, 1987; Schwartz, 2005).
The disease has been a major limiting factor for sustainable
yields especially at small scale farming. In the lowlands of the North

* Corresponding author. Tel.: 55 62 3533 2172; fax: 55 62 3533 2100.

E-mail addresses: gesimaria.costa@embrapa.br (G.R. Costa-Coelho), cafelh@
unb.br (A.C. Caf Filho), murillo.lobo@embrapa.br (M. Lobo).
0261-2194/$ e see front matter 2013 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.cropro.2013.10.006

and Mid-West Brazilian Regions, where web blight is endemic,

yield losses can reach 100%. Even though disease conditions are not
optimal, it still causes low productivity when it occurs together
with other biotic and abiotic sources of plant stress, such as poor
quality seeds (Paula et al., 1996).
There are some reports of resistant P. vulgaris genotypes and
some breeding strategies for web blight tolerance or partial resistance, such as reduction in lesion size (Beaver et al., 2008, 2012;
Montoya, 1997). However, according to Poltronieri and Oliveira
(1990) all genotypes bred for resistance to web blight in other
Latin American regions performed as highly susceptible in the
Brazilian humid tropics. Pathogen variability and quantitative
resistance are considered major impediments for achieving high
levels of disease resistance (Godoy-Lutz et al., 2003; Gonzles et al.,
2012; Takegami et al., 2004). Consequently, to this date, no effective
resistance to web blight has been incorporated into commercial
cultivars (Takegami and Beaver, 2000).

G.R. Costa-Coelho et al. / Crop Protection 55 (2014) 16e20

Disease avoidance by upright plant and open canopy architecture have been noted (Singh and Schwartz, 2010; Takegami and
Beaver, 2000) as options for disease management, but have not
been clearly demonstrated in the eld. Moreover, even moderate
decrease in disease levels based on architectural traits could be
benecial, when combined with other management strategies such
as no-tillage cropping and fungicide sprays (Costa-Coelho et al.,
2012; Galindo et al., 1983a).
Traditional Brazilian common bean cultivars in general exhibit
indeterminate growth, with prostrate or semi-prostrate plants.
Among them, those with the Carioca grain type (beige grains with
dark brown stripes), of the Mesoamerican race of P. vulgaris, are
preferred by consumers in most parts of the country accounting for
approximately 70% of the domestic market. Therefore, those cultivars are responsible for most of the 3.1 million tons of common
beans harvested annually in approximately 3.0 million ha, which
places Brazil as the worlds largest producer and consumer of
Phaseolus beans. Despite the appreciation for cultivars with traditional plant architecture, all relevant domestic breeding programs
are currently focused on releasing cultivars with upright architecture, to facilitate mechanical harvest.
Considering that web blight epidemics are inuenced by long
periods of leaf wetness, dense canopies and plant-to-plant
infection through mycelial bridges (Schwartz, 2005), genotypes
with upright architecture may induce a less favorable environment for the disease, in comparison to those of prostrate growth.
To date, no detailed eld study on the performance of common
bean cultivars with distinct architecture and their reaction to web
blight has been done. Given the lack of information on the subject, our objective was to estimate the progress of web blight
epidemics and yield in 10 cultivars of the Carioca type of determinate bush, indeterminate bush and indeterminate prostrate
habit, for three consecutive rainy seasons in an area endemic to
the disease.
2. Materials and methods
Field trials were carried out in the 2004/05, 2005/06 and 2006/
07 growing seasons in an area naturally infested by T. cucumeris, at
the National Research Centre for Rice and Beans, in Santo Antnio
de Gois, GO, Brazil. The experimental eld (16 280 6000 S, 49 70
0000 W, altitude of 823 m) had a total area of 3600 m2 surrounded by
native forest, where weeds, such as Acanthospermum australe, were
commonly found infected by T. cucumeris (Costa et al., 2007).
Ten common bean cultivars of the Carioca type differing in
growth habit and including one cultivar with an early crop cycle,
were assessed. Cultivars were assigned to three groups according to
Laing et al. (1984). Group I gathered cultivars of upright plants or
determinate bush: cvs. Iapar 81, BRS Horizonte and FT Magnco;
Group II assembled semi-prostrate or indeterminate bush cultivars: cvs. Prola and BRS Pontal; and Group III, included indeterminate prostrate cultivars: cvs. Apor, BRS Requinte, BRSMG
Talism, Carioca, and Carioca Precoce (an early cycle genotype). The
experimental plots consisted of four 5 m rows, spaced 0.45 m apart,
at an average density of 15 plants per meter after emergence. The
experimental design was a randomized complete block with four
replicates.
Fertilization at planting consisted of 500 kg ha1 of NPK 03-1700 Zn. Potassium fertilizer was dispensed supercially, at
80 kg ha1 with KCl, immediately after planting, and nitrogen
topdressing was done at a rate of 100 kg ha1 of urea, equally
divided at 15 and 30 days after planting (DAP). Seeds were treated
just before planting with imidacloprid (200 g 100 kg1 of seeds),
carbendazim thiram (300 mL 100 kg1 of seeds) and pencicuron
(300 mL 100 kg1 seed).

17

Crops were sown yearly in the third week of December when

rain frequency was almost daily, in order to match crop development to weather conditions that favor rapid disease development.
Crops were harvested manually in late February, and yields were
estimated after adjustment of grain moisture to 13%. The experimental eld remained under fallow in between planting seasons.
Web blight severity was recorded weekly with a disease scale,
where 1 no disease symptoms, 3 up to 30% of diseased leaf area,
5 31e60% of diseased leaf area, 7 61e80% of diseased leaf area
and 9 over 80% of diseased leaf area (Van Schoonhoven and
Pastor-Corrales, 1987). Following the method recommended by
Madden et al. (2007), disease scale records were back transformed
to mid-point percentage values for the estimation of the area under
disease progress curve (AUDPC). AUDPCs were estimated according
to Shaner and Finney (1977) and compared through analysis of
variance by the GLM procedure and the Tukey test at P 0.05 for
mean separation, using the SAS 9.1 statistical package (SAS Institute, Cary, NC).
Combined web blight progress curves for each season were
adjusted to logistic and monomolecular models, where regression
residues and R2 values were used to determine the best t for the
disease curves. Models were then linearized and the respective
intercepts and disease progress rates were compared by analysis of
covariance, with the REG and MIXED procedures, as suggested by
Madden et al. (2007). Climate variables (number of rainy days,
rainfall and averages of maximum temperatures) were recorded by
an automatic weather station, located 500 m from the experimental eld.
3. Results and discussion
Severe to very severe web blight epidemics developed in all
cultivars and all three planting seasons, beneted by the wide
distribution of local primary inoculum and weather favorable for
disease development (Fig. 1, Table 1). Typical web blight symptoms
were observed in all plots, with water-soaked symptoms developing from discrete lesions or from mycelial bridges, followed by
abundant microsclerotia production and plant defoliation. Highest
disease progress rates were associated with heavy rainfall (above
20 mm) during the owering/pod lling stages (Table 1). In that
stage, secondary infections were common, as a consequence of rain
splash dispersal of microsclerotia formed on earlier diseased plants
(Galindo et al., 1983b). Such pattern was also reported by Yang et al.
(1990) for soybean (Glycine max (L.) Merr.), where profuse formation of microsclerotia was observed in frequent rainfall periods.
According to Prabhu et al. (1983), web blight incidence may occur at
any plant developmental stage, but it is usually observed in the eld
after blossom, reaching highest intensity at the full owering and
pod lling stages. In the present case, disease progress was also
favored by a plant population that formed closed rows after owering, in all treatments.
In 2004/05 (Fig. 1A), rst web blight symptoms were recorded at
43 DAP. In the second and third seasons (Fig. 1B, C), the disease
became visible soon after the primary leaves were set, at 7 DAP, but
remained stable at low severity levels until 57 and 50 DAP,
respectively, probably due to much drier and somewhat warmer
conditions at late vegetative and owering stages (Table 1). Sparser
canopy during the early phenological stages in the second and third
seasons may also have played a role in reducing web blight levels.
Disease severity was higher in the rst season than in the two
subsequent seasons (p < 0.0001). Overall, a more frequent distribution of rainfall during 2004/05 may explain the higher disease
severities recorded in this season, with fastest disease progress
rates and, ultimately, the total collapse of almost all indeterminateprostrate plants at 65 DAP.

18

G.R. Costa-Coelho et al. / Crop Protection 55 (2014) 16e20

90
80
70
60
50
40
30
20
10
0

Table 2
Area under the disease progress curve (AUDPC) and web blight progress rates on
bean cultivars of different growth habits, averaged over three cropping seasons.

90
80
70
60
50
40
30
20
10
0

16

23

30

37

44

51

58

65

FT Magnfico
Iapar 81
BRS Horizonte
BRS Pontal
Prola
Carioca
BRSMG Talism
BRS Requinte
Carioca Precoce
Apor

16

23

16

23

Growth
habita

Crop cycle

AUDPC

FT Magnco
BRS Horizonte
Iapar 81
Prola
BRS Pontal
Apor
Carioca
BRS Requinte
BRSMG Talism
Carioca Precoce

I
I
I
II
II
III
III
III
III
III

Standard
Standard
Standard
Standard
Standard
Standard
Standard
Standard
Standard
Early

105.10
114.73
113.12
121.36
134.90
194.96
130.19
133.77
122.98
200.48

C.V.%

Disease progress
rate (r)
cb
bc
c
bc
b
a
b
b
b
a

19.4

0.039648
0.042724
0.041200
0.045420
0.049874
0.054901
0.048733
0.050082
0.047017
0.067312

c
c
c
bc
bc
ab
bc
bc
bc
a

18.8

Growth habit according to Laing et al. (1984): I determinate bush,

II indeterminate bush, III indeterminate prostrate.
b
Means followed by the same letter in columns do not differ according to Tukeys
test at P 0.05.

30

37

44

51

58

65

90
80
70
60
50
40
30
20
10
0
2

Cultivar

30

37

44

51

58

65

Fig. 1. Web blight progress curves in 10 common bean cultivars, in an area naturally
infested with Thanatephorus cucumeris in three cropping seasons. A: 2004/05; B: 2005/
06; C: 2006/07. Cultivars from group I (determinate bush, upright plants); II (indeterminate bush plants) and III (plants of indeterminate prostrate growth) are respectively represented by black solid, solid gray and dotted lines.

Although disease levels differed among planting seasons, web

blight progressed at distinct rates for cultivars of Groups I, II and III
(Fig. 1). AUDPC analysis produced a similar cultivar susceptibility
ranking (Table 2) and disease progress rates clustered the different
genotypes into distinct groups (p < 0.05): Group I cultivars had the
lowest AUDPCs and smallest disease progress rates. Semi-prostrate
cultivars BRS Pontal and Prola were generally gathered in an intermediate rank, while prostrate cultivars such as Apor and
Carioca Precoce had the highest AUDPCs and disease progress rates.
Consistent results over a period of three consecutive planting seasons support the recommendation of FT Magnco, IAPAR 81 and
BRS Horizonte cultivars for areas where web blight is endemic and
weather is conducive to the disease. Although the number of

cultivars for each growth habit is small to generalize responses

concerning disease escape and determinate bush cultivars, the
tested genotypes match the standards for plant architecture reported by Faria et al. (2013). In the case of FT Magnco, IAPAR 81
and BRS Horizonte, both t in the upright group comprised of
plants with short guides, high pods, and closer branches, without
bending.
The highest disease levels found in cultivars Apor and Carioca
Precoce are probably related to their respective lodged plants with
long guides, higher leaf area, low pods and open branches, typical
of indeterminate prostrate plants, as reported by Jung et al. (1996)
and Faria et al. (2013). Such plant and canopy architecture allow for
a more humid microclimate, which in turn favors increased leaf
wetness periods and also more frequent contact among plants. The
opposite effect prevails in cultivars of upright growth, where there
is a less frequent and later plant to plant contact, reducing chances
of disease spread through mycelial bridges. The additive effects of
less frequent infections and a less humid environment are expected
to result in slower disease progress rates in Group I cultivars.
Our results support the concept of using disease escape as a
disease management strategy in common beans, as reported earlier
by Ferraz and Caf-Filho (2005) and Schwartz et al. (1987),
respectively for anthracnose (Colletotrichum lindemuthianum) and
white mold (Sclerotinia sclerotiorum). Nechet and Halfeld-Vieira
(2006) also reported signicant differences among cowpea (Vigna
unguiculata (L.) Walp.) genotypes and the severity of web blight due
to plant growth habit, with signicantly more disease in those with
a prostrate habit.
In spite of the obvious polycyclic nature of web blight, the
monomolecular model offered the best t for all disease progress

Table 1
Duration of phenological stages for common bean genotypes with normal cycle and climate variables recorded at the experimental eld at different growth stages during eld
assays in three planting seasons.
DAPa

Phenological stage

Number of rainy days

2004/05

2005/06

2006/07

2004/05

2005/06

2006/07

2004/05

2005/06

2006/07

1e35
36e50b
51e78
Total
Average

Vegetative
Flowering
Pod lling

20
12
21
53

26
01
22
49

22
08
23
53

249.8
125.4
287.7
562.9

512.9
5.8
210.5
729.2

336.4
86.0
172.9
595.3

29.4
26.8
29.5

27.7
31.1
29.5

29.1
31.1
30.0

28.6

29.4

30.1

a
b

Rainfall (mm)

Average of maximum temperatures

( C)

DAP Days after planting.

Early cycle cultivar Carioca Precoce reaches blooming and pod lling in 10 days in comparison to normal cycle cultivars.

G.R. Costa-Coelho et al. / Crop Protection 55 (2014) 16e20

19

Table 4
Grain yield of Phaseolus bean cultivars affected by web blight in an area naturally
infested with Thanatephorus cucumeris during three consecutive cropping seasons.
Cultivar

Fig. 2. Residue dispersion of naturally occurring web blight progress curves on 10

common bean cultivars, tted to the monomolecular and logistic models. The experiment was conducted in a naturally infested eld in three cropping seasons.

curves, in comparison to the logistic model, which consistently

underrated web blight severity in all three cropping seasons
(Fig. 2). Regardless of the inuence of environmental factors on
disease progress rates, all planting seasons were favorable for web
blight development, and the susceptibility ranking of the genotypes
was not affected by season. Therefore, the linearized progress curve
parameters were calculated for each growth habit group and the
combined intercepts and slopes were signicantly different among
Groups I, II and III (Table 3). Intercept and slope parameters estimated by the monomolecular model also supported the differences
between the effects of plant architecture on web blight dynamics.
Slopes, representing disease progress rates, increased consistently
and signicantly from group I cultivars to group III cultivars.
Yields were generally very low, varied among seasons (Table 4)
and harvest had to be conducted at 78 DAP, about 12 days before
normal in all three seasons to avoid total grain loss, regardless the
differences among cultivars. Disease during the pod lling stage
destroyed almost all plants and signicantly reduced yield, especially in the rst season (Table 4). Relatively higher yields in the
second and third seasons may be credited to the relatively lower
disease severities. In the second eld trial, yields ranged from 481
(Carioca) to 978 kg ha1 (BRS Pontal) and in 2006/2007 it ranged
from 414 kg ha1 (Carioca Precoce) to 936 kg ha1 (FT Magnco).
Pereira et al. (1999) also reported devastating yield impacts of web
blight on common beans in Brazil, with productivity falling from
the 446e798 kg ha1 range in the absence of disease to 61e
282 kg ha1 when web blight was present.
Because of the intrinsic variation of potential yields among
representatives of Groups I, II, and III, no clear correlation was
evident between yield and disease severity. BRS Horizonte and

Table 3
Average parameters of web blight progress curves on common bean cultivars in
three planting seasons, according to their different growth habits, and adjustment to
the linearized form of the monomolecular growth model, as in: ln [1/(1  y)] ln[1/
(1  lny0)] r t, where y is disease severity in proportion, r is disease progress rate
and t is time in days.
Growth
habita

Intercept
ln[1/(1  lny0)]

Pr >jtj

Slope (r) disease

progress rate

Pr >jtj

R2

I
II
III

161.42 ab
142.83 b
114.35 c

0.0001
0.0001
0.0022

0.0212 a
0.0227 b
0.0242 c

0.0001
0.0001
0.0001

0.49
0.58
0.58

Growth habit according to Laing et al. (1984): I determinate bush,

II indeterminate bush, III indeterminate prostrate.
b
Means followed by the same letter in columns do not differ according to Tukeys
test at P 0.05.

FT Magnco
BRS Horizonte
Iapar 81
Prola
BRS Pontal
Apor
Carioca
BRS Requinte
BRSMG Talism
Carioca Precoce
Average
C.V%

Growth
habita
I
I
I
II
II
III
III
III
III
III

Grain yield (kg ha1)

2004/05
b

93 ab
158 ab
45 b
261 a
247 ab
69 ab
47 ab
77 ab
210 ab
214.0 ab
142 B
37.0

2005/06

2006/07

606 ab
833 a
508 b
728 a
978 a
506 b
481 b
645 ab
831 a
923.0 a
704 A
24.1

936 a
730 a
647 ab
699 a
504 b
669 ab
602 ab
467 b
486 b
414.3 b
615 A
29.9

a
Growth habit according to Laing et al. (1984): I determinate bush.
II indeterminate bush. III indeterminate prostrate.
b
Means followed by the same lower case letters in columns and higher case
letters in rows do not differ signicantly according to Tukeys test at P 0.05.

Prola were the only cultivars ranking as the most productive in all
three seasons. Under the highest disease severity in 2004/2005, it
was not possible to attribute increased productivity to disease
escape, since highest yields were achieved in cultivars of all three
Groups (I, II and III). However, considering only the 2006/07
planting season, productivity was generally inversely correlated to
web blight severity (p < 0.05). This result was recorded in spite of
the inherent higher yield potential of indeterminate prostrate
plants and can be ascribed to the destructive impact of web blight
in Group III genotypes. Cultivar Carioca Precoce, an early cycle genotype had the highest yield in the 2005/06 season, probably
because its shorter growth cycle allowed for the completion of pod
lling in advance of the other cultivars, thus avoiding the consequences of higher disease levels that prevailed later in the season.
This indicates that short cycle cultivars may also provide an additional mechanism for web blight escape. It is interesting to note
that, judging from the good rank of Group I cultivar disease variables (Table 2, Fig. 1), even moderate web blight severities can
cause high yield losses in common bean (Table 4).
In conclusion, our results showed the relevance of plant architecture as an important trait for web blight management on common bean crops especially when frequent rains and favorable
temperatures prevail. Although upright plant architecture as a
single variable was not sufcient for complete disease management, the consistency of results over three-season eld trials support the recommendation of upright cultivars as a partially effective
measure for web blight management. On the other hand, the option
for prostrate indeterminate cultivars must be made with caution
and combined with other disease management practices to reduce
the expected high disease levels likely to develop under conducive
weather. Effective levels of physiological resistance to web blight
are still desirable, but are notoriously difcult to detect in the
Phaseolus germplasm and cumbersome to incorporate into commercial cultivars. While breeding for resistance to web blight remains an important goal in many Phaseolus breeding programs, the
levels of partial escape observed in the present study may be successfully combined with other disease management measures to
achieve successful integrated management of web blight and sustainable cropping of common beans.
Acknowledgments
This study was supported by the Brazilian National Research
Council for Research and Development (CNPq), grant 578604/2008-

20

G.R. Costa-Coelho et al. / Crop Protection 55 (2014) 16e20

6. First and second authors were also supported by doctorate

scholarship and research fellowship from CNPq.

References
Allen, D.J., 1987. Principal Diseases of Bean in Africa, a Study Guide. Centro Internacional de Agricultura Tropical, p. 32.
Beaver, J., Alameda, M., Rosas, J.C., 2008. Breeding beans for resistance to web blight.
Annu. Rep. Bean Improv. Coop. 51, 30e31.
Beaver, J.S., Zapata, M., Alameda, M., Porch, T.G., Rosas, J.C., 2012. Registration of
PR0401-259 and PR0650-31 dry bean germplasm lines. J. Plant Reg. 6, 81e84.
Costa, G.R., Lobo Jr., M., Caf-Filho, A.C., 2007. Acanthospermum australe hospedeiro alternativo de Thanatephorus cucumeris, agente causal da mela do feijoeiro. Fitopatol. Bras 32, 83.
Costa-Coelho, G.R., Lobo Jr., M., Caf-Filho, A.C., 2012. Epidemiologia da mela e
produtividade do feijoeiro-comum tratado com fungicidas. Summa Phytopathol. 38, 211e215.
Faria, L.C.L., Melo, P.G.S., Pereira, H.S., Del Peloso, M.J., Brs, A.J.B.P., Moreira, J.A.A.,
Carvalho, H.W.L., Melo, L.C., 2013. Genetic progress during 22 years of
improvement of carioca-type common bean in Brazil. Field Crops Res. 142, 68e
74.
Ferraz, L.C.L., Caf-Filho, A.C., 2005. Efeito da arquitetura da planta e densidade de
plantio na incidncia e severidade da antracnose do feijoeiro. Summa Phytopathol. 31, 305e310.
Galindo, J.J., Abawi, G.S., Thurston, H.D., Glvez, G., 1983a. Effect of mulching on web
blight of beans in Costa Rica. Phytopathology 73, 610e615.
Galindo, J.J., Abawi, G.S., Thurston, H.D., Glvez, G., 1983b. Source of inoculum and
development of bean web blight in Costa Rica. Plant Dis. 67, 1016e1021.
Godoy-Lutz, G., Steadman, J.R., Higgins, B., Powers, K., 2003. Genetic variation
among isolates of the web blight pathogen of common bean based on PCR-RFLP
of the ITS-rDNA region. Plant Dis. 87, 766e771.
Gonzles, N., Godoy-Lutz, G., Steadman, J.R., Higgins, B., Eskridge, K.M., 2012.
Assessing genetic diversity in the web blight pathogen Thanatephorus cucumeris
(anamorph Rhizoctonia solani) subgroups AG-1-IE and AG-1-IF with molecular markers. J. Gen. Plant Pathol. 78, 85e98.
Jung, G.W., Coyne, D.P., Skroch, P.W., Nienhuis, J., Arnaud Santana, E., Bokosi, J.,
Ariyarathne, H.M., Steadman, J.R., Beaver, J.S., Kaeppler, S.M., 1996. Molecular
markers associated with plant architecture and resistance to common blight,
web blight, and rust in common beans. J. Am. Soc. Hort. Sci. 121, 794e803.

Laing, D.R., Jones, P.G., Davis, J.H.C., 1984. Common bean (Phaseolus vulgaris L.). In:
Goldsworthy, P.R., Fisher, N.M. (Eds.), The Physiology of Tropical Field Crops.
John Wiley, New York, pp. 305e351.
Madden, L.V., Hughes, G., van den Bosch, F., 2007. The Study of Plant Disease Epidemics. APS Press, St. Paul.
Montoya, C.A., Beaver, J.S., Rodrguez, R., Miklas, P.N., Godoy-Lutz, G., 1997. Heritability of resistance to web blight in ve common bean populations. Crop Sci. 37,
780e783.
Nechet, K.L., Halfeld-Vieira, B.A., 2006. Severidade da mela (Rhizoctonia solani) em
gentipos de feijo-caupi em Roraima. Fitopatol. Bras 31, 71.
Paula Jr., T.J., Vieira, R.F., Arajo, G.A., 1996. Mela (Thanatephorus cucumeris) do
feijoeiro: comportamento de variedades em condies naturais em Leopoldina,
Minas Gerais. Summa Phytopathol. 22, 46e48.
Pereira, R.C.A., Maia, A.S.C., Pinto, E.M., 1999. Ensaio de populaes segregantes de
feijo (Phaseolus vulgaris L.) para resistncia a mela (Thanatephorus cucumeris
Frank (Donk)) no estado do Acre. In: Anais. A parceira nos anos 2000. VI
RENAFE. Goinia-Gois, pp. 343e345.
Poltronieri, L.S., Oliveira, A.F.F., 1990. Mela do feijoeiro: Alternativas de controle.
UEPAE de Belm. Circular Tcnica n 3.
Prabhu, A.S., Silva, J.F.D.A., Correa, J.R.V., Polaro, R.H., Lima, E.F., 1983. Murcha da teia
miclica do feijoeiro comum: epidemiologia e aplicao de fungicidas. Pesqui.
Agropecu. Bras. Braslia 18, 1323e1332.
Schwartz, H.F., 2005. Web blight. In: Schwartz, H.F., Steadman, J.R., Hall, R.,
Forster, R.L. (Eds.), Compendium of Bean Diseases, second ed. APS Press, St. Paul,
pp. 41e43.
Schwartz, H.F., Casciano, D.H., Asenga, J.A., Wood, D.R., 1987. Field measurement of
white mold effects upon dry beans with genetic resistance or upright plant
architecture. Crop Sci. 27, 699e702.
Shaner, G., Finney, R.E., 1977. The effect of nitrogen fertilization on the expression of
slow-mildewing resistance in Knox wheat. Phytopathology 67, 1051e1056.
Singh, S.P., Schwartz, H.P., 2010. Breeding common bean for resistance to diseases: a
review. Crop Prot. 50, 2199e2223.
Takegami, J.C., Beaver, J., 2000. Heritability of web blight resistance in common
bean. Annu. Rep. Bean Improv. Coop. 43, 43e44.
Takegami, J.C., Beaver, J.S., Godoy-Lutz, G., Echvez-Badel, R., Steadman, J.R., 2004.
Inheritance of web blight resistance in common bean. J. Agric. Univ. P. R. 88,
45e54.
Van Schoonhoven, A., Pastor-Corrales, M.A., 1987. Standard System for the Evaluation of Bean Germplasm. CIAT, Cali.
Yang, X.B., Berggren, G.T., Snow, J.P., 1990. Types of Rhizoctonia foliar blight on
soybean in Louisiana. Plant Dis. 74, 501e504.