Improved Resistance Management for Durable Disease Control: A Case Study of Phoma Stem Canker of Oilseed Rape (Brassica napus)

Specific resistance loci in plants are generally very efficient in controlling development of pathogen populations. However, because of the strong selection pressure exerted, these resistances are often not durable. The probability of a resistance breakdown in a pathosystem depends on the evolutionary potential of the pathogen which is affected by: (i) the type of resistance (monogenic and/or polygenic), (ii) the type of reproduction of the pathogen (sexual and/or asexual), (iii) the capacity of the pathogen for dispersal, (iv) the resistance deployment strategy (pyramiding of specific resistances, mixture of cultivars, spatio-temporal alternation), (v) the size of the pathogen population, which is affected by control methods and environmental conditions. We propose the concept of Integrated Avirulence Management (IAM) to enhance the durability of specific resistances. IAM involves a strategy to limit the selection pressure exerted on pathogen populations and, at the same time, reduce the size of pathogen populations by combining cultural, physical, biological or chemical methods of control. Several breakdowns of resistance specific to Leptosphaeria maculans, the causal agent of phoma stem canker have occurred in Europe and in Australia. This review paper examines control methods to limit the size of L. maculans populations and discusses how this limitation of population size can enhance the durability of specific resistances. It proposes pathways for the development of a spatially explicit model to define IAM strategies. Simulation results are presented to demonstrate the potential uses of such a model for the oilseed rape/L. maculans pathosystem.     

Methods for Sampling and Assessment in Relation to the Spatial Pattern of Phoma Stem Canker (Leptosphaeria maculans) in Oilseed Rape

A sound assessment of phoma stem canker symptoms is needed to develop epidemiological, agronomical and physiological studies on the pathosystem. A specific analysis was therefore carried out to: (i) compare four methods of crown canker assessment; (ii) test the among and within assessor repeatability of one of the methods compared; (iii) characterise the spatial pattern of the disease; and (iv) define the sample size required to achieve a given level of disease assessment precision. The methods compared examined the symptoms with different procedures and graded the plants observed into six severity classes. A disease index (DI) summarised the severity distribution observed. Examination of crown cross-sections was the most precise method for assessing crown cankers. The method was repeatable, though an 'assessor effect' was apparent. The disease generally had a random pattern although significant spatial correlations were detected for four out of the fifteen plots studied at the scales examined. A relationship between the coefficient of variation of the DI and the sample size was established, evaluated with experimental field data and exemplified for typical severity distributions.     

World-Wide Importance of Phoma Stem Canker (Leptosphaeria maculans and L. biglobosa) on Oilseed Rape (Brassica napus)

Phoma stem canker is an internationally important disease of oilseed rape (Brassica napus, canola, rapeseed), causing serious losses in Europe, Australia and North America. UK losses of €56M per season are estimated using national disease survey data and a yield loss formula. Phoma stem canker pathogen populations comprise two main species, Leptosphaeria maculans, associated with damaging stem base cankers, and Leptosphaeria biglobosa, often associated with less damaging upper stem lesions. Both major gene and quantitative trait loci mediated resistance to L. maculans have been identified in B. napus, but little is known about resistance to L. biglobosa. Leptosphaeria maculans, which has spread into areas in North America and eastern Europe where only L. biglobosa was previously identified, now poses a threat to large areas of oilseed rape production in Asia. Epidemics are initiated by air-borne ascospores; major gene resistance to initial infection by L. maculans operates in the leaf lamina of B. napus. It is not clear whether the quantitative trait loci involved in the resistance to the pathogen that can be assessed only at the end of the season operate in the leaf petioles or stems. In countries where serious phoma stem canker epidemics occur, a minimum standard for resistance to L. maculans is included in national systems for registration of cultivars. This review provides a background to a series of papers on improving strategies for managing B. napus resistance to L. maculans, which is a model system for studying genetic interactions between hemi-biotrophic pathogens and their hosts.     

Fitness Cost Associated with Loss of the AvrLm4 Avirulence Function in Leptosphaeria maculans (Phoma Stem Canker of Oilseed Rape)

Near-isogenic isolates of Leptosphaeria maculans differing at the AvrLm4 avirulence locus (AvrLm4 or avrLm4) were produced in vitro. Methods for inoculation of leaves of oilseed rape with ascospores or conidia were compared. The ‘ascospore shower’ inoculation was the most efficient method for use when inoculum is limited (e.g. ascospores produced in vitro). It was used in controlled environments to compare fitness of AvrLm4 and avrLm4 isolates at 5, 10, 15, 20 or 25 °C on leaves of oilseed rape cultivars Eurol and Darmor lacking the resistance gene Rlm4, which corresponds to AvrLm4. At all temperatures tested, AvrLm4 ascospores produced more lesions than avrLm4 ascospores. The diameters of lesions produced by AvrLm4 ascospores were greater than those of lesions produced by avrLm4 ascospores. At 15–20 °C, more lesions initiated by AvrLm4 ascospores produced pycnidia than did lesions initiated by avrLm4 ascospores. However, there were no differences between AvrLm4 and avrLm4 isolates in incubation period (from inoculation to appearance of lesions) or rate of mycelial growth in leaves from lesions towards the stems. In field experiments with winter oilseed rape cultivars lacking Rlm4, the frequency of AvrLm4 isolates increased from 5.7% at the phoma leaf lesion stage (autumn) to 20.5% at the stem canker stage (summer) during 2002/2003 and from 7.9 to 11.5% during 2003/2004 growing seasons. Results of controlled environment and field experiments indicate that avrLm4 isolates have a fitness cost compared to AvrLm4 isolates.     

Major Gene and Polygenic Resistance to Leptosphaeria maculans in Oilseed Rape (Brassica napus)

The most common and effective way to control phoma stem canker (blackleg) caused by Leptosphaeria maculans in oilseed rape (Brassica napus) is through the breeding of resistant cultivars. Race specific major genes that mediate resistance from the seedling stage have been identified in B. napus or have been introgressed from related species. Many race specific major genes have been described and some of them are probably identical in B. napus (allotetraploid AACC) and the parental species B. rapa (diploid AA). More work is needed using a set of well-characterised isolates to determine the number of different major resistance genes available. In some B. napus cultivars, there is resistance which is polygenic (mediated by Quantitative Trait Loci) and postulated to be race non-specific. Many of these major genes and Quantitative Trait Loci for resistance to L. maculans have been located on B. napus genetic maps. Genes involved in race specific and polygenic resistance are generally distinct.     

Effects of Severity and Timing of Stem Canker (Leptosphaeria maculans) Symptoms on Yield of Winter Oilseed Rape (Brassica napus) in the UK

The relationships between yield loss and incidence (% plants with stems affected) or severity (mean stem score, 0–4 scale) of stem canker in winter oilseed rape were analysed using data from experiments at Rothamsted in 1991/92, Withington in 1992/93, Boxworth in 1993/94 and Rothamsted in 1997/98. Critical point models and area under disease progress curve (AUDPC) models were better than multiple point models for describing relationships between yield (tha^–1) and incidence or severity of stem canker for the four experiments. Since yield is influenced by many factors other than disease, % yield loss was calculated and critical point models and AUDPC models relating % yield loss to stem canker were constructed. The critical point models for % yield loss on stem canker incidence for three of the four experiments were similar, but differed from that for Rothamsted in 1991/92. There were also no differences between models of % yield loss on AUDPC of both incidence and severity for these three experiments. Therefore, general models of % yield loss (L) against AUDPC of incidence (X) or severity (S) of stem canker from growth stages 4.8 to 6.4 were derived from the combined data sets for the three experiments: L=–0.76+0.0075X (R²=35%, p<0.001), L=0.26+0.53S (R²=37%, p<0.001). The relationships between % yield loss and % plants with different stem canker severity scores at different growth stages were also analysed; the greatest yield losses were generally associated with the largest severity scores, for plants assessed at the same crop growth stage, and were also associated with the early development of stem lesions. Further analyses showed that % yield loss was related to incidence or severity of both basal stem cankers and upper stem lesions in experiments at Boxworth in 1993/94 and at Rothamsted in 1997/98.     

A 10-year Survey of Populations of Leptosphaeria maculans in France Indicates a Rapid Adaptation Towards the Rlm1 Resistance Gene of Oilseed Rape

Leptosphaeria maculans, the cause of stem canker of oilseed rape (OSR), exhibits gene-for-gene interactions with its host plant. The race structure of L. maculans was assessed on the basis of the analysis of 1011 isolates collected in France between 1990 and 2000, with regards to three AVR genes, AvrLm1, AvrLm2 and AvrLm4. The effect of selection pressure, due to large-scale cropping of Rlm1 cultivars, on the evolution of races of the fungus was also evaluated. The results revealed a scarcity or complete absence of isolates harbouring AvrLm2, whereas isolates harbouring AvrLm4 were present at a variable level, that was as high as 17.2–31.2% depending on the sample year and location. When obtained from rlm1 cultivars, isolates harbouring AvrLm1 always represented more than 83% of the populations until the 1997–1998 growing season. As a consequence, the Rlm1 cultivars had been highly efficient at controlling the disease and were grown on an estimated 43.7% of the total French acreage in OSR in 1998–1999. However, the increased commercial success of Rlm1 cultivars was paralleled by a decrease in the proportion of isolates harbouring AvrLm1 in 1997–1998 and 1998–1999. This resulted in less than 13% of isolates harbouring AvrLm1 in populations being collected from rlm1 cultivars in 1999 and 2000, and contributed to the loss of efficiency of the Rlm1 resistance in the field. The present study is an illustration of one round of a `boom and bust' cycle that occurred for a pathosystem where it has never been reported before. These data and the high evolutionary potential of L. maculans are fully supportive of one pathogen species with a high risk of breaking down resistance genes in OSR and suggest that the development of integrated strategies aiming at maximising the durability of novel resistance is now a priority for this pathosystem.     

A Large-Scale Survey of Races of Leptosphaeria maculans Occurring on Oilseed Rape in France

Nine avirulence genes (AvrLm1–AvrLm9) were identified in Leptosphaeria maculans, the causal agent of stem canker of oilseed rape (OSR), combinations of which could theoretically generate up to 512 different races of the fungus. L. maculans displays a high evolutionary potential to adapt to novel resistance genes as illustrated by the Rlm1 breakdown in France, where virulent populations became prevalent within three growing seasons. An improved knowledge of the race structure of the fungal population is therefore needed to ensure a better use of available major resistance genes. The objective of this study was to characterise the L. maculans population structure in France using a large-scale, rationalised sample of isolates. Experimental fields, planted with “trap plants” harbouring no major resistance gene, were sown at 20 locations. Single-pycnidium isolates were collected from leaf lesions that developed in early autumn and 1797 isolates were genotyped at Avr loci. The frequency of AvrLm6 and AvrLm7 was higher than 99%, whereas avrLm2 and avrLm9 alleles were fixed in the population. AvrLm1, AvrLm4, AvrLm5 and AvrLm8 were polymorphic. AvrLm3 isolates were detected at a very low frequency (less than 1%). Only 11 races were identified in France, with one race prevalent, namely Av5-6-7-(8) (i.e. virulent on Rlm1, Rlm2, Rlm3, Rlm4 and Rlm9), representing around 65% of the population. Disparities between the locations sampled were evident at all scales analysed. Some virulent races, such as those harbouring avrLm5, were present before the introduction of the corresponding resistance gene in the commercial OSR crop.     

Effects of Temperature and Wetness Duration on Infection of Oilseed Rape Leaves by Ascospores of Leptosphaeria maculans (Stem Canker)

In controlled environment experiments, ascospores of Leptosphaeria maculans (stem canker) infected oilseed rape (cv. Nickel) leaves and caused phoma leaf spots at temperatures from 8°C to 24°C and leaf wetness durations from 8 h to 72 h. The conditions that produced the greatest numbers of leaf spot lesions were a leaf wetness duration of 48 h at 20°C; numbers of lesions decreased with decreasing leaf wetness duration and increasing or decreasing temperature. At 20°C with 48 h of leaf wetness, it was estimated that one out of four spores infected leaves to cause a lesion whereas with 8 h of leaf wetness only one out of 300 spores caused a lesion. As temperature increased from 8°C to 20°C, the time from inoculation to the appearance of the first lesions (a measure of the incubation period) decreased from 15 to 5 days but leaf wetness duration affected the length of the incubation period only at sub-optimal temperatures. Analyses suggested that, within the optimal ranges, there was little effect of temperature or wetness duration on incubation period expressed as degree-days; the time until appearance of 50% of the lesions was ca. 145 degree-days. A linear regression of % leaves with lesions (P_l) (square-root transformed) on % plants with lesions (P_p) accounted for 93% of the variance: P_l=1.31+0.061P_p. This relationship was also investigated in winter oilseed rape field experiments in unsprayed plots from October to April in 1995/96 (cv. Envol), 1996/97 (cv. Envol), 1997/98 (cvs Bristol and Capitol) and 1998/99 (cvs Apex, Bristol and Capitol) seasons. The linear regression of % leaves with lesions (square-root transformed) on % plants with lesions accounted for 90% of the variance and had a similar slope to the controlled environment relationship: P_l=0.81+0.051P_p. These results were used to examine relationships between the development of phoma leaf spot on plants in winter oilseed rape crops, the incubation period of L. maculans and the occurrence of infection criteria (temperature, rainfall) in the autumns of 1996, 1997 and 1998.     

Cytological responses in the hypersensitive reaction in cotyledon and stem tissues of Brassica napus after infection by Leptosphaeria maculans

A hypersensitive response (HR) occurs after infection of cotyledon, leaf and stem of Brassica napus cv. Surpass 400 by an avirulent strain of Leptosphaeria maculans. The cotyledon or the sixth true-leaf stages of plants were inoculated with pycnidiospores of L. maculans (strain UWA P11). For the first time in this specific pathosystem and its HR, we report condensation of cytoplasm, shrinkage in cell size and nuclear DNA fragmentation in cotyledon tissues, and in stem tissues, shrinkage and condensation of the cytoplasm, chromatin fragmentation and lobing of the nucleus.     

Ability of aLeptosphaeria maculans isolate to form stem cankers on Indian mustard (Brassica juncea) segregates as a single locus

Australian isolates of the blackleg fungusLeptosphaeria maculans, that form cankers on two Indian mustard (Brassica juncea) varieties (Stoke and Zaria) are described. This ability to form cankers on var. Stoke segregates as a single locus in both F₁ and backcross progeny from a cross between twoL. maculans isolates.     

Study of Resistance to Orobanche ramosa in Host (Oilseed Rape and Carrot) and Non-host (Maize) Plants

Orobanche ramosa is a parasitic Angiosperm responsible for severe yield losses in several economical crops. It is a serious threat in oilseed rape in France and Morocco and is appearing in carrot crops in Morocco. In this study, several varieties of oilseed rape and carrot were screened in order to identify resistant cultivars and to characterize the resistance mechanisms involved. All the 15 oilseed rape varieties tested were susceptible. In carrot, the varieties 'Colmar à coeur rouge' and 'Nantaise demi-longue' were susceptible, whereas 'Palaiseau' and 'Buror' were resistant. In the susceptible 'Colmar à coeur rouge' carrot no defence reactions were found and the development of the parasite inhibited carrot tap root formation. In the resistant carrot varieties, the parasite germinated, became attached to the host root but became necrotic before emergence. In 'Buror' carrot, formation of a mechanical barrier was associated with the restriction to the cortex of the parasite. In maize cv. 'Vigni', a non-host of O. ramosa, thickening of xylem vessels, cell divisions in the central cylinder and formation of an encapsulation layer were observed in association with restricted development of Orobanche haustoria.     

Strategies to prevent spread of Leptosphaeria maculans (phoma stem canker) onto oilseed rape crops in China; costs and benefits

Field experiments in Europe have shown that Chinese cultivars of winter oilseed rape ( Brassica napus ) are very susceptible to the pathogen Leptosphaeria maculans (cause of phoma stem canker). Climatic and agronomic conditions in China are suitable for L. maculans since the closely related but less damaging pathogen L. biglobosa occurs on the winter and spring oilseed rape crops there. Major gene resistance to L. maculans is not durable; when introduced into commercial oilseed rape cultivars it is rapidly rendered ineffective by changes in the pathogen population. The threat to Chinese oilseed rape production from L. maculans is illustrated by the way in which L. maculans has spread into other areas of the world where previously only L. biglobosa was present, such as Canada and Poland. Models were developed to describe the spread (in space and time) of L. maculans across Alberta province, Canada, based on survey data collected over a 15-year period. These models were used to estimate the potential spread of L. maculans across the Yangtze river oilseed rape growing areas of China and its associated costs. Short-term strategies to prevent occurrence of severe phoma stem canker epidemics in China include training of extension workers to recognise symptoms of the disease and use of PCR-based diagnostics to detect the pathogen on imported seed. Long-term strategies include the introduction of durable resistance to L. maculans into Chinese oilseed rape cultivars as a component of an integrated disease management programme. The costs of such strategies in relation to costs of a phoma stem canker epidemic are discussed.     

Major gene resistance in Brassica napus (oilseed rape) is overcome by changes in virulence of populations of Leptosphaeria maculans in France and Australia

Resistance of Brassica napus (oilseed rape, canola) conferred by three different major resistance genes has been overcome by changes in virulence of Leptosphaeria maculans populations in France and Australia. In South Australia where B. napus cultivars with major gene resistance derived from Brassica rapa ssp. sylvestris were grown extensively, resistance was rendered ineffective within 3 years of commercial release of the cultivar. Disease severity was higher on cultivars with sylvestris-derived resistance than cultivars with polygenic resistance. This Australian situation is compared to that in France, where resistance conferred by the Rlm1 gene was overcome nation-wide in 5 years under commercial cropping practices, and also where a source of resistance introgressed into B. napus from B. juncea was rendered inefficient in 3 years in experimental field plots near Rennes.     

Assessment of Partial Resistance to Powdery Mildew (Podosphaera pannosa) in a Tetraploid Rose Population Using a Spore-suspension Inoculation Method

Powdery mildew (Podosphaera pannosa) is the most important fungal disease in greenhouse roses and is in practice controlled by fungicides. The creation of novel cultivars with durable resistance to powdery mildew is highly desirable. To understand the inheritance of mildew resistance, a tetraploid rose population with a size of 181 seedlings was obtained by crossing two tetraploid genotypes each having partial resistance. The population and its parents were tested under greenhouse conditions with two well-defined monospore isolates (2 and F1) using artificial inoculation with spore suspensions. Disease score at 11 days post-inoculation, latent period and rate of symptom development were used to describe seedling resistance. The tests for both isolates exhibited a wide and significant variation among genotypes for resistance. The distribution of the genotypic means of the disease scores was continuous and showed a considerable transgression. Statistical analysis, scatter plot of disease scores for the isolates, and correlation analyses indicated that the two isolates differed in pathogenicity. The outcome of the tests showed that the inoculation assay with spore suspensions was a reliable and effective way to screen large numbers of genotypes under greenhouse conditions for genetic and breeding studies. This is the first report on spore-suspension inoculation to be used successfully in rose.     

Susceptibility of Broccoli Cultivars to Bacterial Head Rot: In Vitro Screening and the Role of Head Morphology in Resistance

Head rot is a major disease of broccoli caused by the soft rot pathogens Pseudomonas fluorescens and Erwinia carotovora. Two in vitro pathogenicity tests were evaluated as a methods to identify broccoli cultivars susceptible or resistant to bacterial head rot. One test used mature heads excised from the plant and inoculated with squares of cotton lint which had been soaked in a bacterial suspension. The other test involved stab-inoculating axenically grown seedlings. With the excised head test, susceptible cultivars showed a black soft rot, whilst less susceptible or moderately resistant cultivars showed only watersoaking, or browning and slight softening of the tissue. No cultivar was completely resistant. Ten cultivars were tested, and their susceptibility ratings corresponded with previously recorded field data, with one exception. This laboratory test could be used to screen for susceptibility to head rot in broccoli breeding programmes. The seedling test distinguished differences in aggressiveness among bacterial isolates but not cultivar susceptibility. Increasing head size correlated negatively with disease resistance. Head shape, i.e. cultivars which showed a domed shape rather than a flat shape, was positively correlated with disease resistance. Thus small domed heads are more resistant to head rot than large flat heads. Other morphological characteristics, viz. floret prominence and number, and sepal stomatal number were not correlated with host resistance.     

A Greenhouse Test for Screening Sugar Beet (Beta Vulgaris) for Resistance to Rhizoctonia Solani

Rhizoctonia solani Kühn is a serious plant pathogenic fungus, causing various types of damage to sugar beet (Beta vulgaris L.). In Europe, the disease is spreading and becoming a threat for the growing of this crop. Plant resistance seems to be the most practical and economical way to control the disease. Experiments were carried out to optimise a greenhouse procedure to screen plants of sugar beet for resistance to R. solani. In the first experiment, two susceptible accessions were evaluated for root and leaf symptoms, after being grown in seven different soil mixtures and inoculated with R. solani. The fungus infected all plants. It was concluded that leaf symptoms were not reliable for the rating of disease severity. Statistically significant differences between the soil mixtures were observed, and there were no significant differences between the two accessions. The two soil mixtures, showing the most severe disease symptoms, were selected for a second experiment, including both resistant and susceptible accessions. As in the first experiment, root symptoms were recorded using a 1–7 scale, and a significant expression of resistance was observed. The average severity of the disease in the greenhouse experiment generally was comparable with the infection in field experiments, and the ranking of the accessions was the same in the two types of experiments. It was concluded that evaluation procedures in the greenhouse could be used as a rapid assay to screen sugar beet plants for resistance to R. solani.     

北京地区粘虫对5种杀虫剂的抗药性

采用浸叶法测定了北京地区6个粘虫Mythimna separata(Walker)田间种群对5种不同类型杀虫剂的抗药性。结果表明:与相对敏感品系相比,6个田间种群对5种杀虫剂均表现出不同程度的抗性水平。其中,对氯虫苯甲酰胺(抗性倍数为1.314~4.213)、甲氨基阿维菌素苯甲酸盐(抗性倍数为1.000~4.385)和毒死蜱(抗性倍数为1.083~5.936)表现为敏感至低水平抗性;对虫螨腈(抗性倍数为1.355~20.80)和氯氟氰菊酯(抗性倍数为1.748~13.98)表现为敏感至中等水平抗性。因此,北京地区的粘虫防治应注重将氯虫苯甲酰胺、甲氨基阿维菌素苯甲酸盐和毒死蜱与虫螨腈或氯氟氰菊酯交替或轮换使用,以延缓抗药性的产生与发展。     

Biology and integrated pest management for the banana weevil Cosmopolites sordidus (Germar) (Coleoptera: Curculionidae)

The banana weevil Cosmopolites sordidus (Germar) is the most important insect pest of bananas and plantains (Musa spp.). The larvae bore in the corm, reducing nutrient uptake and weakening the stability of the plant. Attack in newly planted banana stands can lead to crop failure. In established fields, weevil damage can result in reduced bunch weights, mat die-out and shortened stand life. Damage and yield losses tend to increase with time. This paper reviews the research on the taxonomy, distribution, biology, pest status, sampling methods, and integrated pest management (IPM) of banana weevil. Salient features of the weevil's biology include nocturnal activity, long life span, limited mobility, low fecundity, and slow population growth. The adults are free living and most often associated with banana mats and cut residues. They are attracted to their hosts by volatiles, especially following damage to the plant corm. Males produce an aggregation pheromone that is attractive to both sexes. Eggs are laid in the corm or lower pseudostem. The immature stages are all passed within the host plant, mostly in the corm. The weevil's biology creates sampling problems and makes its control difficult. Most commonly, weevils are monitored by trapping adults, mark and recapture methods and damage assessment to harvested or dead plants. Weevil pest status and control options reflect the type of banana being grown and the production system. Plantains and highland bananas are more susceptible to the weevil than dessert or brewing bananas. Banana production systems range from kitchen gardens and small, low-input stands to large-scale export plantations. IPM options for banana weevils include habitat management (cultural controls), biological control, host plant resistance, botanicals, and (in some cases) chemical control. Cultural controls have been widely recommended but data demonstrating their efficacy are limited. The most important are clean planting material in new stands, crop sanitation (especially destruction of residues), agronomic methods to improve plant vigour and tolerance to weevil attack and, possibly, trapping. Tissue culture plantlets, where available, assure the farmer with weevil-free material. Suckers may be cleaned by paring, hot water treatment and/or the applications of entomopathogens, neem, or pesticides. None of these methods assure elimination of weevils. Adult weevils may also invade from nearby plantations. As a result, the benefits of clean planting material may be limited to a few crop cycles. Field surveys suggest that reduced weevil populations may be associated with high levels of crop sanitation, yet definitive studies on residue management and weevil pest status are wanting. Trapping of adult weevils with pseudostem or corm traps can reduce weevil populations, but material and labour requirements may be beyond the resources of many farmers. The use of enhanced trapping with pheromones and kairomones is currently under study. A combination of clean planting material, sanitation, and trapping is likely to provide at least partial control of banana weevil.Classical biological control of banana weevil, using natural enemies from Asia, has so far been unsuccessful. Most known arthropod natural enemies are opportunistic, generalist predators with limited efficacy. Myrmicine ants have been reported to help control the weevil in Cuba, but their effects elsewhere are unknown. Microbial control, using entomopathogenic fungi and nematodes tend to be more promising. Effective strains of microbial agents are known but economic mass production and delivery systems need further development.