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The Role of Entomopathogenic Fungi in Tick Control

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Emily Mesquita, Jéssica Fiorotti, Laura Nobrega Meirelles, Victória Silvestre Bório, Thaís Almeida Corrêa, Adriani da Silva Carneiro Lopes and Patricia Silva Golo

Submitted: 07 January 2026 Reviewed: 02 February 2026 Published: 07 April 2026

DOI: 10.5772/intechopen.1014844

Ticks and Tick-Borne Diseases IntechOpen
Ticks and Tick-Borne Diseases Edited by Muhammad Abubakar

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Ticks and Tick-Borne Diseases [Working Title]

Dr. Muhammad Abubakar, Dr. Shumaila Manzoor and Dr. Abdul Kabir

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Abstract

Tick control is one of the major concerns in animal production, livestock, and public health. Integrated tick management (ITM) is a strategic plan that combines different control methods. Among the non-chemical approaches, biological control using entomopathogenic fungi (EPF) has widespread utilization in agriculture, with a special focus in Brazil. Today, Brazil leads the world’s largest biological control program employing fungus-based bioproducts. There is a great amount of data available on ticks using these microorganisms – in vitro and in vivo results – including semi-field and field assays assessing fungal virulence, tick-immune response to fungal infection, fungal formulation, and their persistence/maintenance in the environment. The dynamics of EPF infection depend on the fungal isolate traits, the interaction of the fungus with the arthropod host, the microclimate where the fungus is applied, and the association with soil and plants. There is, however, a limitation regarding the comparison of agrochemicals versus bioproduct efficacy and what we should expect, particularly due to fungal tolerance to abiotic factors, which thus impacts the acceptance of widespread use. Therefore, the use of biological control with EPF against ticks will rely on each country’s understanding. This chapter aims to update current trends in entomopathogenic fungi for tick control, providing technical and scientific insights that support integrated tick management (ITM) and future regulatory decision-making.

Keywords

  • sustainable tick control
  • integrated tick management
  • metarhizium
  • Beauveria
  • endophytic fungi
  • bioinputs

1. Introduction

According to the Food and Agriculture Organization (FAO) guidelines, there is an increase in tick acaricide resistance in many parts of the world, for both one-host and multi-host ticks [1]. It is stated that the major factors driving the selection of resistance are the high frequency of use and the incorrect use (dosage, application), mainly with products that have the same mode of action. Furthermore, there are associated risks for people in small properties, milk/meat contamination, and environmental pollution [1]. Due to this fact, efforts to establish Integrated Tick Management (ITM) strategies aimed at achieving more sustainable tick control are not new. Biological control is defined as the use of organisms to suppress pest populations [2], and it encompasses several microbial-based approaches, including entomopathogenic bacteria, nematodes, and fungi [3]. Among these, entomopathogenic fungi (EPF) have been selected as one of the main tools within ITM, as their mode of action relies on direct cuticle penetration, without the need for ingestion by arthropods. The development of EPF-based bioproducts has grown steadily in recent years, with particular emphasis on agricultural pest control in Brazil [4, 5]. Worldwide, strategies using fungi against ticks are reported in the United States of America [6], mainly with the commercial strain Met52 targeting the Ixodes ricinus complex, and in Mexico for Rhipicephalus microplus control [7], highlighting different EPF species.

Fungal action follows adhesion on the tick cuticle, germination, penetration into the tick body, colonization of tissues, and finally sporulation on the outside [8]. However, as living organisms, there are limitations in their use regarding fungal viability. For an EPF isolate to be chosen, it needs to demonstrate enough tolerance to heat, UV radiation, and desiccation, besides, of course, high virulence for the target arthropod [4]. Mostly, when EPF do not meet all the requirements, these issues might be addressed by an efficient and proper formulation for each entomopathogenic fungal propagule (i.e., aerial or submerged conidia, blastospores, microsclerotia) as well as with the use of genetic editing techniques.

Moreover, intrinsic traits of EPF isolates play a crucial role in their effectiveness. In ticks, for example, a higher number of fungal propagules is often required compared with insects, a difference that can be explained by both host-related and environmental factors. Host-associated features include cuticle composition [9], tick-associated microbiota [10, 11], and the tick immune response [12]. From an environmental perspective, EPF must overcome established microbial communities on animal skin or in the soil, as well as plant-associated factors such as vegetation type and the bioactive compounds they release. Despite the knowledge in tick control using EPF described further in this chapter, the main challenges are suggested to be (1) identifying the most suitable formulation for each fungal strain and their respective propagules; (2) linking each region's needs with the right bioproduct; and (3) aligning the demands of bioproduct production and efficacy with the required local legislation. Accordingly, this chapter describes the main approaches, recent findings, and remaining knowledge gaps related to the use of EPF within Integrated Tick Management (ITM) strategies. The word “ticks” mentioned throughout the text is related mainly to the Ixodidae family.

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2. Biological assays using fungi

2.1 From laboratory bioassays to semi-field and field evaluations

Laboratory bioassays represent a fundamental step in the initial assessment of the pathogenicity and virulence of EPF for tick control. These evaluations allow the control of experimental variables, screening of new isolates based on their tolerance to simulated abiotic factors, and direct comparisons among isolates, including dose-response analyses under standardized exposure conditions [13, 14]. The results obtained at this stage guide subsequent steps in the validation of EPF isolates. In vivo studies contribute substantially to the understanding of EPF performance against different tick species and under distinct thermal conditions. Borisov et al. [15] demonstrated the pathogenicity of Metarhizium and Beauveria species against Ixodes ticks at lower temperatures (≈14 °C), showing that the evaluated EPF isolates can cause significant mortality even under suboptimal thermal conditions. These results highlight the importance of selecting isolates adapted to the seasonal and climatic variability of target regions. The biology and life cycle of ticks further emphasize the need to move beyond evaluations conducted exclusively under laboratory conditions. Ticks spend part of their life cycle off-host [16], and during this phase, females oviposit in the environment; then larvae and nymphs may remain for weeks in the vegetation or soil while awaiting host passage [17]. The success of biological control, therefore, depends on the prior environmental competence of EPF, from survival in the environment to the establishment of infection in ticks. Beyond isolate selection, the type of propagule (aerial conidia, submerged conidia, blastospores, or microsclerotia) and formulation strategy (e.g., liquid, granular, or encapsulated) become central components of efficacy under semi-natural and field conditions [4, 18–20]. Accordingly, isolate screening should be followed by evaluations of environmental adaptation and propagule stability.

The transition from controlled laboratory bioassays to semi-field and field trials is crucial in the development and validation of bioproducts, where methodological robustness is decisive. This transition depends not only on the biological efficacy of the agent but also on rigorous quality control, the selection of isolates adapted to local conditions, and interdisciplinary approaches that consider biological, ecological, and large-scale production aspects [21]. Semi-field and field trials comprise validation stages that differ in terms of experimental control, ecological complexity, and the ability to generalize results [22].

Semi-field trials are generally conducted in partially controlled environments, such as screened areas, small vegetation plots, and soil-filled containers. These systems allow for the introduction of ticks at defined densities and developmental stages while exposing them to natural substrates and realistic microclimatic conditions [19, 23]. This approach is suitable for evaluating the persistence of fungal propagules in soil, their capacity for in situ germination, and the secondary release of infective conidia under natural environmental conditions [18, 24]. Environmental variables, such as soil moisture, temperature, and vegetation cover, may vary naturally or be experimentally manipulated, allowing for the assessment of persistence, infectivity, and contact between ticks and fungi [25]. The main advantage of semi-field trials is their role as an intermediate scenario between laboratory assays and open-field conditions. These systems make it possible to observe the influence of biotic factors, soil type, and application methods of bioproducts. This approach reduces the risk of failure during the scaling-up of promising formulations, particularly those based on granular or encapsulated formulations, before field application. Studies conducted under these conditions have demonstrated that granular formulations of Metarhizium robertsii based on microsclerotia [19] exhibit greater persistence and performance compared to non-formulated isolates. However, experiments conducted in restricted areas may generate so-called “edge effects” and may not fully capture the heterogeneity of tick mobility or the influence of local fauna [26], requiring caution when directly extrapolating results to natural environments. These trials are conducted in productive environments, such as pastures and rural properties, where natural tick populations are established and continuously interact with their hosts throughout complete life cycles. Field trials incorporate seasonal variation, the presence of vertebrate hosts, soil and vegetation heterogeneity, and unpredictable climatic factors [27]. The results obtained represent the most ecologically realistic approach for evaluating the efficacy of fungal formulations for tick control.

Despite their high relevance, field trials pose significant logistical and methodological challenges, including higher costs, lower experimental control, and greater variability in outcomes. Consequently, structured approaches to methodological validation have been proposed to increase the reliability and reproducibility of data obtained from semi-field and field trials. These approaches include the clear definition of performance criteria, adequate replication, and statistical analyses compatible with complex biological systems. In this context, Matope et al. [26] propose a four-step validation process – preliminary development, feasibility studies, internal validation, and external validation – which provides an organized framework to evaluate control tools in variable biological scenarios, contributing to greater reproducibility and comparability among studies.

Semi-field and field trials allow the identification of isolates and formulations that maintain performance under dual pressure (abiotic effects and host-related challenges), as well as the identification of favorable environmental windows for application – such as relative humidity > 95%, temperatures between 20 and 30 °C, and low UV radiation – which are essential for selecting locally adapted isolates [28]. Thus, laboratory, semi-field, and field approaches should not be interpreted as competing methodologies, but rather as complementary stages within a validation pipeline. Each experimental level presents intrinsic advantages and limitations, mainly related to the degree of environmental control, ecological complexity, and the ability to extrapolate results. Understanding these differences is fundamental for the correct interpretation of experimental data.

2.2 Field-based studies using entomopathogenic fungi for tick control

In contrast to laboratory and semi-field assays, field-based studies allow the evaluation of the impact of EPF not only on isolated individuals but also on tick population dynamics over time. Within this approach, interactions with vertebrate hosts, weather, and management practices adopted during the study are simultaneously considered, providing a more realistic representation of environmental and productive scenarios. Camargo et al. [27] conducted one of the most relevant studies in this field by evaluating the direct application of Metarhizium on cattle, aiming to control the parasitic phase of R. microplus. Although the conidia-based formulation showed partial efficacy, the authors highlighted limitations related to fungal persistence under adverse environmental conditions, emphasizing the challenges inherent to direct applications in open systems. For the control of Ixodes scapularis nymphs in residential landscapes and forest edges, formulations of Beauveria bassiana and M. anisopliae F52 were tested [29]. Under these conditions, treatments with B. bassiana reduced nymphal abundance in urban lawns by ~ 90% in the absence of vegetation barriers and by up to ~ 90% when combined with wood-chip barrier strips. Applications of M. anisopliae F52 in similar areas also resulted in significant reductions in nymph densities in 2002, demonstrating that EPF can effectively impact populations of ticks in natural and peri-urban environments.

The timing of application throughout the year also plays a critical role in EPF efficacy, being more favorable during periods of higher relative humidity and lower ultraviolet (UV) radiation, conditions that are expected to promote conidial germination and persistence in soil and vegetation. This seasonal shift was demonstrated with the report of mortalities of up to 92% in Amblyomma variegatum during the wet season, whereas mortality rates during the dry season were significantly lower (24%), clearly illustrating the direct impact of environmental conditions on EPF efficacy in the field [30].

The application of EPF in pastures, however, requires careful consideration of environmental safety, particularly with respect to non-target species. Evidence indicates that B. bassiana and M. anisopliae exhibit host-dependent specificity, with infection processes conditioned by cuticular cues and the physiological context of the host [31]. Evidence shows that beneficial insects, such as the lady beetle Coccinella septempunctata, display avoidance behavior when exposed to B. bassiana conidia, thereby reducing the risk of contact with EPF and this species [32]. Laboratory evaluations with bees indicate variable effects depending on the species assessed and the route of exposure [33]. Thus, when applied in a targeted manner, using appropriate formulations and during environmentally favorable periods, EPF are considered biologically safe tools for use in pasture systems. The heterogeneity frequently observed in field results should therefore be interpreted as a reflection of the complexity of productive systems rather than as a methodological failure. Despite these challenges, field trials provide essential information that cannot be obtained through laboratory or semi-field studies alone. They allow the identification of operational limitations of fungal formulations, the assessment of tick population suppression under real-world conditions, and the generation of critical data for regulatory processes and for the development of integrated tick management programs.

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3. Formulations based on entomopathogenic fungi for tick control

The persistence of EPF in the environment is closely linked to their ability to tolerate abiotic factors. High or low temperatures, relative air humidity, and ultraviolet radiation can compromise the viability of these organisms under field application conditions [4, 34]. Thus, the development of formulations based on EPF is a crucial strategy that enables longer persistence of these biocontrol agents when exposed to environmental stressors, as well as improving field application for these bioproducts. Formulating EPF involves adding other components to improve the performance of these microorganisms, contributing to greater viability, stability, and control efficacy [35]. Besides enhancing the fungus life cycle in the environment, formulations also increase the viability of EPF propagules during critical stages of industrial production, such as maintenance, storage, and transport. Another advantage is the ease of application in different control strategies (i.e., on animals or soil), as well as the possibility of including additives that stimulate germination and adhesion to the host. These factors together might reduce the frequency of reapplication and reinforce the economic competitiveness and viability of these entomopathogens as biocontrol agents [25, 36].

Different types of fungal propagules, as well as mycelium, can be used as active ingredients in formulations. The development of products containing entomopathogenic fungi (EPF) requires the evaluation of several aspects, including compatibility between components and the selected propagules, shelf life under different storage conditions, fungal efficacy in the field, protection against environmental factors, and adequate dispersion according to the intended use [37]. The ease of application and acceptance by farmers are also important considerations. There are different types of formulations based on EPF, according to solid or liquid forms, mainly defined by their physical state, mode of application, and behavior in water or oil [35, 38]. Solid formulations of mycopesticides are employed as wettable powders (WP), dustable or sprayable powders (DP), granules (GR), ready-to-use baits (RB), water-dispersible granules (WG), and contact powders (CP). In contrast, liquid formulations comprise suspension concentrates (SC), oil-miscible fluid concentrates (OF), ultra-low-volume suspensions (ULV), and oil dispersions (OD) [4, 35, 39].

Wettable powder (WP) formulations are one of the most traditional methods of applying EPF for biological tick control [40]. This formulation consists of dried conidia mixed with inert substances and agents that facilitate dispersion in water, enabling the product to be prepared at the time of application. The main advantages of WP are ease of storage, transport, and preparation, as well as compatibility with commonly used spraying equipment in animal health management [35, 38]. After application, as the fungal propagules come into direct contact with the tick’s surface, it favors fungal fixation and the beginning of the infection process [41]. Due to these characteristics, WP is frequently used as a reference formulation in experimental studies and as a basis for developing other technologies, such as oil-based, encapsulated, or granular formulations. These alternatives aim to increase the stability of the fungi in the environment and improve efficiency in tick control [38]. The formulation of Metarril WP® (a commercial product based on M. anisopliae spores) was reported in 10% or 0.5% mineral oil and showed to be effective against eggs, larvae, and engorged females of Dermacentor nitens in laboratory assays [42]. In a field trial, Camargo et al. [27] evaluated the effectiveness of Metarril SP Organic Plus® (commercial WP product derived from Metarhizium sp.) in controlling R. microplus when added to 10% mineral oil. This oil formulation significantly reduced the average number of ticks collected from cattle housed in paddocks. Granular solid formulations are a promising strategy for tick control using EPF [19]. These GR incorporate the fungal propagules, providing protection against environmental factors while permitting controlled release [35]. Granules are suggested to facilitate handling, reduce the required amount of biomass, and decrease the frequency of applications [37, 43]. Additionally, GR enables direct application to the environment. These facts highlight a possible effective method for controlling ticks found in soil during their non-parasitic phase, prospecting for future fungal-based products.

A granular formulation of M. robertsii microsclerotia or blastospores reduced the number of tick larvae when tested against R. microplus in semi-field conditions; however, persistence on soil was not satisfactory [19], probably due to abiotic factors affecting the propagules outside the capsules’ protection. Besides this, pellets containing inorganic materials and microsclerotia of M. anisopliae preserved fungal viability following fluidized bed drying and short storage at 4 °C [44]. The granules were also effective against R. microplus females and showed outstanding UV–B tolerance in laboratory tests. Technologies using encapsulation are important methods of cell immobilization for biocontrol agent formulations [45, 46]. The ionic gelation encapsulation technique has already been used to produce granules containing EPF for tick control against species such as Ixodes ricinus [47] and R. microplus [20]. When encapsulating Metarhizium pemphigii blastospores in sodium alginate, adding corn starch or chitin, aerial conidia of both granular formulations exhibited high virulence, resulting in 100% mortality of I. ricinus nymphs [47]. Moreover, encapsulation by ionic gelation using 2% and 3% sodium alginate contributed to increasing the shelf life at room temperature as well as the UV–B tolerance and thermotolerance of M. anisopliae LCM S01 conidia [20]. Furthermore, the granulate formulation applied to the soil was effective in controlling R. microplus engorged females in laboratory bioassays.

In the context of liquid formulations, oil-based formulations are the most frequently reported. When applying M. anisopliae, particularly in the form of oil-in-water emulsions, these formulations increased efficacy in the control of R. microplus, especially under outdoor conditions [23, 48]. This type of formulation improves wettability and spreading over the target pest, enhances the bioavailability of the active ingredient, facilitates handling, and provides greater protection against thermal stress and imbibitional damage, in addition to increasing fungal persistence on treated surfaces [38]. An evaluation in 2005 already exhibited trends for cattle tick control using biological oil-in-water emulsions. Researchers tested paraffin oil, palm oil, and emulsifiable adjuvant oils (EAOs) for their ability to improve the performance of M. anisopliae against R. microplus. The study showed germination above 68% after 24 hours and close to 100% after 48 hours. The 10% liquid paraffin formulation of M. anisopliae (EAO) was the most effective, combining moderately high germination after 24 hours with a shorter tick survival time [49]. For Rhipicephalus evertsi evertsi ticks, conidia were formulated in olive oil and water, with the addition of two different sunscreen agents [50]. When conidial survival was assessed after 5 hours of exposure to UV radiation, the oil-based formulation supplemented with the E45 sunscreen showed 40% fungal germination, almost twice conidia when compared to aqueous control. The addition of olive oil and sunscreen agents to formulations provided significant mortality in larvae and unfed adults, besides conidial protection against UV radiation.

Until today, there are only a few data addressing the use of specific formulations – such as ultra-low volume (ULV) suspensions, suspension concentrates (SC), and oil-miscible fluid concentrates (OF) – for the biological control of ticks using EPF. The lack of consistent reports on the use of ULV for mycopesticides may be associated with the fact that this technology is still under development and requires financial investment, while facing limitations related to the stability of fungal propagules and viability after the atomization process [38]. Therefore, most studies focus on oily formulations or emulsions, generally more conceptually similar to OF formulations. Overall, the effectiveness of these formulations is directly related to their ability to protect conidia against adverse environmental factors, as well as the application method employed under experimental or field conditions.

The biopesticide market is experiencing accelerated growth, driven by advances in formulation technologies, increased efficacy of EPFs, and favorable changes in regulatory frameworks, surpassing the expansion rate of conventional chemical insecticides [38]. A compound annual growth rate (CAGR) of between 11% and 16% is estimated for the period from 2025 to 2035, with a market value already exceeding US$ 8 billion in 2024 and projections indicating the possibility of surpassing US$ 20 billion, potentially approaching US$ 30 billion in the early 2030s. Accordingly, EPF occupy a relevant position in the field, although their market share is frequently underestimated in scientific publications. Recent data indicate that bioinsecticides account for approximately 37–40% of the global biopesticide market, with Beauveria bassiana, M. anisopliae, and Akanthomyces lecanii being the main representatives. Based on these estimates, fungal mycopesticides should represent about 16.7% to 18% of the total biopesticide market [38]. The worsening of climate crises represents a challenge for the survival and effectiveness of microorganisms employed in biological control. In this context, improving fungal formulations is crucial for the effectiveness of these biocontrol agents.

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4. Tick-immune response to fungi infection

Ticks are invertebrates and, therefore, lack the adaptive immune system found in vertebrates. Their defense against invading microorganisms, including EPF, relies entirely on innate immunity, which integrates rapid cellular responses mediated by hemocytes with humoral mechanisms involving soluble effector molecules and enzymatic cascades [12, 51, 52]. The main hemocyte types described in ixodid ticks include plasmatocytes, granulocytes, prohemocytes, oenocytoids, and spherulocytes, although their relative abundance and functional specialization may vary among species and developmental stages [12]. Plasmatocytes and granulocytes are generally the most abundant cell types in the hemolymph and are considered the primary effectors during immune challenges. Plasmatocytes are mainly involved in phagocytosis, encapsulation, and interaction with invading pathogens, whereas granulocytes exhibit pronounced secretory activity and are key mediators of pathogen recognition, degranulation, and cytotoxic responses [12].

Fungal infection induces marked quantitative and qualitative changes in hemocyte populations in ticks. Experimental infections with EPF frequently result in an increase in circulating plasmatocytes and granulocytes, reflecting their recruitment and activation during pathogen recognition and elimination. Conversely, reductions in prohemocytes have been reported, suggesting their differentiation into effector cell types during immune activation [12, 52]. Oenocytoids, although less abundant, play a crucial role in the prophenoloxidase (proPO) system and melanization responses, processes often triggered during fungal invasion of the hemocoel. It is suggested that fungal pathogens can disrupt oenocytoid function or reduce their numbers, thereby impairing melanization and facilitating immune evasion [12, 51]. Collectively, these alterations in hemocyte composition highlight the dynamic nature of the tick cellular immune response and underscore the importance of hemocyte-mediated mechanisms in determining susceptibility or resistance to entomopathogenic fungi.

Although some invertebrates display immune priming, where previous pathogen exposure enhances subsequent immune responses [53], there is currently no convincing evidence that ticks develop immune priming against EPF. Similarly, studies in other arthropods, such as Rhodnius pallescens exposed to Beauveria bassiana, have not demonstrated survival benefits attributable to immune priming [54]. Complement-like proteins, including α-macroglobulins and thioester-containing proteins (TEPs), mediate recognition and opsonization of microbes during phagocytosis [55–58]. Ticks can phagocytose EPF, although fungal virulence may damage hemocytes, illustrating an evolutionary arms race between host defense and pathogen activity [12, 59, 60]. Despite these advances, the molecular signaling pathways controlling phagocytosis in ticks remain poorly characterized. Beyond phagocytosis, autophagy contributes to microbial clearance and represents an important innate defense mechanism in invertebrates [61–63]. Nodulation involves multilayer aggregation of hemocytes around invading microorganisms [64]. Moreover, dopamine signaling modulates hemocyte activity and survival following fungal challenge, highlighting a neuro-immune axis in ticks [60, 65].

Melanization is triggered by the activation of the prophenoloxidase cascade, producing melanin and cytotoxic intermediates that damage pathogens [66, 67]. Melanization of fungal structures has been observed during EPF infection in both insects and ticks [68, 69]. Apoptosis also plays a relevant role in the tick immune response during EPF infection. Infection with fungi such as M. anisopliae and B. bassiana has been associated with increased apoptotic activity in tick hemocytes, characterized by nuclear condensation, DNA fragmentation, and activation of caspase-dependent pathways, which may contribute to pathogen control but can also compromise immune efficiency when excessively activated [70]. In parallel, fungal virulence factors and secondary metabolites are known to manipulate apoptotic signaling pathways, promoting hemocyte death and facilitating fungal persistence and dissemination within the host [71–73].

When infection becomes prolonged or systemic, humoral immunity supplements cellular defense. In invertebrates, four major pathways regulate immune gene expression: Toll, IMD, JAK-STAT, and JNK [70, 74]. Most components of the Toll pathway are conserved in ticks [75–78]. Although ticks lack several canonical proteins of the IMD pathway, this pathway can still functionally operate [77, 79, 80]. The JAK-STAT pathway, originally described in mammals [81], is also conserved in ticks [75, 76]. Moreover, ticks produce a restricted but important repertoire of antimicrobial peptides (AMPs), including defensins, hemocidins, microplusins, lysozymes, and ixodidins [82–86]. These peptides exhibit broad antimicrobial activity [87, 88], although their specific contribution to antifungal defense against EPF remains insufficiently explored.

Oxidative stress occurs when oxidants exceed antioxidant buffering capacity [89]. In ticks, heme and iron metabolism play key roles in physiology and immunity [90, 91]. Activation of the prophenoloxidase cascade during fungal infection promotes melanization and generates reactive oxygen species (ROS), which contribute to pathogen killing but may also damage host tissues [66, 92]. Studies in insects show that ROS dynamics during fungal infection vary depending on fungal species and virulence and are influenced by antioxidant enzymes such as superoxide dismutase and catalase [93, 94]. However, ROS biology in tick hemocytes remains poorly investigated [95].

Despite lacking adaptive immunity and some canonical immune pathways found in insects, ticks possess robust, multilayered defenses against EPF. These include hemocyte-mediated cellular responses, immune signaling cascades, complement-like proteins, antimicrobial peptides, melanization processes, and redox metabolism [12, 59, 60]. A deeper understanding of these mechanisms will improve our knowledge of tick–pathogen interactions and support the rational development of fungal-based biological control strategies capable of overcoming tick immune barriers.

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5. Tick-fungi ecology

As ticks spend most of their life cycle off the host and within the environment, effective tick control using EPF requires a comprehensive understanding of tick–fungi–environment interactions. This includes the influence of soil, vegetation, microclimate, and associated microbial communities on fungal persistence and performance. Accordingly, as the use of EPF intensifies, it is crucial to narrow down the fungal association in plant-arthropod-soil interactions, as well as potential competitor microorganisms that may affect fungal action.

Some well-known EPF, such as Metarhizium spp. and Beauveria spp., have been described as endophytic fungi, meaning that they are capable of associating with plant tissues during part of their life cycle without causing harm, while potentially engaging in nutrient exchange [96–100]. A great number of Metarhizium species have been associated with crops only in Brazil, exhibiting outstanding biodiversity and versatility [5], and worldwide this milestone is even higher [101]. More recently, interaction among both plant growth-promoting EPF in grassland plants demonstrates the potential to strengthen pastoral systems, increasing the biomass of roots and shoots with positive endophytic colonization [102]. Here, it is worth mentioning the possible use of switchgrass in the biofuel industry [103].

The phenomenon of nutrient cycling (encompassing nutrient availability and acquisition) induced by endophytic EPF highlights their ecological role in soil–plant systems. In environments where dead arthropods are present in the soil, these fungi can exploit insect-derived resources while simultaneously functioning as nutrient drivers for associated plants. This process reflects a tripartite interaction involving fungi, arthropods, and plants, as EPF are commonly found in soil ecosystems. Beyond their direct effects on nutrient availability – often increasing nutrient pools – certain EPF isolates have been shown to enhance plant nutrient uptake [104]. This occurs because EPF-mediated decomposition of organic matter releases nutrients and minerals in readily available forms, driven by the secretion of hydrolytic and oxidative enzymes [104].

The multifunctional lifestyle (biocontrol agent, bioinoculant, and saprophyte) of EPF is particularly interesting for fungal persistence in the environment, since this close association in the soil-plant environment might work as a reservoir for EPF. In the non-parasitic phase of ticks, engorged females, eggs, and unfed larvae (for monoxenous ticks) or eggs, larvae, nymphs, and adults unfed and engorged (for heteroxenous ticks) are found on the soil, which aligns perfectly with saprophyte EPF biology. Therefore, soil treatment is likely to hinder the tick life cycle even when not directly applied to ticks [23]. However, both ticks and fungi are exposed to environmental conditions. In fact, the susceptibility of EPF propagules (i.e., aerial or submerged conidia, blastospores, or microsclerotia) to abiotic factors is one of the main gaps between a desirable isolate and an effective bioproduct. Since the first 72 hours are crucial for fungal attachment, germination, and penetration [105], it is essential that fungi are protected to remain viable. Accordingly, formulation development is continuously improved to increase fungal tolerance to abiotic stresses, as stated in the Fungi Formulation Section 3.

Soil structure itself can be influenced by the growth and colonization of EPF, as the development of fungal hyphae (fungal biomass) and the secretion of extracellular compounds contribute to improved soil aggregation, water infiltration, aeration, and root penetration [104]. However, soil moisture levels also play a critical role in shaping fungal dynamics. Considering the multiple influences of EPF within soil–plant ecosystems, EPF tend to be less disruptive to soil microbial communities and enzymatic activities than chemical pesticides [106]. These authors reported that nitrogen cycling activity increased following Metarhizium inoculation, while the abundance of pathogenic microorganisms remained unchanged. Although this ubiquitous endophytic trait of EPF may raise concerns among some stakeholders, given their presence within crops commonly consumed in the human diet, an increasing body of evidence supports the safety of EPF for animals and non-target arthropods [34, 107].

In the future perspective of EPF use for the management of tick-borne diseases, it remains unclear whether EPF exhibit similar levels of virulence against ticks infected with tick-borne pathogens, particularly under outdoor conditions. Tick-borne pathogens and their arthropod hosts typically establish highly adapted and efficient associations shaped by long-term coevolution, which may influence host physiology, immune responses, and susceptibility to external stressors. Increasing evidence suggests that microbial symbionts and pathogens can modulate tick fitness, immune pathways, and microbiome composition, potentially affecting interactions with entomopathogenic fungi [11, 108]. Moreover, environmental variables such as temperature, humidity, soil characteristics, and microbial competition further complicate EPF performance under field conditions [109, 110]. Together, these factors highlight the need for integrative studies addressing tick–pathogen–fungus interactions within realistic ecological contexts in order to better predict EPF efficacy and to support their incorporation into integrated tick management strategies targeting tick-borne diseases.

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6. Conclusion

Here, a wide body of results on tick-EPF dynamics is presented, highlighting the ability to improve tick control due to their plasticity, in contrast with the numerous variables that should be noted when applying this strategy in ITM. There are still unpredictable limitations to be addressed as it is applied in outdoor conditions. Therefore, most of the strengths and deficiencies in using EPF for tick control should be considered when producing a potential bioacaricide and deciding when, where, or why to use this approach.

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Acknowledgments

This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior ‐ Brasil (CAPES) ‐ Finance Code 001, providing PhD’s scholarship for Thaís Almeida Corrêa and Laura Nobrega Meirelles. The Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) of Brazil provided scholarships for Victório Silvestre Bório and funding for Emily Mesquita. Rio de Janeiro State Research Support Foundation Carlos Chagas Filho (FAPERJ) provided scholarships for Thaís Almeida Corrêa. Patrícia Silva Gôlo is part of the comitte of Institutos Nacionais de Ciência e Tecnologia- INCT of Innovative Bioinputs.

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Written By

Emily Mesquita, Jéssica Fiorotti, Laura Nobrega Meirelles, Victória Silvestre Bório, Thaís Almeida Corrêa, Adriani da Silva Carneiro Lopes and Patricia Silva Golo

Submitted: 07 January 2026 Reviewed: 02 February 2026 Published: 07 April 2026

© 2026 The Author(s). Licensee IntechOpen. This content is distributed under the terms of the Creative Commons Attribution 4.0 International License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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