1 Introduction

Maize (Zea mays L.) is a globally significant cereal crop, serving as a staple food for millions of people, particularly in sub-Saharan Africa and other developing regions. However, maize production is frequently threatened by a variety of pests, including the Western Corn Rootworm (Diabrotica virgifera virgifera), fall armyworm (Spodoptera frugiperda), and various species of aphids and herbivorous arthropods (Pasquier et al., 2021; Clarkson et al., 2022).

These pests can cause substantial damage to maize crops, leading to significant yield losses and economic hardship for farmers. Traditional pest control methods, such as the use of chemical pesticides, are becoming increasingly unsustainable due to their high costs, environmental impact, and the development of pest resistance (Otieno et al., 2019).

Natural predators play a crucial role in the biological control of maize pests, offering an environmentally friendly and cost-effective alternative to chemical pesticides. Predatory beetles, mites, parasitoid wasps, and other generalist predators can significantly reduce pest populations by preying on them at various stages of their life cycles (Fang et al., 2012).

For instance, the predatory beetle Propylea japonica has been shown to effectively control aphid populations in maize and cotton fields, while the soil-dwelling predatory mite Gaeolaelaps aculeifer has demonstrated potential in managing Western Corn Rootworm larvae. The presence of diverse vegetation and intercropping systems can enhance the effectiveness of these natural predators by providing them with alternative habitats and food sources, thereby promoting their survival and predatory efficiency (Yang et al., 2020; Puliga et al., 2023).

This study evaluates the role of natural predators in controlling maize pests by analyzing their impact on the population dynamics of major maize pests such as the Western Corn Rootworm and Fall Armyworm. It also assesses the effectiveness of habitat manipulation techniques (e.g., intercropping and the use of refuge plants) in enhancing the biological control potential of natural predators, aiming to identify the key factors that influence the success of predator-mediated pest control in maize cropping systems.

2 Natural Predators of Maize Pests

2.1 Identification of key predators

Several natural predators play a crucial role in controlling maize pests (Figure 1). Among these, the predatory beetle Propylea japonica has been identified as a significant predator in maize ecosystems. This beetle is known to inhabit maize plants and travel to adjacent crops like cotton to prey on pests such as aphids (Ramsden et al., 2017; Yang et al., 2020). Other important predators include earwigs, wasps, ants, and spiders, which have been shown to derive their basal carbon from maize and other intercrops, indicating their role in pest regulation. The predator Teretrius nigrescens has been effective in reducing grain losses caused by the pest Prostephanus truncatus in stored maize (Bergvinson et al., 2011).

Figure 1 Comparative effects of arthropod pests on maize crops (Photo credit: Otieno et al., 2019) Image caption: a: Healthy, unaffected maize. b: A stunted maize stand due to attack by arthropod pests. c: A maize cob destroyed by arthropod pests (Adopted from Otieno et al., 2019)

2.2 Predator-prey dynamics

The dynamics between predators and their prey in maize fields are influenced by various factors, including habitat heterogeneity and cropping systems. For instance, intercropping maize with legumes or other crops enhances predator-herbivore feeding linkages, thereby naturally suppressing herbivory (Otieno et al., 2019). Predators like P. japonica exhibit oviposition preferences and habitat selection behaviors that optimize their predation efficiency. They tend to inhabit maize plants with high prey abundance and move to other crops to actively prey on pests (Fang et al., 2012; Ouyang et al., 2020). The use of flower strips as bridge habitats has also been shown to facilitate the movement of predatory beetles from wheat to maize, thereby enhancing their population and predation pressure in maize fields.

2.3 Ecological roles and benefits of predators

Natural predators provide significant ecological benefits by contributing to the biological control of maize pests. The presence of predators like P. japonica and T. nigrescens reduces the need for chemical pesticides, promoting environmentally sustainable pest management practices. The integration of diverse cropping systems, such as intercropping and the use of refuge plants, supports the conservation and effectiveness of these predators. For example, intercropping maize with legumes or planting flower strips can increase predator abundance and diversity, leading to more effective pest suppression (Quispe et al., 2017; Puliga et al., 2023). The use of resistant maize varieties in combination with natural predators has been shown to significantly reduce pest populations and associated crop losses.

3 Mechanisms of Pest Control by Natural Predators

3.1 Direct predation

Direct predation involves natural predators consuming pest insects, thereby reducing their populations. For instance, the predatory beetle Propylea japonica has been shown to actively prey on aphids in maize and cotton fields. Field investigations revealed that P. japonica adults prefer to inhabit maize plants and travel to cotton plants to prey on aphids, indicating a significant role in pest control through direct predation (Fang et al., 2012; Prischmann-Voldseth and Dashiell, 2013). Studies using artificial caterpillars in maize fields across Europe and Argentina demonstrated that invertebrates such as chewing insects, ants, and spiders, as well as vertebrates like birds and small mammals, contribute to pest control by directly attacking pest insects (Ferrante et al., 2019).

3.2 Parasitism and biological control

Parasitism is another crucial mechanism where parasitoids lay their eggs on or in pest insects, leading to the eventual death of the host. In Southern Africa, parasitoids have been identified as more efficient than predators in controlling the fall armyworm (FAW), a major maize pest. The proximity to forests enhances the effectiveness of these natural enemies, suggesting that landscape features can significantly influence parasitism rates (Clarkson et al., 2022). Intercropping systems, such as maize-legume combinations, have been shown to attract parasitoids and other beneficial insects, thereby enhancing biological control and reducing pest populations (Pierre et al., 2023).

3.3 Indirect effects and behavioral changes in pests

Natural predators can also induce behavioral changes in pests, leading to reduced pest activity and damage. For example, the presence of flower strips in wheat-maize rotation fields has been shown to conserve predatory beetles like P. japonica, which then migrate to adjacent maize fields. This conservation strategy not only increases predator abundance but also indirectly affects pest behavior by creating a hostile environment for pests (Lopes et al., 2016; Svobodová et al., 2017). The use of refuge plants next to maize crops has been found to enhance the populations of predators and parasitoids, which in turn exert pressure on pest populations, leading to reduced pest activity and damage (Quispe et al., 2017).

Natural predators control maize pests through direct predation, parasitism, and inducing behavioral changes in pests. These mechanisms are influenced by habitat management practices such as intercropping, the use of refuge plants, and maintaining landscape features like forest proximity, which enhance the effectiveness of natural enemies in integrated pest management programs.

4 Factors Influencing the Effectiveness of Natural Predators

4.1 Environmental factors

Environmental factors play a crucial role in determining the effectiveness of natural predators in controlling maize pests. Temperature fluctuations, for instance, can significantly alter the optimal predator community composition for biological control. Studies have shown that the foraging behavior of arthropod predators is influenced by an interaction between temperature and predator body size, which in turn affects pest control efficiency (Roubinet et al., 2017). Additionally, proximity to forests has been found to positively affect natural enemy-mediated control of pests like the fall armyworm, as diverse vegetation and near-forest proximity enhance the presence and effectiveness of predators such as birds, bats, parasitoids, and generalist predators (Clarkson et al., 2022).

4.2 Agricultural practices

Agricultural practices, including crop management and the use of cover crops, significantly influence the activity and effectiveness of natural predators. For example, mixed cropping systems, such as maize intercropped with sorghum or flower strips, have been shown to enhance the activity density and biocontrol potential of generalist predators compared to conventional monoculture systems (Puliga et al., 2023). Similarly, the use of flower strips as bridge habitats can facilitate the movement of predatory beetles from one crop to another, thereby enhancing pest control in double-cropping systems (Laubmeier et al., 2023). Early season plant cover has also been found to support more effective pest control than insecticide applications, as it increases the abundance of arthropod predators.

4.3 Interaction with other pest management strategies

The interaction between natural predators and other pest management strategies can either enhance or undermine biological control. For instance, the use of non-crop plants as refuges can provide resources that natural enemies need, thereby enhancing their populations and effectiveness in controlling maize pests (Quispe et al., 2017). However, the use of insecticides, particularly broad-spectrum ones, can disrupt the benefits associated with conservation practices by reducing the populations of beneficial arthropods (Rowen et al., 2022). Integrated pest management (IPM) strategies that respond to insect pest risk rather than using insecticides prophylactically have been found to be more disruptive to predator communities than preventative pest management (PPM) strategies. Therefore, promoting biological control through habitat management and minimizing the use of broad-spectrum insecticides can lead to more sustainable pest management outcomes. By understanding and optimizing these factors, farmers can enhance the effectiveness of natural predators in controlling maize pests, leading to more sustainable and environmentally friendly agricultural practices.

5 Case Study

5.1 Overview of East Africa

The selected case study region is East Africa, specifically focusing on the drier agro-ecologies of Kenya, Uganda, and Tanzania. This region has been significantly impacted by the fall armyworm, Spodoptera frugiperda, an invasive pest that has caused substantial damage to maize crops, a staple food in these countries (Midega et al., 2018).

5.2 Specific maize pests targeted

This case study focuses on the following major corn pests. Fall Armyworm (Spodoptera frugiperda) is an invasive pest that has recently invaded Africa, causing extensive damage to maize and other crops. Western Corn Rootworm (Diabrotica virgifera virgifera) is a pest that primarily damages maize roots and has shown resistance to Bt maize in some regions (Gassmann et al., 2011; López-Castillo et al., 2018). Maize Aphid (Rhopalosiphum maidis) is a common pest in maize crops that can be controlled by natural predators.

5.3 Natural predator species in the region

Several natural predator species have been identified in the region, contributing to the biological control of maize pests. Propylea japonica is a predatory beetle that prefers to inhabit maize plants and preys on aphids, providing potential for enhanced biological control in mixed crop systems (Fang et al., 2012; del-Val et al., 2023). Gaeolaelaps aculeifer is a soil-dwelling predatory mite effective against Western Corn Rootworm larvae. Various Predators and Parasitoids, including earwigs, wasps, predatory beetles, ants, and spiders, which have been shown to regulate herbivorous pests in intercropped maize fields.

5.4 Impact on maize pest population and crop yield

The implementation of natural predator-based pest control strategies has shown significant impacts on pest populations and crop yields. The climate-adapted push-pull system resulted in an 82.7% reduction in the number of larvae per plant and an 86.7% reduction in plant damage, leading to a 2.7 times increase in maize grain yields. The predator-in-first strategy using Gaeolaelaps aculeifer showed comparable results to pesticide use, effectively managing the pest population and protecting maize plants (Pasquier et al., 2021). Intercropping and organic farming practices strengthened predator-herbivore trophic linkages, naturally suppressing herbivory and reducing the need for chemical pesticides.

5.5 Lessons learned and implications for broader application

The case study in East Africa provides several key lessons and implications for broader application. The use of natural predators and habitat management strategies, such as intercropping and push-pull systems, can significantly reduce pest populations and increase crop yields, offering a sustainable alternative to chemical pesticides. Strategies like the climate-adapted push-pull system demonstrate the importance of tailoring pest management approaches to local environmental conditions and pest dynamics.The success of these strategies in East Africa suggests that similar approaches could be effective in other regions facing similar pest challenges, promoting sustainable agriculture and reducing reliance on chemical inputs (Midega et al., 2018; Otieno et al., 2019; Pasquier et al., 2021). By leveraging the natural predation mechanisms and enhancing habitat heterogeneity, farmers can achieve effective pest control, improve crop yields, and contribute to sustainable agricultural practices.

6 Challenges and Limitations

6.1 Variability in predator populations

The effectiveness of natural predators in controlling maize pests can be significantly influenced by the variability in predator populations. Factors such as land use and crop management practices can alter the abundance of predatory insects. For instance, the presence of intensively managed crops like orchards can negatively impact predator populations in adjacent fields due to pesticide use, which acts as a sink for predators (Ardanuy et al., 2018; Madeira et al., 2021). The abundance of predators is often temporally and species-specific, influenced by local and landscape structures. This variability poses a challenge in maintaining consistent biological control across different environments and seasons.

6.2 Challenges in field implementation

Implementing biological control strategies in the field faces several challenges. One major issue is the periodic disturbance regimes such as crop planting and harvest, which can lead to frequent mortality events among predator populations (Yang et al., 2020). Conservation strategies, such as planting flower strips, have been shown to help maintain predator populations during these critical periods, but their effectiveness can vary. Moreover, different cropping systems, such as monoculture versus intercropping, can significantly influence the activity and effectiveness of generalist predators. For example, mixed cropping systems tend to support higher predator activity and biocontrol potential compared to continuous monoculture systems (Puliga et al., 2023). However, these effects are not always consistent, making it difficult to standardize practices across different fields and regions.

6.3 Risks of disrupting ecosystem balance

While promoting natural predators can enhance pest control, there is a risk of disrupting the ecosystem balance. Introducing or encouraging certain predator species can lead to unintended consequences, such as the suppression of non-target species or the alteration of existing predator-prey dynamics. For instance, the presence of predatory ants in species-rich assemblages can significantly increase predation rates, but this effect is influenced by local and landscape factors, which can vary widely (Liu et al., 2018; Frizzo et al., 2020). The use of non-crop plants to attract natural enemies must be carefully managed to avoid attracting pests or disrupting other beneficial species (Quispe et al., 2017). Ensuring that these strategies do not negatively impact the broader ecosystem requires careful planning and monitoring.

7 Future Directions and Research Needs

7.1 Enhancing predator efficiency

To enhance the efficiency of natural predators in controlling maize pests, it is crucial to understand their habitat preferences and feeding behaviors. For instance, the predatory beetle Propylea japonica shows a preference for maize plants before moving to cotton to prey on aphids, suggesting that maize can serve as a habitat to support predator populations. The use of flower strips has been shown to facilitate the movement of predatory beetles from wheat to maize, significantly increasing predator abundance in maize fields (Yang et al., 2020). Future research should focus on identifying and optimizing such habitats and resources that can support and enhance predator populations.

7.2 Integrating natural predators with modern pest management techniques

Integrating natural predators with modern pest management techniques can provide a sustainable approach to pest control. For example, the predator-in-first strategy, which involves introducing predatory mites along with an alternative food source, has shown promising results in controlling Western Corn Rootworm populations in maize fields (Pasquier et al., 2021). Moreover, intercropping and organic farming practices have been found to strengthen predator-herbivore trophic linkages, thereby enhancing natural pest suppression. Research should explore the synergistic effects of combining natural predators with other pest management strategies, such as the use of selective herbicides that have minimal impact on natural enemies (Bohan et al., 2011; Zilnik et al., 2023).

7.3 Long-term monitoring and impact assessment

Long-term monitoring and impact assessment are essential to evaluate the effectiveness and sustainability of using natural predators for pest control. Studies have demonstrated the importance of monitoring predator activity and predation pressure across different regions and cropping systems (Ferrante et al., 2019). The proximity of forests has been shown to positively affect natural enemy-mediated control of pests like the fall armyworm, highlighting the need for landscape-level assessments (Clarkson et al., 2022). Future research should focus on developing standardized monitoring methods and conducting long-term studies to assess the impact of natural predators on pest populations and crop yields across various agroecosystems.

8 Concluding Remarks

The research on the role of natural predators in controlling maize pests has highlighted several key findings. The intensification of conventional agriculture negatively impacts biodiversity and the delivery of ecosystem services, including pest control by generalist predators. Studies have shown that mixed cropping systems, such as maize intercropped with runner beans, sorghum, or flower strips, enhance the activity and biocontrol potential of generalist predators compared to monoculture systems. Intercropping and organic farming practices have been found to strengthen predator-herbivore trophic linkages, thereby naturally suppressing herbivory in non-Bt maize fields. The use of flower strips as bridge habitats has also been effective in conserving and enhancing predator populations, such as Propylaea japonica, during crop phenophase changes. Conservation agriculture practices, including zero tillage and residue retention, have been shown to positively influence arthropod community dynamics and enhance biological control potential. The importance of biocontrol for maize production is widely recognized, with experts agreeing on its significance for sustainable agriculture.

The findings from these studies have significant implications for sustainable maize cultivation. The adoption of mixed cropping systems and intercropping can enhance biodiversity and the effectiveness of natural predators in controlling maize pests, reducing the reliance on chemical pesticides and their associated environmental and health impacts. Conservation agriculture practices, such as zero tillage and residue retention, can improve soil quality and provide favorable habitats for beneficial arthropods, further supporting pest control. The use of flower strips as bridge habitats can help maintain predator populations during critical periods, ensuring continuous pest suppression. These strategies not only contribute to sustainable pest management but also promote overall ecosystem health and resilience.

Natural predators play a crucial role in the biological control of maize pests, offering a sustainable alternative to chemical pesticides. The enhancement of predator populations through habitat management, intercropping, and conservation agriculture practices can significantly improve pest control and contribute to sustainable maize production. Future research should continue to explore and refine these strategies, focusing on optimizing predator-prey interactions and understanding the broader ecological impacts. By integrating these practices into maize cultivation, farmers can achieve more sustainable and resilient agricultural systems, ultimately contributing to global food security and environmental sustainability.

Acknowledgments

The authors appreciate the reviewing experts for their insightful suggestions on the manuscript, which benefited the overall quality of the manuscript.

Conflict of Interest Disclosure

The authors affirm that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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