Microbial gatekeepers: midgut bacteria in Aedes mosquitoes as modulators of arboviral transmission and targets for sustainable vector control

Arboviral diseases such as Dengue virus, Zika virus, Chikungunya virus, and West Nile virus pose significant global public health and economic challenges, particularly in tropical and subtropical regions. The absence of effective vaccines and sustainable vector control strategies continues to drive high morbidity and mortality rates. Symbiotic bacteria residing in the mosquito midgut can produce antimicrobial compound, stimulate the host immune response, disrupt nutrient pathways critical for pathogen development, and interfere with the pathogen’s lifecycle and dissemination. Additionally, these microbes may reduce vector reproduction and shorten the lifespan of both immature and adult stages. Genetically modified symbiotic bacteria can release effector molecules that target pathogens without harming mosquitoes. Advances in genomic and metagenomic tools have deepened our understanding of the mosquito gut microbiome. This review highlights current knowledge of gut bacteria and arbovirus interactions and explores strategies to reduce arboviral transmission. Comprehensive literature searches were conducted using global databases, including PubMed, Web of Science, and Scopus, with a focus on English-language publications.

1 Introduction

Vector-borne diseases continue to pose a significant global public health challenge, particularly in tropical and subtropical regions. Despite the implementation of various intervention strategies to control these diseases, their impact remains substantial. According to the World Health Organization (2024) report, vector-borne diseases account for more than 17% of all infectious diseases and cause over 700,000 annual deaths globally. Among these, malaria alone accounts for 249 million cases and 608,000 deaths, while the remaining cases are attributed to arboviral diseases (World Health Organization, 2024).

These diseases are primarily transmitted by mosquitoes belonging to three genera: Anopheles, Culex, and Aedes. Anopheles mosquitoes are vectors for Plasmodium spp. (malaria), Culex primarily transmits filarial worm infections and West Nile virus (WNV). In contrast, Aedes species are the primary vectors of arboviruses, including Dengue virus (DENV), Zika virus (ZIKV), Chikungunya virus (CHIKV), and Yellow fever virus (YFV) (Girard et al., 2020). Arboviruses have emerged as significant public health threats due to their potential to cause explosive outbreaks and severe, sometimes life-threatening, clinical conditions (Challenges in Combating Arboviral Infections, 2024).

Among 950 Aedes species, Aedes aegypti and Aedes albopictus are the most efficient and widespread vectors for DENV, ZIKV, CHIKV, and YFV (De De Curcio et al., 2022; Leta et al., 2018). This is due to their adaptability to urban environments and global distribution, which contribute significantly to arboviral disease transmission. In addition, although Aedes japonicus is not a significant vector for arboviruses to humans, it has been collected from the field and tested positive for WNV, La Crosse, and Usutu viruses (DeCarlo et al., 2020).

Another species, Aedes koreicus, is native to East Asia and has recently become an invasive species in parts of Europe. It has shown potential as a vector for Dirofilaria immitis, Brugia malayi, and CHIKV (Ganassi et al., 2022). In urban areas of northern Italy, this species has been observed feeding on human blood (Montarsi et al., 2022), Further suggesting its role in arboviral transmission. Aedes vexans is another Aedes mosquito species native to Eastern Europe and a potential vector for WNV, ZIKV, and Rift Valley fever virus (RVFV) (Birnberg et al., 2019).

The primary strategies for arbovirus control rely on insecticide-based interventions, such as indoor residual spraying (IRS), space spraying, and the utilization of insecticide-treated bed nets (ITNs). However, the widespread development of insecticide resistance has significantly reduced the effectiveness of these methods (Girard et al., 2020; Minwuyelet et al., 2025). Besides chemical insecticides, vector control through habitat removal, the use of repellents, and other biological controls remain the second line of defense against arbovirus vectors. While these approaches have had some success, no single strategy has proven sufficient to control mosquito populations or eliminate arboviral transmission (Gao et al., 2020).

Considering these challenges, alternative, eco-friendly strategies are being explored. One promising avenue is the manipulation of the mosquito microbiome. Recent studies have revealed that mosquitoes harbor diverse microbiota, particularly in their gut, forming symbiotic relationships mosquito host. This microbiota can influence pathogen transmission by interacting with pathogen antagonistically or indirectly. The gut microbiota plays a crucial role in key physiological and metabolic processes in mosquitoes, including blood digestion, nutrient acquisition, reproduction, and immune modulation (Harrison et al., 2023).

The commensal and pathogenic microbiome colonization in Aedes mosquitos starts in early larval stages, where the aquatic environment plays a critical in shaping microbial community in midgut. During mosquito colonization, a competitive interaction occurs between commensal and pathogenic bacteria for niche establishment. While certain bacterial strains successfully establish stable symbiotic associations within specific mosquito tissues, others persist as pathogens, either causing infections in the mosquito host or exploiting the mosquito as a vector to transmit vector-borne diseases (Cai and Christophides, 2024). Microbial communities also influence development, particularly during the transition from larva to adult (Alfano et al., 2019). Aedes mosquito first instar larvae which grow in aseptic condition cannot survive (Coon et al., 2014). In addition, depletion of the microbiota during the larval stage significantly impairs developmental progression, leading to delayed pupation and adult emergence (Chouaia et al., 2012).

As mosquitoes transition from larvae to adults, microbial communities are maintained through transstadial transmission and environmental exposures such as sugar and blood meals. Both commensal and pathogenic microbes acquired through different feeding regimes and environmental exposure activates systemic immune responses in mosquitoes (Sharma et al., 2020). The interplay between microorganisms for nutrition and resource can modulate robust immune priming in the adult mosquito, notably through the production of antimicrobial peptides such as defensins and cecropins, regulated primarily by the Toll and IMD immune pathways (Cirimotich et al., 2011a).

In Anopheles mosquito gut microbiota induce systemic immunological response that limit the abundance and distribution of microorganism, and RNAi-mediated silencing of AMPs and immune signaling pathways has been shown to result in increased proliferation of the gut microbiota (Dong et al., 2009; Clayton et al., 2014). Similarly in Aedes mosquito proliferation of microbiota following blood meal activate IMD pathway and limits sindbis virus infection (Barletta et al., 2017). Moreover, studies show that certain bacteria in mosquito gut can either enhance or inhibit infections, depending on their interactions with both the pathogen and host immunity (Boissière et al., 2012; Ramirez et al., 2014; Wu et al., 2019).

Recent scientific advancements offer a novel approach to address this long-standing problem by harnessing the potential of gut microbiome in Aedes mosquitoes. A promising technique involves modifying the gut microbiome of mosquitoes to diminish their ability to transmit viruses, which are responsible for arboviral diseases (Dickson et al., 2018; Gao et al., 2020; Hegde et al., 2015).

This review synthesizes current research on the composition and factors related to the gut bacteria of Aedes mosquitoes, revealing its role in influencing arboviral transmission dynamics and evaluating emerging strategies using microbial communities for sustainable vector control. By integrating insights into microbiota-pathogen interactions and innovative interventions, the review aims to bridge gaps in understanding how microbial manipulation can disrupt arboviral spread and address insecticide resistance, ultimately informing next-generation, eco-friendly interventions for global arboviral disease mitigation.

2 Methodology

The literature search focused on primary articles, published between 2010 and 2025. The review research covered topics related to the bacterial composition of Aedes mosquitoes, factors influencing bacterial diversity, interactions between Aedes gut symbiotic bacteria and arboviruses, and their potential role in vector control. Articles were identified using Boolean operators “AND” “OR” and “NOT” in the search strategies. Key words such as Aedes gut microbiota, symbiotic bacteria, arbovirus, and vector control were used either separately or in combination. Studies were excluded if they focused on mosquito vectors other than Aedes mosquitoes, examined non-bacterial components of microbiome, or lacked clear methodologies for bacterial identification. Relevant articles published in English were identified using databases such as PubMed, Web of Science, and Scopus. The final search was conducted between January 30 and February 15, 2025. Data were extracted by analyzing the text, figures, and tables from the included articles. In this review, after examining 219 primary articles, we retrieved 72 articles (see Figure 1).

Figure 1

Figure 1. The flowchart of search and selection of articles for review of Aedes mosquito microbiota, factors and role in modulating arboviral transmission and vector control.

3 Gut microbiota of Aedes mosquito

3.1 Acquisition of gut microbiota in Aedes mosquito

The mosquito microbiome comprises a diverse community of bacteria, fungi, and insect-specific viruses that reside within and may spread through various mosquito tissues (Pascar et al., 2023; Guégan et al., 2018). While the majority of these microorganisms are found within the gut, they are also found in other somatic and germline tissue such as the salivary gland, crop, reproductive tract and cuticle of Aedes mosquitoes (Onyango et al., 2021; Valiente Moro et al., 2013).

Mosquitoes can acquire their microbiota vertically from their parents. Various species of mosquitoes can vertically transmit intracellular bacteria, such as Wolbachia, from one generation to the next (Caragata et al., 2022). In contrast, several studies reported that microbiota are also acquired horizontally from the surrounding environment including aquatic habitat and feeding sources. Additionally, some microbial communities are transmitted via the egg surface (Coon et al., 2016). Upon hatching, first instar larvae ingest fragments of the eggshell, thereby acquiring microbes from the egg-associated microbiome (Gimonneau et al., 2014; Figure 2).

Figure 2

Figure 2. Summary of gut microbiota acquisition through vertical and horizontal transmission.

3.2 Composition of bacteria in Aedes mosquito

Bacteria represent the primary components of the mosquito gut microbiota, followed by fungi, algae, and viruses to a lesser extent (Guégan et al., 2018; Cansado-Utrilla et al., 2021). We identified twenty-five articles that focused on the bacterial composition of Aedes mosquito vectors. The articles included in this study used both culture-dependent and culture-independent methods. Two studies employed culture-dependent techniques, while the others utilized culture-independent approaches based on current molecular strategies such as 16S rRNA gene sequencing and metagenomic analysis, which have become essential tools for characterizing the bacterial microbiota in the mosquito gut.

Both culture dependent and culture independent studies confirmed that Aedes mosquitoes harbor a wide range of both classified and unclassified bacterial taxa associated with the gut (Bennett et al., 2019; Baltar et al., 2023). Among these, the most prevalent bacterial phyla identified in Aedes mosquitoes include Proteobacteria, Firmicutes, Actinobacteria, and Bacteroidetes (Muturi et al., 2021a; Pascar et al., 2023).

Several studies conducted in the United States, consistently reported Proteobacteria as the dominant and highly diverse bacterial phylum in Aedes mosquitoes. Proteobacteria were a highly diverse and dominant phylum in both the midgut and saliva of Aedes mosquitoes (Pascar et al., 2023; Onyango et al., 2021). Similarly, Proteobacteria remained dominant phyla across the mosquito population regardless of variation in their aquatic habitats and blood meal sources (Caragata et al., 2022; Muturi et al., 2018).

The pattern was consistent with findings from India, where Proteobacteria as the dominant phylum in the gut of Ae. aegypti (Yadav et al., 2016; Sarma et al., 2022). Additional studies from diverse locations, including Brazil, Korea, and China, have corroborated the widespread dominance of Proteobacteria in the gut of Aedes mosquitoes (Akintola and Hwang, 2024; Baltar et al., 2023; Lee et al., 2020). This widespread dominance may result either from the insect host’s active recruitment of these bacteria that support its growth or from the greater ability of Proteobacteria to invade and proliferate within new insect hosts compared to other bacterial groups.

One of the most extensively studied genera within the Proteobacteria phylum is Wolbachia, a maternally inherited endosymbiont with critical implications for mosquito biology and vector competence. Studies have consistently reported Wolbachia as an abundant midgut bacterium in Aedes mosquito although the proportion and prevalence vary study and may depend on environmental, geographic, or methodological factors. A study from Spain and São Tomé found Wolbachia to be overwhelmingly dominant with 92.4–98. 8% in Sao Tome and 96.1–97.5% in Spanish samples, with 77.22% of mosquitoes co-infected with both wAlbA and wAlbB strains (Melo et al., 2024).

A similar study from Mexico, showed that Wolbachia accounted for 9.6% of 16S gene sequences, with the abundance 0 to 32% in each sample. A high prevalence of the wAlbB strain, and included genes linked to Cytoplasmic Incompatibility (CI) was detected (Hernández et al., 2024). Molecular approaches using Wolbachia specific primer and strain specific genetic marker essential for strain specific comparison and identification of genes related to CI.

Species and tissue specific occurrence of Wolbachia has been reported in different studies. A metagenomic analysis reported that the Prevalence of Wolbachia was 100% in Ae. albopictus and Cx. pipiens but not in other Aedes or Anopheles species. In addition to this Wolbachia was found to be more abundant in reproductive tissues where as Asaia was predominantly detected in the gut (Ilbeigi Khamseh Nejad et al., 2024). Similar study from Southern China reported that Wolbachia was more abundant in the whole body of Ae. albopictus than in the midgut. Additionally, microbiota network analysis revealed Wolbachia have both positive and negative co-occurrences with other bacterial genera (Lin et al., 2021). For example, Wolbachia and Asaia shows mutual exclusion in species and host tissue level (Rossi et al., 2015). This phenomenon has important implications for symbiont-based vector control strategies.

Studies from South Korea and Malaysia reported that Wolbachia was the most dominant genus, accounting for 98.36% of the midguts of Ae. albopictus with relative abundance in female and 70.5% of the bacterial community in the midgut of Ae. albopictus mosquitoes with relative abundance higher in male, respectively, (Lee et al., 2020; Ilbeigi Khamseh Nejad et al., 2024). Similar study from Brazil observed higher detection rates of Wolbachia in field-collected mosquitoes, particularly during the dry season (Baltar et al., 2023).

Likewise, a study in southern Thailand also reported Wolbachia prevalent in both sexes of Ae. albopictus, with greater abundance in males (Rodpai et al., 2023). Sex-based microbiota profiling, sample pooling, mosquito collection season, and geographical location might cause variation to abundance. Furthermore, due to methodological differences, Wolbachia is an intracellular bacterium that is not grown in artificial media and detected in culture-dependent studies (see Supplementary Table 1).

Another important bacterial genus in the Aedes gut microbiota is Enterobacter. The presence of Enterobacter was detected in the gut of adult Aedes mosquitoes collected from field but not in the egg or larval stages (Hernández et al., 2024). Contrastingly, study from Italy reported Enterobacter was detected in both Bacillus thuringiensis israelensis (Bti) exposed adults and larvae of Ae. albopictus, with a significantly higher abundance in Bti-resistant larvae (Bahrami et al., 2024). Similarly, study from Brazil reported a 3%, prevalence of Enterobacter isolated only from the eggs of Ae. aegypti mosquitoes that obtained from laboratory colony (Gusmão et al., 2010).

The variation in Enterobacter abundance across different developmental stages may be attributed to differences in sample sources and exposure to Bti larvicide, as Enterobacter has been previously associated with increased insecticide resistance. Meanwhile, a study from southern China reported that Enterobacter was present in both the entire body and midgut of both Aedes mosquitoes. In terms of abundance, it was more prevalent in the midgut of female Ae. albopictus than in its whole body (Lin et al., 2021). Additionally, a study from Thailand found that Enterobacter was present in all groups tested for CHIKV infection; however, its presence did not significantly correlate with infection status (Siriyasatien et al., 2024).

A study from the USA has shown that Enterobacter was the dominant genus among the five found in Ae. aegypti mosquitoes that fed on different blood meal sources (Muturi et al., 2021a). On the other hand, studies from India and Madagascar reported that Enterobacter was the second most dominant bacterium isolated in both sugar-fed female and male mosquitoes (Valiente Moro et al., 2013; Yadav et al., 2016). Furthermore, studies from Thailand and India reported that species like Enterobacter cloacae were particularly dominant in both Ae. aegypti and Ae. albopictus across field and lab populations (Yadav et al., 2015; Tuanudom et al., 2021). Enterobacter is symbiotic bacteria commonly detected in the gut of Aedes mosquito regardless of host species, method of isolation, and diet. This promotes microbial stability through beneficial co-occurrences in mosquito guts.

The genus Asaia, another member of the Proteobacteria, also plays a crucial role in the microbiota of Aedes. Studies from Iran detected Asaia in the midgut of field collected Ae. albopictus (Darbandsari et al., 2025). Roman et al. demonstrated that Asaia can accelerate the growth of Ae. aegypti larval development and interact with the broader larval microbiome (Roman et al., 2024). Interestingly, study from Thailand, found Asaia in CHIKV negative and control groups, but not found in infected mosquitoes (Siriyasatien et al., 2024). Similar study from the USA also reported variable Asaia spp. presence in Ae. aegypti populations with differing DENV susceptibility, although the role of Asaia spp. in antiviral defense remained unclear (Chen et al., 2023). The observed difference between infected, and uninfected groups mosquitoes can a possible association implying that Asaia may play a protective or modulatory role in vector competence. Further experimental infection studies are important to elucidate the association. Asaia was the most abundant genus in the Ae. aegypti sample that had been treated with a blood meal containing Amox/Clav and was reported as resistant to it (Van Garcia et al., 2024). It was found in Aedes, Anopheles, and Culex species, with varying prevalence depending on geographical location and mosquito species (Ilbeigi Khamseh Nejad et al., 2024). In Ae. aegypti Asaia was abundant in the crop than in the midgut (Villegas et al., 2023). Its abundance across has been reported at low and fluctuating levels across the regions such as Italy, Spain, and São Tomé (Ilbeigi Khamseh Nejad et al., 2024; Melo et al., 2024).

Other bacterial genera within Proteobacteria frequently detected in Aedes mosquitoes include Pseudomonas, Serratia, Pantoea, Klebsiella, and Aeromonas, as reported by multiple studies across the globe (Brettell et al., 2025; Darbandsari et al., 2025; Pascar et al., 2023; Muturi et al., 2021a; Rosso et al., 2018; Minard et al., 2015).

Firmicutes represent the second most abundant phylum in many studies. A study from the USA reported that Firmicutes accounted for 36.6% of Ae. aegypti microbiota, Bacillus and Clostridium were found in the midgut with Bacillus subtilis being the most dominant species at 42.4% (Pascar et al., 2023). Similar finding was reported in China Bacillus and Clostridium were present in both Bti-resistant and control larvae, with Bacillus being the predominant genus (Bahrami et al., 2024). Firmicutes were also the second most abundant phylum (27.2%) in whole-body microbiota of Ae. albopictus, with Bacillus dominating (22.9%). In contrast, tissue specific comparative analysis showed Bacteroidetes as the second most prevalent phylum, indicating variation in microbial composition across different tissues(Lin et al., 2021).

In contrast, Actinobacteria was the second most dominant phylum (11.3%), followed by Firmicutes (10.3%), Bacteroidetes (5%) and Cyanobacteria (1.3%) in Ae. aegypti. In this study Bacillus, Lysinibacillus, and Clostridium as common genera detected in adult (Hernández et al., 2024). Similarly high levels of Actinobacteria were detected in both laboratory-reared and field-collected Ae. albopictus (Tuanudom et al., 2021). Acinetobacter consisted of 17% of Ae. albopictus bacterial community, while Bacteroidetes was the least represented phylum, characterized by a single species, Chryseobacterium rhizoplanae, isolated from blood-fed individuals (Yadav et al., 2016).

Actinobacteria and Firmicutes were commonly found in larvae and breeding sites, however the mosquito gut appears more selective toward these bacterial groups. For example, Staphylococcus, Bacillus, and Clostridium are more likely associated with hindgut or body surface than midgut lumen (Ngo et al., 2016). Under laboratory condition larvae fed controlled larval diet, organic matter is limited, Firmicutes are less supported, whereas Actinobacteria tend to persist and adapt well to these stable, low-diversity microbiota environments.

Bacteroidetes were present in lower abundance in most studies, but its enrichment in mosquito gut associated with bloodmeal, Elizabethkingia with a dominant genera (Sharma et al., 2020). Variation in microbial abundance between species and across geographic regions has also been reported by Pascar et al. (2023). Bacteroidetes were 4.7 and 1.5% of Actinobacteria in Ae. aegypti mosquitoes. Actinobacteria and Bacteroidetes were present in Ae. albopictus mosquitoes in low abundance, but their abundance was high in Culex mosquitoes (Akintola and Hwang, 2024).

Similarly, Bacteroidetes were detected in both the midgut and saliva, bacteria belonging to the genus Elizabethkingia were enriched in ZIKV-infected midguts. In contrast, Wolbachia was abundant in non-infected midguts (Onyango et al., 2024). Elizabethkingia enrichment in infected mosquito midguts suggests a host-pathogen interaction, potentially involving an antiviral mechanism that influences viral replication. While Wolbachia prevalence in uninfected mosquitoes associates with mosquito immunity and suppressing arbovirus infection and replication.

In addition to the factors related to the abundance of certain bacteria in mosquito characterization of bacteria could biased by the techniques used for studying microbiota such as DNA extraction method, primer selection, sequencing platform and bioinformatics pipeline. Level of variability within the 16S rRNA genes also making it difficult to distinguish them in species or strain level. This might cause underestimation or over estimation of certain bacteria (see Supplementary Table 1).

3.3 Factors that shape gut microbiota of Aedes mosquito

A total of twenty-seven articles were retrieved that examined the various factors influencing the mosquito microbiome. Recent studies have shown that the microbial communities of Aedes mosquitoes vary significantly depending on several intrinsic and extrinsic factors, including mosquito species, developmental stage, sex, larval diet, and the environment of the breeding site.

For instance MacLeod et al. (2021) found that adult mosquitoes emerging from larvae reared on a nutrient-rich diet exhibited a significantly higher bacterial load in both their midguts and breeding water. Specifically, increased dietary abundance was associated with elevated levels of Enterobacteriaceae and Flavobacteriaceae and a decrease in Sphingomonadaceae. Larval nutrition not only affects growth and development but also influences microbial colonization. A significant increase in Enterobacteriaceae in larvae-fed pelleted diets however, Flavobacteriaceae levels remained essentially unchanged (Linenberg et al., 2016).

Martinson and Strand (2021) showed that larvae fed a complete bacterial community alongside nutrient-rich food exhibited distinct microbial profiles. Similarly, variation in midgut bacterial communities across developmental stages, sexes, and feeding conditions has been reported. For example, Acinetobacter pitti was abundant in sugar-fed females and larvae, while Pseudomonas monteilii dominated in blood-fed mosquitoes. Pantoea was prominent in adult males, whereas Chryseobacterium rhizoplanae, the only Bacteroidetes species isolated, was found exclusively in blood-fed Ae. albopictus (Yadav et al., 2016).

Environmental exposure during larval or adult stage also plays a significant role in diversity of microbiota. Scolari et al. found that over 60% of the bacterial genera was conserved in both larval and adult Ae. albopictus were also present in breeding site water (Scolari et al., 2021). Similarly, Alfano et al. (2019) reported that 84% of the bacterial communities in the mosquito gut were varied across breeding sites, larvae, pupae, and adults, with notable shifts in dominant taxa from the larval to adult stages.

Juma et al. (2021) observed that larval sampling environments significantly influenced microbial communities in Ae. triseriatus and Ae. japonicus, with Dysgonomonas being the dominant genus in Ae. triseriatus, while Mycobacterium and Carnobacterium were dominant in Ae. Japonicaus. Unclassified Comamonadaceae was dominant in water samples (Rodpai et al., 2023) confirmed that the composition of microbiota varies significantly across developmental stages and between Ae. aegypti and Ae. albopictus. While transstadial transmission of microbiota was observed, adult mosquitoes showed a reduced bacterial load compared to larvae. Microbiota also varies species to species, Wolbachia was more abundant in Ae. albopictus, whereas Blautia was enriched in Ae. aegypti.

Blood feeding has a profound effect on gut microbiota. Sarma et al. (2022) demonstrated a significant difference in the gut microbiota of Ae. aegypti depending on feeding status: Rhodobacterales and Neisseriales were enriched in mosquitoes fed with human blood, while Caulobacterales dominated in unfed mosquitoes. Supporting this finding, Muturi et al. (2021a) reported that the blood source influenced the composition of midgut microbiota. For example, newly emerged adults and those fed on chicken, rabbit, and human blood were characterized by Leucobacter, Chryseobacterium, Elizabethkingia, and Serratia, respectively, Whereas sugar-fed mosquitoes harbored more Pseudomonas.

Salgado et al. (2024) reported lower microbiota diversity in blood-fed mosquitoes compared to sugar-fed ones, with blood digestion dominated by Enterobacterales, followed by a rise in Elizabethkingia anopheles post-digestion. LaReau et al. (2023) highlighted taxonomic and functional differences between axenic mosquitoes colonized by environmental bacteria and those reared in insectaries. The former showed greater diversity and dynamic shifts during blood feeding and could even perform hemolysis in culture.

The composition and diversity of microbial communities in both larvae and adult mosquitoes are influenced by the colonization of microorganisms. Frankel-Bricker et al. (2020) demonstrated that the fungal colonization of the gut by Zancudomyces culisetae in larvae reduced microbial diversity in adults and affected the transmission of specific bacterial genera. Similarly, (Yin et al., 2025) demonstrated that inoculation with Escherichia coli, Staphylococcus aureus, and Beauveria bassiana altered the midgut microbiota across different stages.

Wei et al. (2017) reported that B. bassiana infection in mosquitoes induced gut dysbiosis, increasing bacterial load while reducing diversity. The gut became dominated by Acinetobacter, Serratia and Asaia, with Serratia marcescens overgrowth leading to translocation into the hemocoel and increased mortality. Wolbachia infection in Ae. aegypti also caused microbiome shifts and negatively interacted with other taxa (Pascar et al., 2023). Notably, Serratia was enriched in Wolbachia-infected mosquitoes, while Pseudomonas and Acinetobacter dominated in Wolbachia-free individuals (Balaji et al., 2021).

Viral infections also influence gut microbiota. DENV infection modulates bacterial abundance in Ae. aegypti, upregulating Desulfovibrionaceae and Enterococcus gallinarum while reducing overall bacterial load (Zhao et al., 2022). Similarly, (Ramirez et al., 2012) showed that DENV infection significantly decreases the overall bacterial load in the midgut of Ae. aegypti mosquito.

In addition to viral infections, chemical insecticides also significantly alter the microbiota of mosquitoes. Arévalo-Cortés et al. (2020) observed reduced gut diversity in both ZIKV-infected and lambda-cyhalothrin–resistant mosquitoes. Bacteroides vulgatus were enriched in ZIKV-infected groups, while Pseudomonas viridiflava and Clostridium ramosum were found in resistant mosquitoes. Additionally, Wei et al. (2023) demonstrated that pyrethroid exposure resulted in microbial enrichment or depletion, with genera such as Butyricimonas, Prevotellaceae, Anaerococcus, and Pseudorhodobacter significantly reduced.

Resistance mechanisms also drive microbiota shifts. Ae. aegypti resistant to permethrin showed different gut microbiota compared to susceptible strains (Muturi et al., 2021a). Viafara-Campo et al. (2025) found that deltamethrin-resistant females and temephos-treated larvae had distinct microbiota, with Enterobacter predominant in untreated females and resistant larvae, Bacillus exclusive to larvae, and Serratia, Cedecea neteri, and Elizabethkingia exclusive to resistant females. Sun et al. (2024b) similarly reported a higher abundance of certain gut symbiotic bacteria in deltamethrin field-resistant adults compared to sensitive adults; however, both field-resistant and field-sensitive adult mosquitoes exhibited significantly reduced gut microbiota diversity compared to laboratory-sensitive adults.

Antibiotic exposure also alters gut microbiota. A study by Qing et al. (2020) reported that ampicillin exposure in Ae.albopictus across developmental stages caused gut dysbiosis, particularly in adult females. Van Garcia et al. (2024) demonstrated that the ingestion of antibiotics during blood meals reduced microbial diversity, particularly in field-collected mosquitoes. Co-exposure with DENV-modified bacterial composition: Pseudomonas and Asaia decreased, while Enterobacter increased. Minard et al. (2015) also observed that larval antibiotic exposure led to a reduction in Elizabethkingia, elimination of Chryseobacterium and an increase in Wolbachia in adults.

Environmental pollutants, such as polycyclic aromatic hydrocarbons (PAHs), can also impact gut microbiota (Antonelli et al., 2024) reported stage-specific effects of chronic PAH exposure in Ae. albopictus with a greater impact on larvae. PAH exposure enriched bacterial families capable of PAH degradation, altering competitive dynamics in the gut. Moreover, CRISPR/Cas9 mediated deletion of bacteria ompA genes impaired colonization capability (Hegde et al., 2019).

Geographic distribution and environment also influence the composition of microbiota. Minard et al. (2015) found that mosquitoes invading new geographic areas had reduced microbial diversity compared to those from native regions. Brettell et al. (2025) showed that Ae. aegypti reared in different insectaries from eggs laid at the same time exhibited significantly different gut microbiota despite similar development.

Similarly, Ae.albopictus collected from Spain and São Tomé shared core microbiota but had location-specific genera, including different Wolbachia strains (Melo et al., 2024). Baltar et al. (2023) observed differences between lab colonies and field-collected mosquitoes, with gut microbiota diversity decreasing from wet to dry seasons (see Figure 3; Supplementary Table 2).

Figure 3

Figure 3. Factors that shape the gut microbiome composition of Aedes mosquitoes.

3.4 Role of mosquitoes’ guts microbiota in modulating pathogen transmission

During blood feeding, mosquitoes might ingest pathogens, particularly Plasmodium parasites and/or arboviruses, which first enter the mosquito’s midgut. These pathogens penetrate the midgut epithelial cells, spread into the hemocoel, and ultimately cross the salivary gland barrier, gaining access to the saliva for transmission during subsequent bites (Mueller et al., 2010). Vector competence refers to the intrinsic ability of a mosquito to acquire, maintain, and transmit pathogens to another host. This is a complex biological trait influenced by various intrinsic and extrinsic factors, including the mosquito’s genetics and associated microbiota. The interaction between the mosquito genotype and its microbiota plays a crucial role in modulating vector competence (Cansado-Utrilla et al., 2021). Sixteen recent articles addressed the role of mosquito microbiota in pathogen transmission.

Studies have shown that the presence of a certain bacteria particularly a member of Rickettsiaceae, Enterobacteriaceae, and Flavobacteriaceae family can be corelated with reduced arboviral infection in mosquitoes(Kukutla et al., 2014; Apte-Deshpande et al., 2012; Moreira et al., 2009). The underlying mechanism by which the gut microbiome in aedes mosquito is not fully understood, but they are believed to involve both direct and indirect interactions.

Gut microbiota in mosquitoes can modify the gut environment by secreting antiviral metabolites and modulating the mosquito’s immune response, thereby inhibiting arbovirus entry, replication, and transmission. For example, Rosenbergiella YN46, found in field-collected Ae. albopictus has been shown to colonies the mosquito gut consistently. This bacterium secretes glucose dehydrogenase (RyGDH) enzyme, which changes glucose to gluconic acid during blood digestion. The accumulation gluconic acid in mosquito gut change the gut lumen to acidified environment which inactivate viruses and significantly inhibits invasion of DENV and ZIKV gut epithelial cells (Zhang et al., 2024) Similarly, Enterobacter hormaechei B17 (Eh_B17), a symbiotic gut bacterium, consistently colonizes the midgut of female mosquitoes after transplantation. Eh_B17 produces metabolite sphingosine, which significantly inhibits the early stages of DENV and ZIKV entry into host cells (Sun et al., 2024a).

Symbiotic bacteria can inhibit pathogen transmission by computing pathogen essential resources that are important for growth, replication and transmission. Wolbachia, an intracellular symbiotic bacterium, is widely used in mosquito control strategies and demonstrates antiviral properties. Transient somatic infections with Wolbachia strains wAlb and wMel significantly reduced Mayaro virus (MAYV) infection and viral titters in a strain-specific fashion. However, Wolbachia causes enhancement to Sindbis virus infection (Dodson et al., 2024). Wolbachia, alters cholesterol metabolism by diverting host resources from the mevalonate (MVA) pathway and downregulating cholesterol esterase genes, which are typically upregulated during ZIKV infection. This metabolic disruption depletes lipid droplets and inhibits ZIKV replication within mosquito cells (Edwards et al., 2023).

Wolbachia infection primarily blocks virus transmission, the mechanism is not fully explored yet, it could be activating the mosquito’s innate immune system or outcompeting with intracellular resource. Wolbachia strain NC-wMel, derived from crosses between Australian wMel females and New Caledonian wild-type males, and wMel-Sg from Singapore significantly reduced susceptibility to and blocked transmission of ZIKV, DENV, and CHIKV in Ae. aegypti. Notably, mosquitoes infected with NC-wMel exhibited complete CI and efficient maternal transmission (Pocquet et al., 2021; Tan et al., 2017). Similarly, populations of Wolbachia-infected Ae. aegypti (wMel), both in the field and in the laboratory, showed a significant reduction in DENV transmission potential and experienced an extended extrinsic incubation period of 4–7 days (Carrington et al., 2018).

Mosquito associated symbiotic bacteria also modify arboviral transmission by altering the expression or function of conserved mosquito proteins required for viral entry, replication and attachment. For example, Wolbachia (wAlbB) inhibits DENV-2 replication, Aag-2 Cells. wAlbB inhibited virus genome replication by blocking synthesis of the viral negative-strand RNA. In addition to this wAlbB inhibit DENV binding to Aag-2 cells by downregulating transcription of host membrane binding protein dystroglycan and beta-tubulin (Lu et al., 2020). Pelo protein is a conserved protein in insects involved in immune regulation, promoting Drosophila C virus replication in D. melanogaster (Wu et al., 2014). Wolbachia-infected Ae. aegypti females (wMelPop-CLA), showed reduced expression of Pelo and altered subcellular localization, which could potentially contribute to decreased DENV replication (Asad et al., 2018).

Symbiotic bacteria in the mosquito gut can produce natural toxins, antiviral compounds, or metabolites that prevent viruses from attaching to the gut lining and promote the degradation of viral genomes before attachment. The previous ingestion of Chromobacterium sp. Panama (Csp_P) by mosquitoes significantly reduced susceptibility to P. falciparum and DENV infection, both in vitro and in vivo (Ramirez et al., 2014). In support of this, Csp_Panama exhibits an inhibitory effect on DENV replication both in mosquitoes and in-vitro. Neutral protease and amino-peptidase enzymes destabilize the virus by degrading the viral envelope protein. This degradation of the viral envelope protein inhibits viral attachment to the host cell (Saraiva et al., 2018).

Likewise, Chromobacterium sp. Beijing (Csp_BJ), isolated from Ae. aegypti produces two antiviral effectors, CbAE-1 and CbAE-2, with conserved lipase domains. These lipases disrupt viral envelopes, thereby inactivating DENV, Japanese encephalitis virus (JEV), YFV, and ZIKV. Furthermore, high doses of Csp_BJ administered orally result in significant mortality in mosquitoes (Yu et al., 2022). Prostaglandins (PGs), immune-active lipids, are produced by midgut tissues in response to microbiota and play crucial roles in mosquito immunity. Enterobacter cloacae triggers PG production in the midgut of Ae. aegypti and in Aag2 cells, which in turn enhance antiviral immune responses against DENV (Barletta et al., 2020).

Introduction of symbiotic bacteria isolated from mosquito guts of antibiotic-treated mosquito shows a significant role in modulating viral replication.,by boosting mosquitoes innate immune system, particularly the upregulation of AMPs, and upregulation of immune pathway leading to reduced viral infection and viral titters. Furthermore, microbial competition between symbiont and viruses in the gut creates a hostile environment for viral replication. Proteus sp. and Paenibacillus sp. were introduced through blood meals and significantly reduced DENV infection and viral titters in aseptic mosquitoes. Notably, sugar meal supplementation with Proteus spp. also decreased DENV infection rates (Ramirez et al., 2012).

Similarly, Elizabethkingia anopheles aegypti colonize Ae, albopictus resulted in lower average ZIKV infections and reduced viral loads in Vero cell assays for ZIKV, DENV, or CHIKV (Onyango et al., 2021). Lysinibacillus spp., previously recognized for its larvicidal activity, was recently shown to reduce ZIKV viral loads in the head and thorax of Ae. aegypti, with no detectable virus in the saliva following forced feeding (Do Nascimento et al., 2022; see Table 1).

Table 1

Table 1. Summaries of the role of mosquitoes’ gut microbiota in pathogen transmission prevention.

3.5 Role of mosquito’s guts microbiota in vector control

Eight articles addressed the use of mosquito microbiota in strategies for vector control. Applications of microbial-based approaches suppress the Aedes mosquito population. For example, an independent evaluation of Wolbachia-infected male (WIM) mosquito releases in Harris county, Texas, showed that CI induced by Wolbachia significantly reduced Ae. aegypti populations by over 90%. Similarly, large-scale field releases of Ae. aegypti mosquitoes infected with the wMel strain of Wolbachia have led to the stable establishment of the bacterium in local mosquito populations, with a consistent prevalence of over 60% (Lozano et al., 2022).

Similarly, large-scale field releases of Ae. aegypti mosquitoes infected with the wMel strain have led to the stable establishment of the bacterium in local mosquito populations, with a consistent prevalence of over 60%. Due to the large-scale establishment of Wolbachia, the incidence of dengue has been reduced (Velez et al., 2023), resulting in a 38% decrease in dengue cases and a 10% reduction in chikungunya cases (Ribeiro Dos Santos et al., 2022).

Other studies have also reported that introgression, which involves crosses between wild Wolbachia-infected Ae. albopictus males (carrying the wild wPip strain) and naturally infected wAlbA/B females lead to complete bidirectional CI, as shown by 0% egg hatch rates. The life history traits in these wild-wPip crosses were similar to those observed in laboratory crosses between lab-wPip males and wild wAlbA/B females (Lejarre et al., 2025). Similarly, the presence of Wolbachia strain wMelM in female Ae. aegypti triggers fitness costs that disrupt egg retention and prevent oviposition (Ross et al., 2025). Introgression of the genetic background from a wild population into a Wolbachia-infected line capable of producing incompatible males (Cholvi et al., 2024).

A pilot study conducted in southern Mexico tested the integration of the Sterile Insect Technique (SIT) and the Incompatible Insect Technique (IIT) using wAlbB-infected Ae. aegypti males. These mosquitoes were mass-reared, irradiated for sterilization, and released in urban areas. After release rates resumed at the five-month mark, the intervention led to an 88.4–89.4% reduction in indoor Ae. aegypti presence and an overall population suppression rate ranging from 50 to 75.2% (Martín-Park et al., 2022). Similarly, combined use of IIT and SIT through the mass release of male Ae. albopictus mosquitoes resulted in a 62% decrease in larval abundance and a 65% decrease in adult populations over the course of a year (Zheng et al., 2019).

Wolbachia-based vector control has shown great promise in reducing arbovirus transmission and mosquito populations. Field releases in endemic areas have significantly decreased disease incidence. However, large-scale, sustainable implementation requires coordinated multidisciplinary collaboration, standardized methodologies, and long-term ecological monitoring to adapt to variable field conditions and maintain success (O’Neill et al., 2019; Nazni et al., 2019).

Beyond Wolbachia-based interventions, some resident bacteria in mosquito influence the physiology of mosquito species; cause mortality, induce the sterility and extent mosquito development. Chromobacterium sp. (Csp_P), Chromobacterium sp. Panama (Csp_P), isolated from field-derived Ae. aegypti showed strong entomopathogenic effects. Larval exposure to Csp_P in breeding water and adult consumption of the bacterium resulted in high mosquito mortality (Ramirez et al., 2014).

A recent study on bacteria and their metabolites isolated from Aedes mosquitoes demonstrated significant larvicidal activity against Ae. aegypti larvae. Among the most promising genera were Bacillus spp., Enterobacter spp. and Stenotrophomonas spp. (De Oliveira et al., 2024). Rajagopal and Ilango (2021) studied the effect of Exiguobacterium spp. (specifically E. aestuarii and E. profundum) on Ae. aegypti larvae. Exposure to different bacterial concentrations significantly prolonged larval development (from 11.41 to 14.78 days) and resulted in reduced fecundity and egg hatchability. Similarly, the Rahnella aquatilis isolate RAeA1, found throughout the tissues of Ae. albopictus was shown to impair female reproductive physiology. Inoculating adult mosquitoes with RAeA1 resulted in disrupted egg production and ovarian development due to reduced levels of ecdysteroids and vitellogenin hormones, which are essential for successful reproduction (Gu et al., 2025).

The gut microbiota of Aedes mosquitoes has been explored for its potential to control arbovirus through Para-transgenesis, which involves the genetic engineering of symbiotic microorganisms to express antipathogen effector molecules. Symbiotic bacterium Serratia AS1 has been genetically engineered to express effector molecules targeting pathogens. Mosquitoes harboring engineered Serratia demonstrated significant inhibition of Plasmodium and ZIKV infections in both Anopheles and Aedes mosquitoes (Hu et al., 2025; Table 2).

Table 2

Table 2. Summaries of role of gut microbiota in vector control.

4 Discussion

In response to the challenges of vector-borne disease and the rapid development of insecticide resistance, integrated mosquito management (IMM) strategies have become increasingly important. IMM advocates for a multifaceted approach that combines chemical, biological, and environmental tools to reduce mosquito populations sustainably. Among biological control methods, bacterial larvicides like Bti and Lysinibacillus sphaericus are widely used (CDC, 2024). These bioinsecticides target larvae specifically, leaving a minimal impact on non-target organisms.

Additionally, the WHO recommends the use of symbiotic bacteria, such as Wolbachia and other microorganisms, to reduce the transmission of arboviral pathogens by interfering with viral replication in mosquito vectors (World Health Organization, 2016). Meanwhile, due to growing scientific interest in targeting the mosquito gut microbiota as a novel approach to control arboviral disease, this emphasized the potential of symbiotic gut bacteria in Aedes mosquitoes as a novel tool for inhibiting pathogen transmission and enhancing vector control.

The gut microbiota also plays a crucial role in mosquito immunity and resistance to pathogens. The presence of bacteria in the midgut can antagonize infectious agents, such as DENV and Plasmodium, acting as a negative factor in the vectorial competence of the mosquito (Onyango et al., 2021; Pocquet et al., 2021). Additionally, gut bacteria are involved in regulating reactive oxygen species (ROS) levels, which are essential for controlling pathogen growth and maintaining mosquito resistance to infections (Cirimotich et al., 2011b). In Anopheles mosquito bacteria like Enterobacter have been shown to enhance ROS production and reduce plasmodium survival in the midgut (Dennison et al., 2016).

The interaction between pathogenic and non-pathogenic microorganism started in the early stage of mosquito development by modulating the basal level of immune gene expression associated with immune response, tissue homeostasis, gut physiology, and metabolism. This microbiota-induced gene expression leads to a more rapid and robust immune response upon pathogen challenge. In Drosophila melanogaster, commensal bacteria upregulate antimicrobial peptide genes via the Imd pathway, enhancing resistance to subsequent infections (Broderick and Lemaitre, 2012). Similarly, in Aedes mosquito symbiotic bacteria’s elevated expression of several immune marker genes, including the Toll pathway related genes and modulating DENV infection (Xi et al., 2008).

Additionally, microbial interactions within the gut microbiome of Aedes mosquitoes are complex and involve mechanisms that enable them to evade mosquito immune responses. For example, the gut microbiome in mosquitoes utilizes C-type lectins (mosGCTLs) to counteract the bactericidal activity of antimicrobial peptides (AMPs) (Pang et al., 2016). This mechanism enables the microbiome to maintain homeostasis and colonize the mosquito’s gut successfully. Similarly, oral ingestion of bacteria triggers a robust immune response, notably antimicrobial peptides, to combat the bacteria (Lhocine et al., 2008). This suggests that interactions between the mosquito immune system and symbiotic bacteria can enhance immune priming, thereby strengthening the mosquito’s immune response against subsequent infections.

Even though the application of symbiotic bacteria for blocking pathogen transmission and suppressing mosquito populations has shown effectiveness under laboratory conditions, its implementation in field settings remains limited. One of the key issues is that bacterial communities are not static; they vary significantly across mosquito species, life stages, environmental conditions, host genome and sex, and dietary regimes (Guégan et al., 2018).

For example, the source of blood meal and mixed blood feeding influence gut bacterial community composition in mosquitoes, potentially affecting pathogen acquisition and transmission (Muturi et al., 2021b). In Anopheles mosquitoes difference in larval diet affects causes a change in the abundance of midgut Enterobacteriaceae influencing the prevalence and intensity of P. berghei in adults (Linenberg et al., 2016). Blood meal increases bacteria’s antioxidant activity by disturbing the compositional harmony of the consortium; this dysbiosis of microbial community may increase mosquito permissiveness for pathogenic infection.

The developmental transition from larvae to adults involves substantial remodeling of the gut and its microbiota. During the larval stage, mosquitoes develop in aquatic environments, where they acquire a diverse range of environmental bacteria. Variations in water temperature, pH, oxygen availability, and other physicochemical properties across different aquatic habitats significantly influence microbial growth and, consequently, shape the larval gut microbiota leads to ecological unpredictability in vector control (Fu et al., 2023).

Furthermore, larval exposure to different bacterial communities can result variation in adult gut microbiota, immune responses, and pathogen transmission (Dickson et al., 2017). Transient microbes present in the larval aquatic environment can be carried over to the adult stage and influence mosquito vector competence. Mosquitoes reared in environmental water containing a diverse microbial community exhibit reduced competence for Zika virus (ZIKV) transmission compared to those reared in laboratory water with limited microbial diversity (Louie and Coffey, 2021).

However, during pupation, the gut undergoes physiological renewal, including the elimination of existing microbial content via the mechanism and the replacement of the larval gut epithelium. Despite this turnover, some bacteria are retained and transmitted transstadial, contributing to the adult microbiota (Fu et al., 2023; Alfano et al., 2019). The instability of microbiota across mosquito development reduces the predictability and reproducibility of microbiota-based vector control strategies in field.

Gut symbionts also present a promising platform for delivering anti-pathogenic effectors through genetic engineering to reduce disease transmission. This method is both cost-effective and scalable, as these engineered symbionts can stably colonize various mosquito vector species and be sustained within mosquito populations through vertical, horizontal, and transstadial transmission, thereby minimizing the need for repeated reintroduction (Ratcliffe et al., 2022).

Even though genetically engineered symbionts hold great promise for targeting arbovirus and Plasmodium transmission and for suppressing mosquito populations, several challenges must be addressed before this approach can be widely implemented in the field. These challenges include fitness costs and genetic instability, ecological risks, horizontal gene transfer and non-target effects, as well as regulatory, ethical, and social concerns (Ratcliffe et al., 2022). To overcome these obstacles and responsibly release genetically modified mosquitoes, a multidisciplinary risk assessment, strong community engagement, and adaptive management strategies are essential to ensure sustainability and public acceptance.

Beyond pathogen suppression, gut microbiota also influences insecticide resistance. Symbiotic bacteria, such as Serratia oryzae, can enhance resistance to deltamethrin in Ae. albopictus by upregulating metabolic detoxification genes (Wang et al., 2025). This dual role, supporting both detoxification and immune defense, highlights the need to better understand microbial contributions to resistance mechanisms and their implications for control strategies. Additionally, studying the composition and functional mechanisms of the microbial community to insecticide resistance will be crucial for identifying microbial markers that could complement existing vector surveillance tools (Mantzoukas and Eliopoulos, 2020).

5 Conclusion

This review provides an overview of the complex and dynamic relationship between the gut microbiota of Aedes mosquitoes and the transmission of arboviral diseases such as Dengue, Zika, and Chikungunya. It also highlights the urgent need for innovative and sustainable vector control strategies. Different symbiotic bacteria species and strains that are taxonomically affiliated with core phyla, including Proteobacteria, Firmicutes, Actinobacteria, and Bacteroidetes, have been isolated from the gut of Aedes mosquitoes play pivotal roles in modulating vector competence. Understanding the factors shaping mosquito gut microbiota is the main point to developing innovative vector control strategies. Since gut bacteria influence pathogen transmission, manipulating these microbes could reduce vector competence or boost mosquito resistance. Continued research on host-pathogen interactions is vital for advancing next-generation public health vector control tools. Continued research into the mechanisms by which gut microbes interact with both their hosts and pathogens is essential for developing next-generation tools for vector control and public health. In addition to bacterial-based therapies, entomopathogenic fungi like Beauveria bassiana and Metarhizium anisopliae have shown promise in lowering mosquito populations and upsetting the balance of gut microbes, which reduces vector fitness and viral susceptibility. Further research is needed using a biomolecular approach to detect the role of gut microbes, such as viruses and fungi, as well as the mechanisms that inhibit the role of pathogenic microbes, as well as the mechanisms of competition and dominance between germs in the mosquito body, which can be the basis for vector control.

Moreover, symbiotic bacteria like Wolbachia have shown great promise in large-scale vector control by reducing arbovirus transmission and mosquito populations. Field releases of Wolbachia-infected mosquitoes have already led to significant declines in disease incidence in endemic areas. Although the reviewed studies offer compelling insights, translating microbiome-based research into scalable public health interventions requires further multidisciplinary collaboration. There is still a significant knowledge gap regarding the dynamics of microbiota in natural environments, especially when field conditions and ecological diversity are present. Long-term monitoring, evaluation of non-target impacts, and standardized microbiome manipulation techniques are necessary to further this strategy. The effective integration of mosquito gut microbiota into public health practice requires multidisciplinary research to inform interventions and continuous field evaluation within vector control programs.

Further research is needed to elucidate the mechanisms by which microbiota influences pathogen transmission fully and to explore potential applications in mosquito control efforts.

Author contributions

AW: Writing – review & editing, Conceptualization, Investigation, Writing – original draft. AS: Validation, Data curation, Writing – review & editing, Conceptualization. AG: Writing – original draft, Investigation, Visualization. MF: Visualization, Investigation, Writing – original draft. NB: Visualization, Investigation, Writing – original draft. AM: Visualization, Writing – original draft, Investigation. MCu: Visualization, Writing – review & editing, Investigation. GA: Conceptualization, Validation, Data curation, Writing – review & editing. DN: Visualization, Data curation, Writing – review & editing. MCo: Writing – review & editing, Validation, Data curation, Investigation. DY: Data curation, Project administration, Writing – review & editing, Conceptualization. RM: Conceptualization, Writing – review & editing, Project administration, Supervision. GP: Writing – review & editing, Validation, Project administration, Supervision, Conceptualization.

Funding

The author(s) declare that no financial support was received for the research and/or publication of this article.

Conflict of interest

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The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision.

Correction note

This article has been corrected with minor changes. These changes do not impact the scientific content of the article.

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Supplementary material

The Supplementary material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fmicb.2025.1656709/full#supplementary-material

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