Phylogenetic and morphological identification of Colletotrichum boninense: a novel causal agent of anthracnose in avocado

In recent years, anthracnose has become a significant disease affecting avocado fruit in the state of Michoacan, Mexico, where it significantly reduces fruit quality and commercial yield. Anthracnose has been assumed to involve Colletotrichum gloeosporioides and C. acutatum as causal agents. However, because of the increasing incidence of anthracnose, a more precise identification of the Colletotrichum spp. involved in this disease has become desirable. During the years 2004–2007, avocado fruits of different sizes exhibiting brown‐black and reddish spots on the pericarp and soft rot in the mesocarp, were gathered from orchards in nine counties. Fungal isolates were cultured on potato dextrose agar, and among these, 31 were selected for molecular, morphological and pathogenicity analyses. The molecular approaches used sequence typing of the internal transcribed spacer region and the partial nuclear large ribosomal subunit, allowing the unequivocal identification of C. gloeosporioides (71%), C. acutatum (16%) and C. boninense (13%). This last species has not been previously reported as being associated with anthracnose symptoms in avocado fruits anywhere in the world. Various morphological characteristics such as the size and shape of conidia were determined, as well as the conidial mass colour. Pathogenicity tests performed with all three species were conducted by inoculating healthy fruits. In each case, identical symptoms developed within 3 days of inoculation. Knowledge of the Colletotrichum populations in the Michoacan state, including the newly encountered avocado pathogen C. boninense, will facilitate further studies addressing the relationships between these Colletotrichum spp. and their avocado host.     

我国小麦秋苗条锈病发生规律及其区间菌源传播关系

全国冬小麦秋苗均可遭受来自西北和西南越夏区条锈菌的侵染与危害, 条锈菌在寄主上进一步繁殖和发展, 引起本地或者外地小麦条锈病流行。不同麦区秋苗条锈病发生时间、发生程度及其菌源传播规律各不相同, 年度间亦存在差异。根据小麦条锈病发生流行频率、病菌越冬和越夏情况、秋季菌源和春季菌源的有无与多少、提供时间及其影响范围与作用, 结合地理生态条件、气候特点、小麦种植区划与栽培模式等, 将中国小麦条锈病发生流行区域划分为8个明显不同的生态区系, 即关中、华北春季流行区; 成都平原、江汉流域冬季繁殖区; 西北、川西北越夏易变区; 云贵高原越夏冬繁区; 新疆冬春麦常发区; 西藏高原青稞、小麦常发区; 南方晚播冬麦偶发区; 内蒙古、东北春麦偶发区。通过病害实地调查、病菌群体遗传多样性和高空气流轨迹分析, 进一步揭示了区间菌源传播关系。陇南、陇中、陇东、宁南、海东、陕西宝鸡以及川西北和云贵高原等地区离越夏区较近, 冬小麦播种较早, 秋苗条锈病发生早、发病重, 秋季随西北气流传播到平原冬麦区和海拔较低的冬麦区侵染危害秋播麦苗, 其菌源数量对全国小麦条锈病发生流行程度起着至关重要的作用, 是中国小麦条锈病的秋季菌源基地, 面积约67万hm2; 成都平原、江汉流域、陕南、豫南、云贵坝区等麦区, 离条锈菌越夏区相对较远, 小麦播种期也较晚, 秋苗发病较晚较轻, 但冬季气候温和, 雨露条件充沛, 条锈菌在冬季可以不断侵染和繁殖, 在早春可积累大量菌源, 然后向北部和西部广大麦区扩散传播, 引起小麦条锈菌春季侵染, 是中国小麦条锈病的春季菌源基地, 面积约200万hm2。云贵高原越夏冬繁区可为我国广大麦区特别是长江中下游麦区提供部分菌源, 荆州等鄂东南地区是云贵菌源向长江下游麦区传播的中转站。云南与甘肃之间存在大量的基因流, 推测云南可能是中国小麦条锈菌重要的起源中心。西南和西北秋季菌源对长江流域麦区的相对重要性有待进一步研究明确。     

Alternative inoculum sources for fire blight: the potential role of fruit mummies and non‐host plants

Fire blight is the most damaging bacterial disease in apple production worldwide. Cankers and symptomless infected shoots are known as sites for the overwintering of Erwinia amylovora, subsequently providing primary inoculum for infection in the spring. In the present work, further potential sources of inoculum were investigated. Real‐time PCR assays covering a 3‐year‐period classified 19·9% of samples taken from fruit mummies as positive. Bacterial abundance in fruit mummies during autumn, winter and spring was up to 10⁹ cells per gram of tissue and correlated well with later infection rates of blossoms. Blossoms of non‐host plants growing close to infected trees were also shown to be colonized by E. amylovora and to enable epiphytic survival and propagation of bacteria. The results indicate a potential role of fruit mummies and buds in overwintering and as a source of primary inoculum for dissemination of the pathogen early in the growing season. Non‐host blossoms may also serve as an inoculum source in the build‐up of the pathogen population. Both aspects may contribute significantly to the epidemiology of E. amylovora. The significance of infected rootstocks as an inoculum source is also discussed. Fruit mummies might be used to determine pathogen pressure in an orchard before the beginning of the blooming period.     

影响柿树炭疽菌孢子萌发、附着胞形成及致病性的环境因子

 在凹玻片上测定了不同浓度的葡萄糖溶液对柿树炭疽菌(Colletotrichum gloeosporioides)的分生孢子萌发和附着胞形成率的影响,结果表明:分生孢子萌发率随葡萄糖浓度升高而增加,但附着胞形成率下降,芽管长度增加,附着胞的直径几乎没有变化;随着时间的延长,孢子萌发率和附着胞形成率都有所增加。不同pH对分生孢子萌发和附着胞形成率的影响结果显示,分生孢子在pH 2.0~9.0的溶液中可以萌发,并产生附着胞;最适分生孢子萌发和附着胞形成的pH是在5.0~6.0。不同pH处理的致病试验结果说明。在23℃时病斑在pH 4.0~8.0条件下都可产生;致病试验结果也发现,17℃时,pH6.0处理能发病,但不形成分生孢子团,pH 5.0处理不发病;15℃时,pH 5.0和pH 6.0的处理都不能发病。温度对菌丝生长的影响显示,菌落生长最适温度是25℃左右,高温抑制菌落生长。寄主表面侵染结构扫描电镜观察结果表明,芽管长度变化很大,芽管可以纵向沿着脊或沟延伸,也可横向通过脊沟;附着胞均在沟底或近底部形成。     

Fungicidal potential of methoxylated flavones from citrus for in vitro control of Colletotrichum gloeosporioides, causal agent of anthracnose disease in tropical fruits

Four polymethoxylated flavones (3,5,6,7,3',4'-hexamethoxyflavone, 3,5,6,7,8,3',4'-heptamethoxyflavone, 5,6,7,8,4'-pentamethoxyflavone and 5,6,7,8,3',4'-hexamethoxyflavone) were isolated and characterized from cold-pressed orange oil. Their antifungal activities were evaluated against Colletotrichum gloeosporioides (Penz) Penz & Sacc, a major plant pathogen of fruits that causes significant damage to crops in tropical, sub-tropical and temperate regions. Methoxylated flavones were effective in inhibiting mycelial growth of the fungus. As flavone concentration increased, mycelial growth decreased. 5,6,7,8,3',4'-Hexamethoxyflavone completely inhibited the growth of Cgloeosporioides at a concentration of 100 microg ml(-1).     

Epidemiology,histopathology and aetiology of olive anthracnose caused by Colletotrichum acutatum and C. gloeosporioides in Portugal

Anthracnose is an important disease affecting mature olive fruits, causing significant yield losses, and poor fruit and oil quality. In Portugal, high anthracnose incidence was recorded during 2003–2007 with 41% of 908 orchards surveyed displaying disease symptoms. In another 14% of the orchards, the pathogen was recorded in symptomless plants. Disease severity was on average 36%, frequently reaching 100%. In Portugal, anthracnose is endemic to neglected orchards of susceptible cultivars, but under favourable conditions it can also severely affect less susceptible cultivars. Pathogens were genetically heterogeneous, with Colletotrichum acutatum genetic group A2 as the most frequent (80%), followed by group A4 (12%) and group A5 along with C. gloeosporioides (3–4%), while groups A3 and A6 of C. acutatum were sporadic. Important geographic variations were observed in the frequencies of these populations, accompanied by year‐to‐year populational shifts. Epidemiology and histopathology studies showed the presence of the pathogens on vegetative organs year‐round, particularly on olive leaves and branches, and on weeds. These represent inoculum reservoirs where secondary conidiation occurs, and conidia are then dispersed by spring rains reaching flowers and young fruits or by autumn rains reaching pre‐mature fruits. Unripe fruits were colonized without showing symptoms up to penetration of the cuticle, but further colonization and symptom production was completed only as fruits matured. These findings challenge current control practices, particularly the timing of fungicide treatment, and contribute to improved disease management.     

Inoculum prevalence, host infection and biological control of Colletotrichum acutatum: causal agent of blueberry anthracnose in British Columbia

To identify the causal organism of anthracnose (ripe-rot), which reduces yield and postharvest quality of blueberries grown in British Columbia, Canada, 80 isolates were recovered from diseased fruits collected from commercial blueberry fields during 2002–04 and identified as Colletotrichum acutatum using colony morphology, growth rate and species-specific PCR primers. In vitro incubation of replicated sets of inoculated detached berries at various temperatures produced infection at temperatures of 7–30°C, with an optimum at 20°C. Colletotrichum acutatum could not survive on the soil surface as mummified berries but the pathogen was detected mostly within flower buds and less so in blueberry twigs and fruit trusses. Infection of developing flower buds in May–June of the preceding growing season gave the highest inoculum recovery in the following year. Two commercial fungal biocontrol agents, Prestop ( Gliocladium catenulatum ) and PlantShield ( Trichoderma harzianum ), each reduced anthracnose development in 2003 and 2004 by up to 45% when sprayed three times onto plants between flowering and fruit ripening.     

Role of rainfall in the development of coffee berry disease in Coffea arabica caused by Colletotrichum kahawae,in Cameroon

The development of coffee berry disease (CBD) epidemics (caused by Colletotrichum kahawae) in Cameroon was monitored over two successive years (2004 and 2005) on coffee trees protected from rainfall by transparent plastic sheets and on unprotected control trees. This work was done to assess how rain affected disease development when it did not fall directly onto the coffee trees and to determine the influence of primary inoculum on the severity of CBD. Weekly observations over the 2 years showed that there were 1·1% diseased berries on coffee trees completely protected from rainfall, compared with 45% diseased berries on unprotected coffee trees. Disease severity on unprotected trees during the 2 years of the experiment was estimated at 53% diseased berries, compared with 27% on trees only protected in the first year. These results confirmed rainfall as one of the key physical factors in the development of Arabica CBD. They also provided evidence of a subsequent effect of protecting coffee trees from rainfall in 2004 on the severity of CBD in 2005. This suggested some practices that might lead to very effective cultural control of CBD in regions where severe epidemics of the disease occur.     

Screening of weed extracts for antifungal properties against Colletotrichum lagenarium,the causal agent of anthracnose in cucumber

The antifungal activity of the leaf extracts from 203 weed species was investigated by performing a bioassay using cucumber plants and Colletotrichum orbiculare. The leaf extracts from four families, namely, Urticaceae, Onagraceae, Commelinaceae, and Solanaceae, showed a relatively stronger inhibition of the anthracnose lesions in cucumber plants when compared with the other families investigated in the study. A remarkable inhibition of anthracnose infection in cucumber leaves was observed with the extracts from the following 19 weed species: Boehmeria nipononivea and Boehmeria longispica, Persicaria scabra, Ranunculus japonicus and Ranunculus sceleratus, Cardamine flexuosa, Oenothera biennis, Aeschynomene indica, Indigofera pseudo‐tinctoria, Torilis scabra, Calystegia japonica, Solanum americanum, Bidens pilosa, Gnaphalium japonicum, Kalimeris yomena, Bromus catharticus, Cynodon dactylon, Alopecurus aequalis, and Scirpus tabernaemontani. In particular, it is noteworthy that the extracts from C. dactylon, K. yomena, and S. americanum completely inhibited anthracnose infection in cucumber.     

Molecular methods for the detection and quantification of Pestalotiopsis and Neopestalotiopsis inoculum associated with macadamia

Pestalotiopsis blight and leaf spot in macadamia are caused by species of Pestalotiopsis and Neopestalotiopsis. Conidia produced by these pathogens are mostly dispersed by wind and rain splash and are the primary infecting propagules. Rapid detection and quantification of airborne conidia are essential to determine the seasonal dynamics of inoculum and critical infection periods in macadamia orchards. A quantitative PCR (qPCR) using a new primer pair and a hydrolysis probe was developed to detect and quantify airborne conidia of Pestalotiopsis and Neopestalotiopsis species. The amplification efficiency of the qPCR was 96.4% and the assay had a limit of detection of 400 fg of Pestalotiopsis/Neopestalotiopsis gDNA, which corresponds to approximately 10 conidia. The qPCR assay coupled with spore trapping was used to monitor airborne conidia dispersal under field conditions. The results showed that the assay is specific and sensitive for detecting and quantifying Pestalotiopsis/Neopestalotiopsis conidia in macadamia orchards. The detection and quantification of the pathogen inoculum will improve our understanding of disease epidemiology and the ability to manage these diseases in macadamia.     

Cytogenomic characterization of Colletotrichum kahawae,the causal agent of coffee berry disease,reveals diversity in minichromosome profiles and genome size expansion

Colletotrichum kahawae is an emerging fungal pathogen, which has recently undergone a speciation process from a generalistic ‘C. gloeosporioides species complex' background by acquiring the unique capacity to infect green coffee berries, thus causing coffee berry disease. This is a severe and widespread disease in Africa and an imminent threat to Arabica coffee cultivation in Asia and America, if the pathogen enters those continents. Genetic diversity within C. kahawae is low but notorious differences in pathogen aggressiveness have been described. This work characterized two cytogenomic traits (genome size and minichromosome profiles) of a collection of C. kahawae isolates, representing the breadth of its genetic diversity and distinct aggressiveness classes, along with closely related taxa. The results obtained constitute the first flow cytometry‐based genome size estimation in the genus Colletotrichum and show a c. 8 Mb genome size expansion between C. kahawae (79·5 Mb on average) and its closest relatives (71·3 Mb), corroborating evidence indicating that C. kahawae (i.e. the coffee berry disease pathogens) should remain as a distinct species. Results have also shown the presence of two to five minichromosomes in C. kahawae, suggesting a positive relationship between the number of minichromosomes and the level of aggressiveness of the different isolates analysed, while no correlation could be established between aggressiveness and whole genome size. Overall, these results may be the basis for the identification of pathogenicity/aggressiveness‐related factors in such minichromosomes, and may provide clues to the characterization of specific markers for aggressiveness classes.     

Effect of humidity and temperature on conidial germination and appressorium development of two Philippine isolates of the mango anthracnose pathogen Colletotrichum gloeosporioides

A comparison of rates of germination and appressorium formation by an isolate of Colletotrichum gloeosporioides on mango leaves, fruit surfaces and cellophane membranes showed that behaviour was broadly similar on all three substrates. Frequency of appressorium formation was slightly higher on cellophane membranes, and both hyaline and melanized appressoria were formed. Only melanized appressoria were formed on mango surfaces. In vitro experiments on membranes showed comparative differences in physiological behaviour with temperature for two Philippine isolates of C. gloeosporioides . The most stimulatory temperature for production of appressoria differed in isolates I-2 and I-4 (25 and 20°C, respectively). At 30°C more appressoria became melanized than at lower temperatures, but the frequency of formation of penetration pegs was highest at 25°C. Conidia of C. gloeosporioides germinated on cellophane membranes at relative humidities as low as 95%, but the percentage of conidia germinating and forming appressoria increased as the RH approached 100%. Approximately 18% of conidia of C. gloeosporioides I-2 held at 62 and 86% RH for 4 weeks retained viability, and some were capable of forming appressoria when placed at 100% RH. These results have implications for epidemiological models for disease control.     

Field studies of anthracnose symptoms and pathogen infection in different phases of the persimmon growing season

Anthracnose is the main disease of persimmon and is caused by Colletotrichum spp. The study of field epidemiology is essential for the development of efficient management of this disease. In this study, we investigated infection by Colletotrichum spp. throughout the persimmon growing season to understand the host–pathogen interactions better. We observed the production of primary inoculum of persimmon anthracnose and described how epidemics progress from secondary infections during the fruit crop season. The field study was carried out in an organic orchard with three susceptible persimmon cultivars, Fuyu, Kakimel and Jiro, for three consecutive seasons. Our results indicate that the pathogen survives in 1-year-old shoots, which are the sources of primary inoculum. Later that growing season, the inoculum reaches flowers and new shoots, developing symptoms and producing the secondary inoculum. Fruit drop was also observed, with or without symptoms of anthracnose, throughout the plant cycle. In some of the symptomless fruit, collected from the plant and from the ground where they had fallen, latent infections of Colletotrichum spp. were detected. Shoots, flowers, immature and ripened fruit remained infected throughout the growing season, producing conidia that could lead to new secondary infections within and among plants. The incidence of anthracnose in fruit at harvest and postharvest proved to be less relevant for disease management. Practices for chemical and cultural control of the disease throughout the persimmon growing season are discussed.     

Dynamics of the ascospore dispersal of Stagonosporopsis citrulli,a causal agent of gummy stem blight of cucurbits

Sexually reproduced, airborne ascospores of Stagonosporopsis citrulli may play a role in its dispersal. S. citrulli causes gummy stem blight (GSB), one of the most important foliar diseases of cucurbits. Four studies were conducted with S. citrulli to investigate for how long ascospores are released and how far they can be dispersed from a source field. In the first study, colonized watermelon debris was sampled during three seasons and samples were assayed for ascospore release. Ascospores were detected 292, 313, and 306 days after inoculation of the source. In the second study, the active release of ascospores from pseudothecia in a Petri dish was monitored for 7 days. The release of ascospores decreased by ≤90% from 1 day after the start of the assay until 7 days after. In the third study, trap plant assays were conducted to measure the dispersal gradient of ascospores up to 366 m from the source. Generally, frequency of pathogen recovery from trap plants decreased with increasing distance from the source. The ascospore dispersal data fitted the exponential model better than the power law model. In the final study, dispersal experiments were conducted under controlled conditions. The incidence of GSB decreased with increasing distance, up to 55 m, from the source. It was concluded that ascospores of S. citrulli can serve as primary inoculum for epidemics and could easily spread among fields. Debris from cucurbit crops can be the source of ascospores for up to 10 months and should be cleared expeditiously.     

SNP-based genotyping and whole-genome sequencing reveal previously unknown genetic diversity in Xanthomonas vasicola pv. musacearum,causal agent of banana xanthomonas wilt,in its presumed Ethiopian origin

For decades, Xanthomonas vasicola pv. musacearum (Xvm) has been an economically important bacterial pathogen on enset in Ethiopia. Since 2001, Xvm has also been responsible for significant losses to banana crops in several East and Central African countries, with devastating consequences for smallholder farmers. Understanding the genetic diversity within Xvm populations is essential for the smart design of transnationally reasoned, durable, and effective management practices. Previous studies have revealed limited genetic diversity in Xvm, with East African isolates from banana each falling into one of two closely related clades previously designated as sublineages SL 1 and SL 2, the former of which had also been detected on banana and enset in Ethiopia. Given the presumed origin of Xvm in Ethiopia, we hypothesized that both clades might be found in that country, along with additional genotypes not seen in Central and East African bananas. Genotyping of 97 isolates and whole-genome sequencing of 15 isolates revealed not only the presence of SL 2 in Ethiopia, but additional diversity beyond SL 1 and SL 2 in four new clades. Moreover, SL 2 was detected in the Democratic Republic of Congo, where previously SL 1 was the only clade reported. These results demonstrate a greater range of genetic diversity among Xvm isolates than previously reported, especially in Ethiopia, and further support the hypothesis that the East/Central Africa xanthomonas wilt epidemic has been caused by a restricted set of genotypes drawn from a highly diverse pathogen pool in Ethiopia.     

Monitoring the occurrence of tomato bacterial spot and range of the causal agent Xanthomonas perforans in Iran

A 2‐year comprehensive field survey was conducted across major tomato‐growing areas of Iran. Two hundred and thirty‐four tomato fields and six tomato‐producing greenhouses were surveyed for the potential presence of bacterial spot disease. Five hundred and ninety‐six tomato samples with and without symptoms were analysed. While Xanthomonas spp. were found in association with tomato plants both with and without symptoms from five surveyed counties, the bacterial spot disease was observed only in plants from three of them. Only strains isolated from plants with symptoms induced disease symptoms on tomato, while those isolated from symptomless plants caused symptoms only on cabbage and common bean. None of the isolates caused disease symptoms on pepper and eggplant. Phylogenetic analysis showed that X. perforans is the causal agent of tomato bacterial spot in Iran, although X. campestris and X. axonopodis were also associated with symptomless tomato plants. All X. perforans isolates in this study were sensitive to streptomycin, copper sulphate and copper oxychloride at concentrations of 50 mg L^?1, 200 mg L^?1 and 0.8 g L^?1, respectively. Unlike the type strain of X. perforans, isolates in this study did not produce bacteriocin against other Xanthomonas spp., nor were they detected using the usual species‐specific primer pair Bs‐XpF/Bs‐XpR. This suggests an atypical nature of X. perforans strains in Iran, which leads to the hypothesis that X. perforans strains in Iran may have a separate origin to those causing disease epidemics elsewhere. The aggregated dispersal pattern of the diseased tomato fields signifies the seedborne introduction of the pathogen into the country.     

Growth and sporulation of <Emphasis Type="Italic">Stemphylium vesicarium</Emphasis>, the causal agent of brown spot of pear,on herb plants of orchard lawns

The inoculum sources of ascospores of Pleospora allii and of conidia of its anamorph Stemphylium vesicarium were investigated in relation to the brown spot disease epidemiology on pear. Dead and living leaves of three pear varieties (Abate Fétel, Conference and William), seven grasses (Poa pratensis, Festuca rubra, Festuca ovina, Lolium perenne, Digitaria sanguinalis and Setaria glauca) and Trifolium repens, which are used in pear orchard lawns, were inoculated with conidia of Stemphylium vesicarium virulent on pear and incubated under controlled-environment. Stemphylium vesicarium was always re-isolated from dead leaves of the considered plants, but not from symptomless green or yellowish living leaves. The fungus was occasionally re-isolated from leaf segments showing unspecific necrosis. Inoculation of pear leaves with isolates from grasses demonstrated that the fungus did not lose pathogenicity. Pseudothecia, ascospores and conidia were produced on all the dead inoculated leaves; differences between specimens were found for phenology of pseudothecia, their density and size, and for the number of conidia produced. Pseudothecia were produced faster in the lawn species than in pear leaves, and their density was higher, especially for S. glauca, L. perenne and P. pratensis. Ascospore maturation and ejection was more concentrated for the pseudothecia developed on pear leaves than for those on F. ovina and S. glauca. All the lawn species produced more conidia than pear leaves.     

Epidemiology of Fusarium Infection and Deoxynivalenol Content in Winter Wheat in the Rhineland, Germany

Details of our long-term research programme concerning the epidemiology of Fusarium spp. and mycotoxin production are summarized. Evaluation of the occurrence of Fusarium spp., mainly on winter wheat (Triticum aestivum), was carried out by investigating Fusarium infection and mycotoxin contamination. Two to 15% of grains were infested during 1995–1998 at three climatologically differing localities of the Rhineland, Germany. Disease progress was accelerated by rainfall during the flowering season. The species most frequently isolated were Fusarium avenaceum, F. poae, F. culmorum and F. graminearum. The mean deoxynivalenol (DON) content varied from 19gkg^–1 (1995) to 310gkg^–1 (1998) and was not always correlated with disease severity. Organic farming systems showed lower rates of infection with ear blight and lower mycotoxin contamination than conventional farming systems.     

Overwintering and epidemiology of Puccinia dracunculina,the causal agent of rust in open tarragon fields

Rust, caused by Puccinia dracunculina, is the main foliar disease of open‐field tarragon (Artemisia dracunculus) crops in Israel. As not much is known about the biology or epidemiology of this pathogen, the long‐term objective of the current study was to accumulate the knowledge needed to develop an effective, environmentally friendly means of adequately managing the disease. Puccinia dracunculina is an autoecious brachy‐form pathogen, but it is not known whether the life cycle is completed under field conditions in Israel. Field observations and greenhouse studies revealed that although the telial stage is produced, the pathogen overwinters in the uredinial stage. In vitro experiments were used to quantify the temperature and wetness requirements for urediniospore germination and to calculate the daily duration of conducive weather (DDCW); DDCW was defined as the number of hours during which temperature ranged between 15 and 25°C and RH was >90%. Cumulative DDCW values (CDDCW) were a good predictor of disease under natural conditions in two growing seasons. Disease onset occurred when CDDCW values reached a level of 10 and the relationship between log CDDCW values and season‐long severity values (in logit) was highly significant, explaining 90·6% of the variation.