Endophytic Fungi-Mediated Defense Signaling in Maize: Unraveling the Role of WRKY36 in Regulating Immunity against Spodoptera frugiperda.

Seed priming with beneficial endophytic fungi is an emerging sustainable strategy for enhancing plant resistance against insect pests. This study examined the effects of Beauvaria bassiana Bb20091317 and Metarhizium rileyi MrCDTLJ1 fungal colonization on maize growth, defence signalling, benzoxazinoid levels and gene expression. The colonization did not adversely affect plant growth but reduced larval weights of Spodoptera frugiperda. Maize leaves treated with M. rileyi exhibited higher levels of jasmonic acid, jasmonoyl-Isoleucine, salicylic acid, and indole acetic acid compared to control. B. bassiana and M. rileyi accelerated phytohormone increase upon S. frugiperda herbivory. Gene expression analysis revealed modulation of benzoxazinoid biosynthesis genes. We further elucidated the immune regulatory role of the transcription factor zmWRKY36 using virus-induced gene silencing (VIGS) in maize. zmWRKY36 positively regulates maize immunity against S. frugiperda, likely by interacting with defense-related proteins. Transient overexpression of zmWRKY36 in tobacco-induced cell death, while silencing in maize reduced chitin-triggered reactive oxygen species burst, confirming its immune function. Overall, B. bassiana and M. rileyi successfully colonized maize, impacting larval growth, defense signalling, and zmWRKY36-mediated resistance. This sheds light on maize-endophyte-insect interactions for sustainable plant protection.

Competitive interactions between invasive fall armyworm and Asian corn borer at intraspecific and interspecific level on the same feeding guild.

Interspecific competition is an important factor in the population dynamics and geographical distribution of insect populations. Fall armyworm (FAW), Spodoptera frugiperda, an invasive species, and the Asian corn borer (ACB), Ostrinia furnacalis, a native pest species are major pests on maize in China, posing a threat to maize yield and grain quality. A series of laboratory, greenhouse, and field experiments were conducted to elucidate the competitive interactions between FAW and ACB. In the laboratory experiments, FAW exhibited aggressive behaviors more frequently when compared with ACB, while the latter species exhibited defense behaviors more frequently. Higher intraspecific competition was recorded in the FAW conspecific rather than in ACB, particularly in the 6th larval stage. FAW had a higher interspecific competitive advantage through intraguild predation over ACB, resulting in partial or complete displacement ACB when initially the ratio of the 2 species was 1 : 1. The interspecific competition also had significantly influenced on the population parameters, defensive enzymes, and nutrient of these 2 species. Competitive interaction proved that the response of superoxide, catalase, and soluble protein in FAW were significantly increased, whereas the total sugar content in both species was substantially decreased. Survival rate, and the plant damage that co-infested by both species varied significantly among the sequential combinations under greenhouse and field conditions. FAW consistently exhibited stronger intraspecific aggression than ACB under laboratory and field conditions when co-existing on the same feeding guild. These findings contribute to efforts toward the improvement of integrated pest management programs for FAW, in decision making for invasive and native pests' management strategies to reduce the high risks of FAW and ACB outbreaks.

Management of Spodoptera frugiperda J.E. Smith Using Recycled Virus Inoculum from Larvae Treated with Baculovirus under Field Conditions.

Fall armyworm (FAW) is a major pest of maize and causes huge losses. Chemical pesticides are the commonly used control strategy among farmers. The efficacy of baculoviruses against FAW has been proven; however, farmers may not be able to afford the products. The use of farmer-produced baculovirus mixtures could provide an opportunity for a nature-based solution for FAW at a low cost. This study evaluated the potential of recycled virus inoculum from FAW larvae treated with a commercial baculovirus (Littovir) for the management of FAW under laboratory and field conditions. In the laboratory, the virus from 25, 50, 75 and 100 FAW larvae caused variable mortality among FAW instars. The highest mortality (45%) among 1st-3rd instars was caused by Littovir followed by recycled virus inoculum from 100 FAW larvae (36%). Under field conditions, even though recycled virus inoculum did not offer adequate protection against FAW damage, the maize yield was comparable to that of commercial insecticide-treated plots and similar to that of control plots. This study has shown the potential use of recycled virus inoculum from infected larvae for the management of FAW. This would offer the farmers a sustainable and affordable option for the management of FAW as it would require the farmers to purchase the commercial baculovirus once and collect larvae from treated plots for repeat applications.

Assessment of the socio-economic impacts associated with the arrival of apple snail (Pomacea canaliculata) in Mwea irrigation scheme, Kenya.

BACKGROUND: In Kenya, rice (Oryza sativa L.) is mainly produced under irrigation by small-scale farmers. Mwea irrigation scheme (MIS) in Kirinyaga County accounts for 80-88% of rice production. Here, rice is the main source of livelihood and a source of revenue generation for the county. However, a recently established invasive freshwater snail, Pomacea canaliculata (Lamarck) (family: Ampullariidae), a species of apple snail, presents a serious threat to rice production. RESULTS: Household surveys, focus group discussions and key informant interviews highlight apple snail as a serious problem in MIS. Households that observed at least a moderate level of infestation (>20% of cultivated area) experienced significant reductions in rice yield (~14%) and net rice income (~60%). Farmers reported increased use of chemical pesticides for management of apple snail. In addition, the cost of hired labor for physical removal of egg masses and snails is resulting in substantial negative effects on net income. Farmer age, area of land owned, responsibility for decision-making, receipt of extension advice, training, and membership of a farmer organization, were all statistically significant variables to explain farmers awareness of the need for area-wide apple snail management. CONCLUSION: Strategies to limit the spread of apple snail are urgently needed. A Multi-Institutional Technical Team (MITT) has been established to spearhead management efforts and consolidate advice to farmers on how to manage apple snail. However, without action to mitigate spread, the consequences could be disastrous for rice production and food security in Kenya, and for other rice growing regions across Africa. © 2023 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Integrated management of Spodoptera frugiperda 6 years post detection in Africa: a review.

The introduction of fall armyworm (FAW) Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae) on the African continent has led to paradigm shifts in pest control in maize systems, occasioned by year-round populations. The discovery of resident parasitoid species adapting to the new pest significantly informed decision-making toward avoiding highly hazardous synthetic insecticides to control the pest. A number of biopesticides have shown promise against the fall armyworm, providing a new arsenal for the sustainable management of this invasive pest. However, a few knowledge gaps remain for a fully integrated and sustainable FAW-management approach, particularly on host-resistance potential.

Comparing, evaluating and combining statistical species distribution models and CLIMEX to forecast the distributions of emerging crop pests.

BACKGROUND: Forecasting the spread of emerging pests is widely requested by pest management agencies in order to prioritise and target efforts. Two widely used approaches are statistical Species Distribution Models (SDMs) and CLIMEX, which uses ecophysiological parameters. Each have strengths and weaknesses. SDMs can incorporate almost any environmental condition and their accuracy can be formally evaluated to inform managers. However, accuracy is affected by data availability and can be limited for emerging pests, and SDMs usually predict year-round distributions, not seasonal outbreaks. CLIMEX can formally incorporate expert ecophysiological knowledge and predicts seasonal outbreaks. However, the methods for formal evaluation are limited and rarely applied. We argue that both approaches can be informative and complementary, but we need tools to integrate and evaluate their accuracy. Here we develop such an approach, and test it by forecasting the potential global range of the tomato pest Tuta absoluta. RESULTS: The accuracy of previously developed CLIMEX and new statistical SDMs were comparable on average, but the best statistical SDM techniques and environmental data substantially outperformed CLIMEX. The ensembled approach changes expectations of T. absoluta's spread. The pest's environmental tolerances and potential range in Africa, the Arabian Peninsula, Central Asia and Australia will be larger than previous estimates. CONCLUSION: We recommend that CLIMEX be considered one of a suite of SDM techniques and thus evaluated formally. CLIMEX and statistical SDMs should be compared and ensembled if possible. We provide code that can be used to do so when employing the biomod suite of SDM techniques. © 2021 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Fall armyworm in Botswana: impacts, farmer management practices and implications for sustainable pest management.

BACKGROUND: Since first reported in Botswana, fall armyworm (FAW) continues to be a threat to crop production. This study aimed to estimate impacts of FAW on yield and farmers' livelihoods in Botswana, and to obtain data that could be extrapolated to national level. Further, farmer knowledge of the pest, management practices and pesticide use for FAW management were assessed. RESULTS: In fact, 76% of the 220 respondents had seen FAW in their farms in the 2018/2019 cropping season, affecting almost the entire and about half of cultivated area for maize and sorghum, respectively. Thus, 51% of the respondents implemented FAW control measures, with chemical pesticides (27%) being the most common management against FAW. Only 33% of respondents in 2018/2019 were food self-sufficient, as opposed to 80% in an ordinary year, with farmers who reported not to have been affected by FAW more likely to be insufficient with food (88%) compared to 60% of the farmers who reported FAW attack. Drought was ranked the major stress experienced by the famers (35%), and also showed significant yield reducing effects on maize yield with pest and diseases reported second most important. Pesticides (20%) and training on pest management (18%) were the top ranked needs by farmers interviewed. CONCLUSIONS: This study confirms the impact and threat of FAW to crop production in Botswana. Chemicals remain the go-to control option by a majority of the farmers. Other low-risk technologies exist and are proposed for adoption in the management of FAW. Of note is the acknowledgement that a single control strategy will not be effective against FAW and as such integrated pest management (IPM) on an area-wide scale is needed to achieve best results. Mass awareness, training and demonstration will be required to achieve this.

The potential global distribution of the papaya mealybug, Paracoccus marginatus, a polyphagous pest.

BACKGROUND: The papaya mealybug, Paracoccus marginatus, is a highly polyphagous invasive pest that affects more than 200 plants, many of which are of economic importance. We modelled the potential distribution of P. marginatus using CLIMEX, a process-oriented, climate-based niche model. We combined this model with spatial data on irrigation and cropping patterns to increase the real-world applicability of the model. RESULTS: The resulting model agreed with known distribution points for this pest and with broad areas where P. marginatus has been reported, but for which no GPS data were available. Our model highlights the potential expansion of P. marginatus into novel areas in Central and East Africa, as well as further expansion in Central America and Asia, as these areas are highly climatically suitable, and have large expanses of suitable crop hosts. It also highlights areas, such as the central and eastern states of the USA as well as the western provinces of China, that are suitable for seasonal invasions of P. marginatus. CONCLUSION: Our results offer refined resolution on areas with high potential for invasion by P. marginatus. © 2020 Society of Chemical Industry.

Understanding smallholders' responses to fall armyworm (Spodoptera frugiperda) invasion: Evidence from five African countries.

Fall armyworm (FAW) is a new invasive pest that is causing devastating effects on maize production and threatening the livelihoods of millions of poor smallholders across sub-Saharan Africa and Asia. Using unique survey data from 2356 maize-growing households in Ghana, Rwanda, Uganda, Zambia and Zimbabwe, we examined how smallholder farmers are fighting this voracious pest. In particular, we assessed the FAW management strategies used by smallholders, socio-economic factors driving the choice of the management options, the complementarities or tradeoffs among the management options, and the (un)safe pesticide use practices of farmers. Results showed that smallholder farm households have adopted a variety of cultural, physical, chemical and local options to mitigate the effects of FAW, but the use of synthetic pesticides remains the most popular option. Results from multivariate probit regressions indicated that the extensive use of synthetic pesticides is driven by household asset wealth, and access to subsidised farm inputs and extension information. We observed that farm households are using a wide range of pesticides, including highly hazardous and banned products. Unfortunately, a majority of the households do not use personal protective equipment while handling the pesticides, resulting in reports of acute pesticide-related illness. Our findings have important implications for policies and interventions aimed at promoting environmentally friendly and sustainable ways of managing invasive pests in smallholder farming systems.

Distribution of Candidatus Liberibacter species in Eastern Africa, and the First Report of Candidatus Liberibacter asiaticus in Kenya.

Huanglongbing (HLB) is a serious disease of Citrus sp. worldwide. In Africa and the Mascarene Islands, a similar disease is known as African citrus greening (ACG) and is associated with the bacterium Candidatus Liberibacter africanus (Laf). In recent years, Candidatus Liberibacter asiaticus (Las) associated with the severe HLB has been reported in Ethiopia. Thus, we aimed to identify the Liberibacter species affecting citrus, the associated vectors in Eastern Africa and their ecological distribution. We assessed the presence of generic Liberibacter in symptomatic leaf samples by quantitative PCR. Subsequently, we sequenced the 50 S ribosomal protein L10 (rplJ) gene region in samples positive for Liberibacters and identified the species by comparison with public sequence data using phylogenetic reconstruction and genetic distances. We detected generic Liberibacter in 26%, 21% and 66% of plants tested from Uganda, Ethiopia and Kenya, respectively. The rplJ sequences revealed the most prevalent Liberibacters in Uganda and Ethiopia were LafCl (22%) and Las (17%), respectively. We detected Las in Kenya for the first time from three sites in the coastal region. Finally, we modelled the potential habitat suitability of Las in Eastern Africa using MaxEnt. The projection showed large areas of suitability for the pathogen in the three countries surveyed. Moreover, the potential distribution in Eastern Africa covered important citrus-producing parts of Ethiopia, Kenya, Uganda and Tanzania, and included regions where the disease has not been reported. These findings will guide in the development of an integrated pest management strategy to ACG/HLB management in Africa.

Parasitoid Complex of Fall Armyworm, Spodoptera frugiperda, in Ghana and Benin.

Keywords: biological control; Chelonus bifoveolatus; Coccygidum luteum; Telenomus remus; Trichogramma; West Africa.

Evidence of Leaf Consumption Rate Decrease in Fall Armyworm, Spodoptera frugiperda, Larvae Parasitized by Coccygidium luteum.

Biological control is one of the best options for the sustainable management of the invasive maize pest Spodoptera frugiperda in Africa. However, there is limited knowledge of the efficacy of native natural enemies of S. frugiperda and their potential use in integrated pest management. The endoparasitoid wasp Coccygidium luteum is one of the natural enemies of S. frugiperda in Africa. This study assessed, under laboratory conditions, the effect of C. luteum on the leaf consumption rate of its host. Fifty first instar S. frugiperda larvae were exposed to C. luteum for oviposition and the maize leaf consumption rate of parasitized larvae was assessed and compared to 50 unparasitized larvae from the same cohort. Coccygidium luteum completed a generation, from egg to adult emergence, in 16.7 days. The leaf consumption rate of parasitized S. frugiperda larvae declined gradually compared to unparasitized larvae and the overall consumption reduction by parasitized S. frugiperda larvae was 89%. Our findings show that C. luteum could reduce damage caused by S. frugiperda to maize farms but, prior to its use in biological control programmes, further studies are needed to assess potential parasitism rates in field conditions and develop a cost-effective mass production system.

Farmer perception of fall armyworm (Spodoptera frugiderda J.E. Smith) and farm-level management practices in Zambia.

BACKGROUND: This paper documents farmer perceptions and management practices for fall armyworm (Spodoptera frugiderda J.E. Smith), providing a baseline for the development of sustainable pest management strategies. RESULTS: 91% of farmers correctly identified fall armyworm, and reported it as the most important maize pest during 2016/2017 cropping season, affecting nearly half of cultivated area. Estimated maize yield loss during the season, attributed to fall armyworm was 28%. A majority of farmers (60%) used pesticides for fall armyworm control, along with other cultural/physical practices - hand picking and crushing egg masses/caterpillars (36%), and application of ash/sand in the funnel (19%). Farmers used various pesticide active ingredients, and protective measures were inadequate; >50% of farmers did not use any protective measures while spraying. Significantly more male than female farmers used pesticides (P = 0.05), and the reverse was true for cultural practices. Significant maize yield differences (P = 0.001) were observed by gender, attributed to differences in utilization of production inputs/practices. At least 77% of farmers received and shared agricultural advice, which can be optimized to spread information on fall armyworm management options. CONCLUSION: Increased use of pesticides to manage fall armyworm poses health and environmental risks, besides the high cost for farmers and governments. Research into cultural and indigenous practices used by farmers will offer opportunities for alternative and sustainable management practices. Research efforts should pay attention to gender differences in access to resources and inputs. Tackling fall armyworm at the farm level, and averting yield losses will require integrated messaging addressing other production risks. © 2019 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Telenomus Remus, a Candidate Parasitoid for the Biological Control of Spodoptera Frugiperda in Africa, is already Present on the Continent.

The fall armyworm, Spodoptera frugiperda, a moth originating from tropical and subtropical America, has recently become a serious pest of cereals in sub-Saharan Africa. Biological control offers an economically and environmentally safer alternative to synthetic insecticides that are being used for the management of this pest. Consequently, various biological control options are being considered, including the introduction of Telenomus remus, the main egg parasitoid of S. frugiperda in the Americas, where it is already used in augmentative biological control programmes. During surveys in South, West, and East Africa, parasitized egg masses of S. frugiperda were collected, and the emerged parasitoids were identified through morphological observations and molecular analyses as T. remus. The presence of T. remus in Africa in at least five countries provides a great opportunity to develop augmentative biological control methods and register the parasitoid against S. frugiperda. Surveys should be carried out throughout Africa to assess the present distribution of T. remus on the continent, and the parasitoid could be re-distributed in the regions where it is absent, following national and international regulations. Classical biological control should focus on the importation of larval parasitoids from the Americas.

Spatial Distribution of Bactrocera dorsalis and Thaumatotibia leucotreta in Smallholder Avocado Orchards along Altitudinal Gradient of Taita Hills and Mount Kilimanjaro.

Avocado (Persea americana) fruits are an important source of income and a nutritious food for small-scale growers and other stakeholders involved in farming along the Afrotropical highlands of Taita Hills and Mount Kilimanjaro in Kenya and Tanzania, respectively. Avocado fruits are infested by several insect pests, namely the Asian invasive fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae), and the false codling moth, Thaumatotibia leucotreta Meyrick (Lepidoptera: Tortricidae). However, there is inadequate information on the distribution patterns of these pests in small-scale avocado cropping systems in the East African highlands. This study was initiated to generate a spatial distribution map of B. dorsalis and T. leucotreta in avocado orchards at Taita Hills and Mount Kilimanjaro in Kenya and Tanzania, respectively. The two pests were monitored by using their respective parapheromone lures for two years between August 2012 and July 2014. Fruit damage was assessed by computing the proportion of infested fruits for B. dorsalis, whereas the damage score was used for T. leucotreta. Our results indicated that the mean number of B. dorsalis per trap per day differed significantly across elevation, being highest in lowland zone for both Taita Hills (15.90) and Mount Kilimanjaro (24.45). Similarly, the percentage infestation of ground collected fruits by B. dorsalis varied with altitude, being lowest at highlands above 1500 m.a.s.l. (0.66% and 0.83% for Taita Hills and Mount Kilimanjaro, respectively). Conversely, the mean number of T. leucotreta did not vary with altitude in either study area. However, the damage score for T. leucotreta infestation was significantly lower in the highlands of both transects (7.0% and11.1% for Taita Hills and Mount Kilimanjaro, respectively). These findings describe spatial trends that are important in formulating strategies aimed at suppressing the populations of B. dorsalis and T. leucotreta in East African avocado cropping systems.

Risk assessment and spread of the potentially invasive Ceratitis rosa Karsch and Ceratitis quilicii De Meyer, Mwatawala & Virgilio sp. Nov. using life-cycle simulation models: Implications for phytosanitary measures and management.

Integrative taxonomy has resolved the species status of the potentially invasive Ceratitis rosa Karsch into two separate species with distinct ecological requirements: C. rosa "lowland type" and the newly described species Ceratitis quilicii De Meyer, Mwatawala & Virgilio sp. nov. "highland type". Both species are tephritid pests threatening the production of horticultural crops in Africa and beyond. Studies were carried out by constructing thermal reaction norms for each life stage of both species at constant and fluctuating temperatures. Non-linear functions were fitted to continuously model species development, mortality, longevity and oviposition to establish phenology models that were stochastically simulated to estimate the life table parameters of each species. For spatial analysis of pest risk, three generic risk indices were visualized using the advanced Insect Life Cycle Modeling software. The study revealed that the highest fecundity, intrinsic rate of natural increase and net reproductive rate for C. rosa and C. quilicii was at 25 and 30°C, respectively. The resulting model successfully fits the known distribution of C. rosa and C. quilicii in Africa and the two Indian Ocean islands of La Réunion and Mauritius. Globally, the model highlights the substantial invasion risk posed by C. rosa and C. quilicii to cropping regions in the Americas, Australia, India, China, Southeast Asia, Europe, and West and Central Africa. However, the proportion of the regions predicted to be climatically suitable for both pests is narrower for C. rosa in comparison with C. quilicii, suggesting that C. quilicii will be more tolerant to a wider range of climatic conditions than C. rosa. This implies that these pests are of significant concern to biosecurity agencies in the uninvaded regions. Therefore, these findings provide important information to enhance monitoring/surveillance and designing pest management strategies to limit the spread and reduce their impact in the invaded range.

Resolution of the Identity of 'Candidatus Liberibacter' Species From Huanglongbing-Affected Citrus in East Africa.

'Candidatus Liberibacter asiaticus', the bacterium associated with citrus Huanglongbing (HLB), was reported from Uganda and tentatively from Tanzania, posing a threat to citriculture in Africa. Two surveys of citrus expressing typical HLB symptoms were conducted in Uganda, Kenya, and Tanzania to verify reports of 'Ca. L. asiaticus' and to assess the overall threat of HLB to eastern and southern African citrus production. Samples were analyzed for the presence of 'Candidatus Liberibacter' species by real-time PCR and partial sequencing of three housekeeping genes, 16S rDNA, rplJ, and omp. 'Ca. L. africanus', the bacterium historically associated with HLB symptoms in Africa, was detected in several samples. However, samples positive in real-time PCR for 'Ca. L. asiaticus' were shown not to contain 'Ca. L. asiaticus' by sequencing. Sequences obtained from these samples were analogous to 'Ca. L. africanus subsp. clausenae', identified from an indigenous Rutaceae species in South Africa, and not to 'Ca. L. asiaticus'. Results indicate a nontarget amplification of the real-time assay and suggest that previous reports of 'Ca. L. asiaticus' from Uganda and Tanzania may be mis-identifications of 'Ca. L. africanus subsp. clausenae'. This subspecies was additionally detected in individual Diaphorina citri and Trioza erytreae specimens recovered from collection sites. This is the first report of 'Ca. L. africanus subsp. clausenae' infecting citrus and being associated with HLB symptoms in this host.

Evidence for competitive displacement of Ceratitis cosyra by the invasive fruit fly Bactrocera invadens (Diptera: Tephritidae) on mango and mechanisms contributing to the displacement.

Bactrocera invadens Drew, Tsuruta & White (Diptera: Tephritidae) invaded Kenya in 2003. Before the arrival of B. invadens, the indigenous fruit fly species Ceratitis cosyra (Walker) was the predominant fruit fly pest of mango (Mangifera indica L.). Within 4 yr of invasion, B. invadens has displaced C. cosyra and has become the predominant fruit fly pest of mango, constituting 98 and 88% of the total population in traps and mango fruit at Nguruman, respectively. We tested two possible mechanisms responsible for the displacement namely; resource competition by larvae within mango fruit and aggression between adult flies. Under interspecific competition, larval duration in B. invadens was significantly shorter (6.2 +/- 0.6-7.3 +/- 0.3 d) compared with C. cosyra (8.0 +/- 1.2-9.4 +/- 0.4 d). Pupal mass in C. cosyra was affected by competition and was significantly reduced (7.4 +/- 0.3-9.6 +/- 0.6 mg) under competitive interaction compared with the controls (12.1 +/- 1.5-12.8 +/- 1.1 mg). Interspecific competition also had a significant adverse effect on C. cosyra eclosion, with fewer adults emerging under co-infestation compared with the controls. Interference competition through aggressive behavior showed that fewer C. cosyra (3.1 +/- 0.8) landed on mango dome compared with the controls (14.2 +/- 1.5) when adults were mixed with B. invadens adults in Plexiglas cages. Similarly the number of times C. cosyra was observed ovipositing was significantly lower (0.2 +/- 0.2) under competitive interaction compared with the controls (6.1 +/- 1.8). Aggressive encounters in the form of lunging/ head-butting and chasing off other species from the mango dome was higher for B. invadens compared with C. cosyra. Our results suggest that exploitative competition through larval scrambling for resources and interference competition through aggressive behaviors of the invader are important mechanisms contributing to the displacement of C. cosyra by B. invadens in mango agroecosystems.