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Beyond Insects: Emerging Arthropod Reservoirs of Current and

Novel Entomopathogens for Insect Pest Management – A Review

Babatunde A. Kelly,

Taiye A Jemilaiye,

Funmi Olatujoye,

Olayinka E. Omoseyin,

Marcus O. Bello,

David O.

Adegbola,

David O. Ajibola,

Esther L. Jacob,

Oluwabunmi G. Adeusi,

Department of Microbiology, Adekunle Ajasin University Akungba-Akoko, Nigeria

Department of Microbiology, Wesley University Ondo, Nigeria

DOI:

https://dx.doi.org/10.51584/IJRIAS.2025.101100071

Received: 11 November 2025; Accepted: 18 November 2025; Published: 18 December 2025

ABSTRACT

With the increasing resistance of insects to chemical pesticides, regulatory pressure on chemical pesticides, and

the need for environmentally friendly and benign pest management systems, efforts are geared towards the search

for new entomopathogens. Traditional bioprospecting focuses on diseased insects in sourcing for microbes to be

used in formulating biopesticides, yet a wide range of non‑insect arthropods—copepods, arachnids (spiders and

mites), myriapods (millipedes, centipedes), and other arthropod lineages—harbour microbial pathogens (fungi,

oomycetes, microsporidia, bacteria, and viruses) with potentials to control insect pests. This manuscript reviews

the ecological rationales, discovery methods, promising taxa, safety considerations, and the processes for

sourcing novel entomopathogens from alternative arthropod reservoirs other than insects. We gave evidence that

spider‑pathogenic fungi (e.g., Gibellula spp.), entomophthoralean fungi associated with springtails and mites

(e.g., Pandora, Conidiobolus), oomycetes found on arthropods in aquatic environment (e.g., Lagenidium,

Leptolegnia), as well as microsporidia linked with aquatic crustaceans (copepods) all represent yet-to-be-fully

explored resources for both crop and veterinary pest management. We outline an integrated discovery framework

which combines high‑throughput field sampling, specialized culture strategies, functional screening in Galleria

mellonella and target pests, multi‑omics (amplicon, shotgun metagenomics, metabolomics), and genome mining

for virulence/secondary metabolite clusters. Finally, we highlight a regulatory and formulation roadmap to

de‑risk development while safeguarding non‑targets and aquatic ecosystems.

Keywords: entomopathogens; alternative reservoirs; arachnids; copepods; microsporidia; biological control

INTRODUCTION

Insect are known to be sometimes detrimental to economic crops, livestock and human health systems.

Controlling them appears arduous majorly due to their resistance to chemical insecticides which has continued

to escalate. Biological control using entomopathogens of fungal, viral, nematodal and bacterial origins do

provide an alternative to deal with this anomaly due to their distinct modes of action as well as lower resistance

risk (Bihal et al., 2023; Qin et al., 2023). When sourcing for biocides to be used as candidates of biopesticides,

researchers do focus on morbid insects and the edaphic environment. However, non‑insect arthropods have

equally been known to host rich, sometimes highly specialized pathogen communities that have been

taxonomically clarified and re‑evaluated in recent years (Hajek et al., 2024, 2025; Gryganskyi et al., 2024).

Systematically mining these alternative reservoirs could expand the spectrum of candidate agents for controlling

economically important insects and arachnids such as mosquitoes, locusts and ticks. These new candidates can

equally yield new metabolites (e.g., destruxins, proteases, chitinases) with bioinsecticidal value (Barelli et al.,

2022; Kuhnert et al., 2022; Vasquez et al., 2024).

Rationale For Exploring Non-Insect Arthropod Reservoirs of Entomopathogens

Non‑insect arthropods are known to share ecological niches, food webs, and fauna with target pest insects.

Predation, scavenging, and shared substrates (leaf litter, soil, biofilms, aquatic habitats) create reservoirs where

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host‑jump or closely related pathogens can be found (Bass et al., 2021; Pavanelo et al., 2023). In the aquatic

environment, oomycetes like Lagenidium and Leptolegnia parasitize aquatic arthropods and mosquito larvae,

thriving in ephemeral ponds and rice fields (Kaczmarek et al., 2021; Sila et al., 2023). Earlier research even

traced Lagenidium giganteum to mixed collections of copepods and mosquito larvae (Scholte et al., 2004).

Away from aquatic environment, the known spider‑pathogenic Hypocreales (e.g., Gibellula) and

Entomophthorales together infect spiders and mites; their infection process (cuticle penetration, toxin secretion)

mirrors that of an insect entomopathogenic fungus thus suggesting transferable virulence determinants (Joseph

et al., 2024; Sacco et al., 2023; Evans et al., 2025).

In the case of Collembola and mites, Entomophthoromycotina (e.g., Conidiobolus, Pandora) shows a narrow

host ranges yet a relatively broad arthropod coverage including springtails and mites—taxa sharing soils with

many crop pests (Gryganskyi et al., 2024; Sacco et al., 2023).

Myriapods such as millipedes act as hosts to actinomycetes and fungi with antagonistic activities and

biosynthetic capacity for metabolites potent against insects as well as their symbionts (e.g., Streptomyces

volatiles) (Stulanovic et al., 2025).

Potential Entomopathogen Groups from Alternative Arthropod Hosts

Oomycetes from aquatic arthropods

Lagenidium giganteum is a mosquito‑parasitizing oomycete capable of producing biflagellate zoospores that

actively seek larval hosts; re‑isolations from African rice fields demonstrate current relevance (Sila et al., 2023).

Biosafety studies do indicate variable non‑target effects, thus necessitating modern tiered toxicological and risk

assessment (Nestrud et al., 1994). They are known to produce hydrolases which facilitate infection (Bell &

Sayre, 1989).

Aedes and Culex larvae have been reported to succumb to Leptolegnia chapmanii. The agent often exhibits

temperature‑dependent efficacy and compatibility with other Integrated Pest Management practices. (Kaczmarek

et al., 2021).

Oomycete infections found in planktonic copepods also indicates an overlooked reservoir; these parasites share

aquatic niches with mosquito larvae, warranting targeted baiting and metagenomic surveillance (García et al.,

2018; Bass et al., 2021).

Entomophthorales from arachnids, mites and springtails

Erynioideae is a subfamily within the family Entomophthoraceae (order Entomophthorales) comprising obligate

arthropod-pathogenic fungi. Members within this subfamily are highly evovled entomopathogens capable of

causing rapid, often explosive epizootics in their hosts. Genera such as Pandora, Furia, Zoophthora,

Strongwellsea include around a hundred arthropod pathogens with many narrow host ranges. Using modern

phylogenomics, their taxonom, potential biocidal usage and host breadth can be better revised and understood

including records from mites and Collembola (Gryganskyi et al., 2024; Hajek et al., 2024; Sacco et al., 2023).

Spider‑pathogenic Hypocreales

The Gibellula spp which is a strict araneopathogens is getting increasingly reported from diverse climates. The

recent isolation and culture of Gibellula floridensis using modern methods has made researchers to better

understand its mode‑of‑action and metabolite discovery (Joseph et al., 2024; Evans et al., 2025). New species

continue to be discovered inside this group thus implying chemical/virulence novelty.

Bacterial symbionts and actinomycetes from myriapods and mites

Actinomycetes (e.g., Streptomyces) associated with myriapods produce a combination of both insecticidal and

certain antifungal volatiles. In addition, chitinolytic bacteria from both marine and terrestrial arthropod niches

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(e.g., Curtobacterium) exhibit antifungal activity as well as chitinase repertoires relevant to insect cuticle

degradation (Vasquez et al., 2024; Lai et al., 2024). Such microbes may act directly as entomopathogens or as

synergists/formulation adjuvants.

DISCOVERY AND SCREENING PIPELINE

Field sampling strategy

The first steps towards culturing new biocides involves identifying suitable habitats where the hosts are present.

Different habitats possess differing qualities which support the growth of different hosts and biocides depending

on their physiological and metabolic needs. Ecological habitats can be big or small in size including leaf litter,

riparian zones, temporary ponds, rice fields, bromeliad phytotelmata, cave drip zones, and under‑leaf

microhabitats frequented by spiders and mites. The diverse arthropod hosts (spiders, mites, copepods,

millipedes) can be sampled together with environmental substrates (soil, detritus, biofilms) (Joseph et al., 2024;

Bass et al., 2021) as all of these can house the biocides of interest.

Timing, seasonality and microclimate

Once a suitable habitat has been located, it is preferable to target humid, cooler morning periods for optimal

spore discharge especially for fungi within the Entomophthorales group. As regards the season, it is advisable

to target the rainy season and for sampling to be done during the post‑rain windows for oomycete zoospores

(García et al., 2018; Gryganskyi et al., 2024).

Isolation techniques and culture‑dependent enrichment

Most entomopathogens according to literature possess a relatively longer regeneration time which gives other

microbes especially the saprobes an earlier start when being cultured. By the time the entomopathogens are ready

to grow, the nutrient source would have been depleted and the culture space occupied. As a result, baiting

procedure through the direct use of sentinel hosts (mosquito larvae, Galleria mellonella larvae, Tenebrio molitor

pupae) in situ and ex situ to lure and attract pathogens from arthropod washes and microhabitats (Giammarino

et al., 2024). This attraction reduces contaminants and when the baits are cultured directly, it eliminates the

growth of unwanted microbes to a large extent.

Generally, entomopathogens sometimes require specialized media to be able to grow. For example,

semi‑selective plates are required for culturing oomycetes and certain entomopathogenic fungi. Media might

require supplementation using antibiotics to suppress the growth of unwanted bacteria colonies. For some strains,

oil overlays might be needed for optimal growth. ‑supplemented PDA/SDA; oil overlays for hydrophobins).

Some cultures also require incubation at 95–100% relative humidity (RH) using a saturated-moisture chamber

to ensure maximal water availability and prevent desiccation stress. This near-saturation humidity level is critical

for optimal sporulation, germination, and pathogen development, especially for entomopathogenic fungi and

other moisture-dependent microbes (Kaczmarek et al., 2021; Gryganskyi et al., 2024).

Molecular Approaches for The Identification of Novel Entomopathogens

Identifying novel entomopathogens greatly depends on molecular biology techniques, which usually offer a

much higher taxonomic resolution compared to morphological or ecological characteristics alone. The

identification process which solely relies on phenotypic traits alone is usually insufficient, particularly for

microbial groups with similar morphological features or cryptic species complexes which so far has not been

extensively studied. Therefore, confirmation using molecular methods is essential to reliably establish the

novelty of any putative entomopathogen (Hajek & St. Leger, 1994).

Polymerase chain reaction (PCR)-based amplification and sequencing of conserved genetic markers—such as

ITS rDNA for fungi, 16S rRNA for bacteria, and COI for insect-associated protists—provide definitive insights

into phylogenetic placement and species boundaries. Multi-locus sequence typing (MLST) and whole-genome

sequencing (WGS) have further improved taxonomic resolution, enabling differentiation among closely related

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strains and detection of previously unrecognized diversity (Kepler et al., 2014). Genomic techniques are

currently the gold standard for characterizing novel taxa and confirming their evolutionary distinctiveness for

many entomopathogenic fungi (Araújo & Hughes, 2016).

All things considered, reliable and repeatable identification is ensured by combining molecular technologies

with traditional taxonomy. Molecular characterization is essential for verifying the originality of

entomopathogens isolated from insect hosts since the designation of novel species depends on clear genetic

evidence.

ADVANCED FORMULATION

Next-generation formulations are primarily designed for protection and persistence. Key advancements that can

be employed for novel entomopathogens include Nanoformulations which entails the usage of nanoparticles

(e.g., chitosan, zeolite, or titania-based) as carriers which significantly shields the active microbial component

(spores, conidia, or nematodes) from degradation by UV light, temperature extremes, and desiccation. These

carriers can also provide a controlled-release system, synchronizing the EP release with the target pest's activity

cycle and thus extending the product's effective window in the field (Wu et al., 2023).

Oil-Based and Emulsion Formulations is another significant formulation method that can still be used for

emerging fungal biopesticides, formulations such as inverted emulsions (water-in-oil) enhance spore tolerance

to dry and warm conditions, improving germination and penetration of the insect cuticle (Batta, 2016). Oil-based

adjuvants can also improve foliar persistence for entomopathogenic nematodes.

The use of biodegradable carriers like hydrogels, gums (e.g., gum katira biogel), and biopolymers (e.g., cellulose

or gellan gum) can also be employed as it offers a safe, environmentally responsible way to package and protect

EPNs and microbial spores, replacing less eco-friendly synthetic alternatives.

Solid Formulations such as freeze and spray-drying technologies are being optimized to produce solid microbial

formulations that offer extended shelf life and easier transportation, crucial for commercial viability (Shapiro-

Ilan & Lewis, 2024).

DELIVERY STRATEGIES FOR NOVEL ENTOMOPATHOGENS

Innovation in the use of EPs has been sparked by the drive for precision agriculture, especially for pests that are

hard to reach. The majority of delivery strategies attempt to maximize target coverage, minimize waste

throughout the procedure, and assure correct application and successful interaction with target pests. Drones and

sophisticated precision spraying equipment have been used in the application of EPs in order to achieve this

goal.

Additionally, a relatively new and specialized method called "living insect bombs" releases live insect hosts that

have already been infected with EPNs. Effectively serving as mobile delivery systems, these "infected hosts"

transport the subsequent generation of entomopathogens into obscure environments (such as wood-boring

tunnels or beneath bark) that are inaccessible to traditional sprays (Gumus et al., 2015).

For soil and root pests, techniques like seed coating and the use of granular soil applications ensure EPNs or

fungal spores are delivered directly into the rhizosphere (root zone), their natural habitat, where they can

effectively intercept soil-dwelling pests.

To enhance immediate efficacy and broaden the target spectrum, EPs are increasingly being formulated for use

in Integrated Pest Management (IPM) programs, often in synergy with compatible, low-dose chemical

insecticides or other BCAs like parasitoids (Koller et al., 2023; Ma et al., 2025).

BIOSAFETY, NON‑TARGET RISK, AND ONE‑HEALTH FRAMING

Usage of entomopathogens in pest control is considered safe and nonlethal to the environment. However, a lot

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of toxicological and safety analyses is required to guarantee and ensure this not just on the biocides but also on

the formulation and delivery materials required. Every new candidate to be considered must be subjected to these

tests. Rigorous tiered testing is essential especially for aquatic systems (fish, amphibians, crustaceans) and

beneficial arthropods. Recent One‑Health analyses underscore the need for exposure modelling and case

surveillance for EPF/oomycete bioproducts (de Sousa et al., 2025; Nestrud et al., 1994).

REGULATORY AND STEWARDSHIP ROADMAP

A crucial pathway that is still complicated and varied worldwide is the regulatory roadmap for New Biocontrol

Agents (BCAs). Even the comparatively older entomopathogens still experience these complex regulatory

process which are initially created for chemical pesticides due to their toxicity. One major issue associated with

the regulatory framework of BCAs is the absence of specific, uniform rules for live creatures, which usually

leads to expensive and time-consuming regulatory processes that hinder commercialization and farmer uptake

(EFSA, 2025; Ravensberg, 2011). Creating simplified, risk-based assessment matrices that concentrate on

environmental safety, non-target effects, and human health concerns unique to the BCA's mode of action is part

of the reform movement that can facilitate the timely approval and usage of novel BCAs obtained from noninsect

sources (Wu et al., 2023). At the same time, stewardship is crucial, emphasizing the proper use of BCAs by

creating farmer-friendly e-learning materials, increasing regulators' and farmers' ability thus guaranteeing quality

promoting the successful long-term integration of BCAs into Integrated Pest Management (IPM) systems

(Hoeschle-Zeledon et al., 2013).

CONCLUSIONS AND FUTURE DIRECTION

Alternative arthropod reservoirs are fertile ground for discovering next‑generation entomopathogens and

synergists. Integrating ecological sampling with multi‑omics, genome mining and pragmatic screening can

accelerate the identification of strains with suitable virulence, specificity and manufacturability. Priorities

include moving beyond simple insect toxicity towards multi-functional biological agents that are deeply

integrated into sustainable agriculture. Others areas of focus are geared towards overcoming current limitations

including environmental instability, slow action and narrow host range. Although narrow hast range can be

advantageous in cases where specific pests are targeted in the midst of other beneficial species.

Formulations and deliver methods can also be improved upon with modern formulation science and discovery

of new delivery materials. The future of entomopathogens (EPs), is set to be dramatically accelerated and

transformed by Artificial Intelligence (AI) and its sub-fields, like Machine Learning (ML) and Deep Learning

(DL). AI’s ability to analyze massive, complex biological and ecological datasets will address key bottlenecks

in the discovery, optimization, production, and deployment of these biological control agents.

AI will assist with Predicting Virulence and host specificity by analyzing the genomes and transcriptomes of

novel EP isolates (e.g., fungi, bacteria, viruses) against large databases of known virulent factors and insect

immune responses. It will also assist with Novel Toxin Identification as DL models can be trained to analyze

the vast metabolome data of EPs (e.g., secondary metabolites from Metarhizium or Beauveria) to virtually screen

and prioritize novel bioactive micropeptides and toxins with insecticidal properties. AI can also be useful

in.Optimizing Multi-Role Agents in the case of dual-role pathogens (biopesticide and biofertilizer/antagonist).

Process Optimization and Mass Production can also be AI driven. Key environmental parameters such as

temperature, dissolved oxygen and media composition can be monitored in mass production bioreactors for EP

fungi (spore production) or Entomopathogenic Nematodes (EPNs) thus ensuring consistent quality, scalability,

and cost-efficiency (Ravensberg, 2011).

ACKNOWLEDGMENTS

The authors acknowledge the valuable contributions of Prof. Omoya, F. O. of the Federal University of

Technology Akure as well as Dr. Makinde O. of Adekunle Ajasin University Nigeria whose insights improved

the quality of this review. We also appreciate the institutional support and access to scientific literature made

possible by Adekunle Ajasin University Nigeria that made this work possible. No external funding was received

for the preparation of this manuscript.

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