Studies on prevalence of Hantavirus in small mammals in Southeast Asia: A systematic review and meta-analysis

Zixiao Guo; Hongxin Pan; Nini Wang; Yang Xiao; Qianwen Zhang; Changchun Ren; Puyu Liu; Qun Wu; Lijun Cai; Yang Cheng; Weixia Li; Dingwei Sun

doi:10.1371/journal.pntd.0014075

Abstract

Background

This study systematically analyzed the prevalence of Hantavirus (HVs) in small mammals across Southeast Asia to evaluate the risks of this zoonotic disease.

Methods

We searched the Web of Science, PubMed, Embase, Scopus, and Cochrane Library for studies published up to 6 February 2025, extracted data from 28 eligible studies.

Results

Meta-analysis revealed a pooled prevalence of HVs was 6.07% (986/11,806) in small mammals in Southeast Asia, with the highest prevalence in Indonesia (17.49%) and Singapore (10.53%). The prevalence was higher in samples detected by Enzyme-Linked Immunosorbent Assay (10.68%) and in rodents (8.07%).

Conclusions

The results of our study highlight the urgency of strengthening surveillance in trade networks with Southeast Asia, particularly in high-risk areas such as Indonesia and Singapore, to mitigate the threat of imported zoonotic diseases.

Author summary

Hantaviruses (HVs) are zoonotic pathogens that can cause severe diseases in humans, such as Hantavirus pulmonary syndrome (HPS) and Hemorrhagic Fever with Renal Syndrome (HFRS). Understanding the prevalence of HVs in small mammals is critical for assessing public health risks, especially in regions with high biodiversity and frequent international trade. In this study, we conducted a systematic review and meta-analysis of 28 studies to estimate the prevalence of HVs in small mammals across Southeast Asia. We found an overall prevalence of 6.07%, with higher rates in Indonesia and Singapore. Certain species, such as Rattus norvegicus showed particularly high infection rates. Detection methods also influenced prevalence estimates, with immunofluorescence assays reporting higher rates than molecular methods. Our findings underscore the need for enhanced surveillance in high-risk areas and trade networks to prevent the cross-border spread of Hantaviruses.

Citation: Guo Z, Pan H, Wang N, Xiao Y, Zhang Q, Ren C, et al. (2026) Studies on prevalence of Hantavirus in small mammals in Southeast Asia: A systematic review and meta-analysis. PLoS Negl Trop Dis 20(3): e0014075. https://doi.org/10.1371/journal.pntd.0014075

Editor: Jonas Klingström, Linköping University: Linkopings universitet, SWEDEN

Received: August 26, 2025; Accepted: February 23, 2026; Published: March 12, 2026

Copyright: © 2026 Guo et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Data Availability: All relevant data are available within the manuscript and its supplementary files.

Funding: This work was supported by the Hainan Province Science and Technology Special Fund (ZDYF2023SHFZ134 to DS; ZDKJ2021035 to DS) (https://kjt.hainan.gov.cn/). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Competing interests: The authors have declared that no competing interests exist.

Introduction

Orthohantaviruses, hereafter referred to as Hantaviruses (HVs), are negative-sense RNA viruses that reside within the subfamily Mammantavirinae of the family Hantaviridae, according to the most recent taxonomy from the International Committee on Taxonomy of Viruses (ICTV; https://ictv.global/) [1]. All human-pathogenic HVs belong to this genus, which includes the etiologic agents responsible for both Hemorrhagic Fever with Renal Syndrome (HFRS) and Hantavirus Pulmonary Syndrome (HPS) [2]. HV is a zoonotic pathogen transmitted by small mammals, which can cause HPS, a severe acute respiratory illness in the Americas primarily. The symptom is generally severe and associated with a mortality higher that 25% [3]. In the United States, the estimated case fatality rate is approximately 35% [4,5].The clinical features of HFRS are mainly fever, varying degrees of bleeding tendency, renal failure, headache, back pain, abdominal pain, and hypotension. Infection results from exposure to aerosols contaminated with HVs [6]. Annually, approximately 150,000–200,000 HFRS cases are reported worldwide, with a mortality as high as 15% [7,8]. HFRS is mainly prevalent in Asia and Europe, with China being the country most severely affected by the disease globally. Studies show that China accounts for more than 90% of all reported HFRS cases worldwide [9]. It is worth noting that since 2000, the incidence of HFRS in China has remained high, consistently ranking first in the world [6].

The primary vectors for HVs are small mammals, such as bats and rodents [10]. HVs have a wide range of hosts, which are large in number and often coexist with domestic animals or humans, thereby posing a significant risk of HVs infection to humans. Research indicates that vectors for HVs transmission include Rattus norvegicus, Apodemus agrarius, Rattus flavipectus, among others [11]. Therefore, monitoring the prevalence of small mammals for HVs is crucial for analyzing, assessing, and warning against HFRS. With global climate change and the deepening of China’s exchanges with the international community in transportation, trade, and tourism, pathogen-carrying organisms and the pathogens they carry can be introduced between international ports via vehicles, containers, goods, mail, etc., increasing the risk of imported diseases and potentially triggering domestic transmission [12]. Furthermore, studies have also postulated that HVs infections introduced through international freight channels via rodents may serve as a primary etiology [2].The frequent detection of rodents on incoming transport in China poses a direct risk of pathogen introduction, as evidenced by reports of HVs detection in these imported animals [13].

Hainan is located in the tropical region of China, which is favorable for the reproduction of small mammals. The rodent density exceeds 11.67% using fluorescence Polymerase Chain Reaction method [1]. According to the China Public Health Science Data Center, the number of HFRS cases in Hainan Province exhibited an increasing trend from 2014 to 2019 (https://www.phsciencedata.cn/Share). With the development of the Free Trade Port, the risk of imported HVs carried by small mammal hosts is higher, as small mammals can enter Hainan via ships, aircraft, etc. Current research is scattered across countries such as Vietnam, Thailand, and Indonesia, with no studies estimating the prevalence of HVs across the entire Southeast Asian region. Therefore, it is essential to identify HVs infection in small mammals and clarify the distribution of the virus in small mammals across Southeast Asia.

To understand the prevalence is critical for implementing effective control measures and facilitate the development of targeted preventive strategies for HVs, we conduct a comprehensive systematic review and meta-analysis to evaluate the prevalence of HVs in small mammals. Specifically, we explored the prevalence of HVs in small mammals across Southeast Asia, analyzing variations by geographic subregions, detection methods, host species, publication years, and sample sizes.

Materials and methods

Protocol and registration

The systematic review and meta-analysis were conducted based on the Preferred Reporting Items for Systematic and Meta-analysis (PRISMA) protocols [14] and prospectively registered on PROSPERO (CRD420251063857).

Search strategy

A computerized search was performed in the Cochrane Library, PubMed, Embase, Web of Science, and Scopus databases for publicly available literature on surveys of HVs in small mammals in Southeast Asia. The search period covered the establishment of the databases up to 6 February 2025. The search terms were (Rodent OR Mouse OR Rats OR Mice OR Cricetid OR Bat) AND (Hantaviruses OR Hantaan virus OR Hemorrhagic fever with Renal Syndrome OR HFRS OR Hantavirus infection OR Hantavirus disease OR Hantavirus detection OR Hantavirus prevalence OR HPS OR Hantavirus Pulmonary Syndrome), with the region restricted to Southeast Asia.

Literature inclusion and exclusion criteria

Inclusion criteria: (1) The study type is an observational study, including Cross-Sectional Studies (CSS); (2) Studies involving small mammals (bats, shrews, and rodents) carrying HVs; (3) Detection results must reflect natural infection (excluding experimental infections); (4) The research location in Southeast Asia, including Vietnam, Laos, Cambodia, Thailand, Myanmar, Malaysia, Brunei, Singapore, Indonesia, East Timor, and the Philippines.

Exclusion criteria: (1) Unable to extract or convert key parameters such as sample size, number of positives, or positive rate; (2) Interventional studies, reviews, case reports, conference abstracts, non-original studies, and duplicated publications.

Literature screening and data extraction

Two reviewers carried out the extraction and recording of data from each chosen study independently. In case of any discrepancies between the reviewers or any ambiguity regarding the suitability of a study, additional reviewers were consulted to resolve the issue. Information was recorded as follows: First author, Year of publication, Country, Research subject type, Detection method, Total sample size, Number of HVs-positive cases, Study design type, and Study quality. Microsoft Excel 2021 was used for data management.

Quality assessment

The quality assessment of included cross-sectional studies was conducted using the cross-sectional study evaluation criteria recommended by the Agency for Healthcare Research and Quality (AHRQ) [15]. Scores of 8–11, 4–7, and 0–3 represented high, fair, and poor quality, respectively. Two independent reviewers conducted the evaluations, and any disagreements resolved through discussion or by a third reviewer.

Statistical analysis

We quantified the prevalence of HVs in small mammals in Southeast Asia through meta-analysis using R4.5.0. Study heterogeneity was evaluated by I² statistics to determine the selected model (random-effects for I² > 50%, otherwise fixed-effects). Publication bias was assessed by funnel plots, Egger’s regression test, Begg’s rank correlation tests [16]. Robustness was verified by sensitivity analysis excluding individual studies sequentially [17]. All effect estimates are reported as 95% confidence intervals (CI) and p < 0.05 defined statistical significance.

Results

Database search

A total of 403 studies were identified from database searches. After removing 122 duplicates, 281 studies were screened. A total of 229 studies were excluded based on titles and abstracts. Full-text assessment was conducted for 47 studies (5 were not retrieved), of which 19 were excluded. Finally, 28 studies were included in the meta-analysis(Fig 1) [2,3,5,7, 18–41].

Download:

Fig 1. Flowchart illustrating the study selection process for the meta-analysis.

https://doi.org/10.1371/journal.pntd.0014075.g001

Study characteristics

Ten studies (36%) were published after the year 2015, and 4 studies (14%) were published before 2000. All selected 28 studies (100%) were cross-sectional studies. Sample sizes varied widely, with the smallest sample reported being 53 individuals surveyed in Malaysia [34] and the largest sample being 1311 individuals surveyed in Vietnam [40]. Overall agreement for the rating of the quality of reporting and methodology between the two assessors was 93% (S1 Table). Twenty-two articles were high quality literature, and 6 articles were fair quality literature (S2 Table).

Meta-analysis

Prevalence of HVs in small mammals in Southeast Asia.

A random-effects model was used for the meta-analysis due to the high degree of heterogeneity in prevalence of HVs in small mammals in Southeast Asian (I² = 97.6%, p < 0.01). The results showed that the prevalence of HVs in small mammals in Southeast Asian was 6.07% (95%CI, 3.80%-9.57%, p < 0.05) (Fig 2).

Download:

Fig 2. Forest map of the prevalence of Hantavirus in small mammals in Southeast Asia.

https://doi.org/10.1371/journal.pntd.0014075.g002

Subgroup analysis of countries.

Firstly, studies spanning multiple countries were disaggregated into individual units for analysis. A total of 35 studies were ultimately included (Table 1 and S1 Fig). The prevalence of HVs in Indonesia, Singapore, Thailand, Cambodia, Malaysia, Vietnam were 17.49% (95%CI, 8.90%-31.51%, p < 0.01), 10.53% (95%CI, 0.56%-70.97%, p < 0.01), 4.36% (95%CI, 2.22%-8.42%, p < 0.01), 5.50% (95%CI, 2.39%-12.16%, p < 0.01), 4.32% (95%CI, 1.06%-15.97%, p < 0.01), 4.25% (95%CI, 2.11%-8.36%, p < 0.01), respectively. Only one study was identified for Laos and the Philippines, and the prevalence in Myanmar was not statistically significant (p > 0.05). Significant differences in prevalence of HVs were observed between countries (p < 0.05) (S1 Fig).

Download:

Table 1. Prevalence of Hantavirus in small mammals in different subgroups analysis.

https://doi.org/10.1371/journal.pntd.0014075.t001

Subgroup analysis of detection methods.

Firstly, studies employing multiple detection methods were disaggregated into individual units for analysis. A total of 30 studies were ultimately included (Table 1 and S2 Fig). The 24 studies were divided into three groups based on the detection methods used: IFA, ELISA, RT-PCR. The remaining six studies that used cannot distinguish methods or other methods were excluded, and a subgroup analysis was then performed. The prevalence for IFA, ELISA, RT-PCR were 9.35% (95%CI, 4.85%-17.26%, p < 0.01), 10.68% (95%CI, 4.05%-25.33%, p < 0.01), and 2.29% (95%CI, 0.87%-5.91%, p < 0.01), respectively, with ELISA showing the highest rate. A marginal p-value (p < 0.05) was reported among different detection methods (S2 Fig).

Subgroup analysis of species.

Firstly, studies encompassing multiple species were disaggregated into individual units for analysis. A total of 33 studies were ultimately included (Table 1 and S3 Fig). The prevalence of HVs in rodents, shrews, bats were 8.41% (95%CI, 5.35%-12.99%, p < 0.05), 3.90% (95%CI, 1.09%-13.05%, p < 0.05), 1.64% (95%CI, 0.41%-3.16%, p > 0.05), respectively. Significant differences in prevalence of HVs were observed between species (p < 0.05) (S3 Fig).

Subgroup analysis of publication years.

Subgroup analysis by publication year included a total of 28 studies (Table 1 and S4 Fig). The prevalence of HVs in 1980–1999, 2000–2009, 2010–2019, and 2020–2025 group were 11.99% (95%CI, 4.64%-27.63%, p < 0.05), 5.08% (95%CI, 2.79%-9.07%, p < 0.05), 3.86% (95%CI, 1.97%-7.45%, p < 0.05), 11.32% (95%CI, 2.07%-43.48%, p < 0.05), respectively. Not significant differences in prevalence of HVs were observed between publication years (p > 0.05) (S4 Fig).

Subgroup analysis of year of Sample size.

Subgroup analysis was divided into three groups based on sample size: small sample group (<100), medium sample group (100–500), and large sample group (>500) (Table 1 and S5 Fig). The prevalence of HVs in the small sample group, medium sample, large group were 9.02% (95%CI, 1.97%-32.87%, p < 0.05), 5.78% (95%CI, 2.96%-10.98%, p < 0.05), 5.00% (95%CI, 2.45%-9.94%, p < 0.05), respectively. Not significant differences in prevalence of HVs were observed between sample sizes (p > 0.05) (S5 Fig).

Sensitivity analysis and publication bias

The findings were reliable as the exclusion of any single study did not greatly change the overall results. Tests for publication bias gave mixed results: Egger’s test suggested possible bias (p < 0.001), but Begg’s test did not (p = 0.8911). The funnel plot was visually asymmetric (S6 Fig), and imply possible bias or differences between studies. The funnel plot generated by the trim-and-fill method estimated five missing studies (Fig 3). The adjusted effect increased by 11% (from -2.7392 to -2.4298) but stayed within an acceptable range after adding these. The corrected effect size remained statistically significant, demonstrating that the findings are robust despite potential existence of publication bias.

Download:

Fig 3. Trim and fill funnel plot of the prevalence of Hantavirus in small mammals in Southeast Asia.

https://doi.org/10.1371/journal.pntd.0014075.g003

Discussion

This meta-analysis indicated that the prevalence of HVs in small mammals in Southeast Asia (6.07%) is marginally higher than that reported in China (2.58–6.39%) [9,42]. The transmission of HVs is primarily related to the population of small animal hosts. And the population is closely associated with the hosts’ reproductive capacity, living environment, and climatic conditions. Southeast Asia is recognized as the origin of rodents, with more that 35 species of rodents in this region [26]. The temperature and humidity is high in this region, it could create favorable conditions for the survival and reproduction of small mammals. It leads to an increase in the population of small mammals, thereby the high prevalence of HVs [43].

According to the results of subgroup analysis by country, significant differences were observed in the prevalence of HVs in small mammals. Indonesia and Singapore have higher prevalence, while other countries have showed lower values. Indonesia has large areas of farmland and a suitable climate, while Singapore is densely populated and generates substantial waste [39]. Both of these factors provide ideal habitats for small mammals and contribute to a higher prevalence of HVs. This elevated prevalence poses a direct international risk, as highlighted by a confirmed case of human HVs infection imported from Indonesia to Germany [44]. The absence of positive cases in the Philippines may be related to factors such as the limited number of studies and small sample size. A geographical distribution heat map analysis showed that the prevalence of HVs in small mammals in low-latitude countries (Singapore, Indonesia) is significantly higher than in other Southeast Asian countries (Fig 4).The warm and humid climate of low-latitude regions facilitates small mammal reproduction, leading to higher population densities and enhanced survival of HVs [43]. Chinese research has found that small mammal populations tend to be larger in low-latitude regions [45]. However, other studies have shown that habitat quality has a greater impact on population size than latitude [46]. We suggest that future studies should explore the mechanism by which latitude affects the prevalence of HVs in small mammals.

Download:

Fig 4. Geographic distribution heatmap of prevalence of Hantavirus in small animals in Southeast Asia.

All geographic data processing and visualization were performed with R software (version 4.5.0), using base map data from Natural Earth (public domain, https://www.naturalearthdata.com/) retrieved via the rnaturalearth package.

https://doi.org/10.1371/journal.pntd.0014075.g004

Both IFA and ELISA detection methods showed higher prevalence than RT-PCR, which is consistent with Obando-Rico et al. [47]. The reason for this result may be that IFA and ELISA detect antigens or antibodies, thereby reflecting prior exposure of the virus and a relative higher prevalence. While RT-PCR detects the current HVs infection in small mammals.

This study found a significant difference in the prevalence of HVs between rodents and shrews, with rates of 8.07% and 3.90%, respectively (p < 0.05), while no statistically significant was observed between the prevalence of bat and rodents, and of bats and shrews (p > 0.05). A total of 10,423 rodents were included in this study, far exceeding the sample size of bats and shrews.

To further explore the species-specificity of HVs transmission, we conducted a more in-depth subgroup analysis. The results indicate that R. norvegicus, Bandicota indica, R. rattus, and Suncus murinus are the primary vectors for HVs transmission in Southeast Asia, with their respective being 14.01% (95%CI, 9.28%–20.61%, p < 0.01), 10.42% (95%CI, 9.28%–20.61%, p < 0.01), 4.01% (95%CI, 9.28%–20.61%, p < 0.01), and 3.90% (95%CI, 9.28%–20.61%, p < 0.05) (S7 Fig). Among these species, R. norvegicus exhibited the highest prevalence of HVs, which can be attributed to its status as a dominant species with a broad distribution spanning from tropical to arctic climates [39,48]. Their high adaptability facilitates the transmission of HVs.

In Southeast Asia, distinct HVs seropositivity patterns were observed across small mammal species: B. indica, R. norvegicus, R. exulans, R. rattus, and S. murinus are widely distributed and show relatively high prevalence of HVs [39]. In contrast, R. argentiventer, R. tiomanicus, and Hipposideros spp. have restricted distributions, and their prevalence of HVs may display regional fluctuations driven by habitat fragmentation and host specificity (S8 Fig). Notably, B. indica, R. rattus, and S. murinus are highly abundant in particular geographic regions. Yet, due to inadequate sampling efforts and constraints in research priorities, these species remain poorly investigated in current literature, potentially masking their actual role in HVs epidemiology.

Notably, the HVs data of bats should not be directly pooled with that of rodents or shrews for comparative analysis, given the substantial disparities in research contexts. Unlike the aforementioned small mammals, for which both molecular (RT-PCR) and serological (IFA/ELISA) data are available, current prevalence of HVs in bats is limited to genome detection by RT-PCR alone, and lacked to serological validation (e.g., antibody detection). The number of studies focusing on bats is significantly smaller than that on rodents, resulting in insufficient research coverage and limited data support. Therefore, the absence of statistical significance in the prevalence of HVs in bats observed in this study more likely underestimated the limitations of current research rather than reflects a true absence of epidemiological risk. The primary reasons for this finding are the insufficient number of studies on bats, coupled with a severe scarcity of serological data [3,33,35,36]. As bats are recognized reservoirs for numerous viruses, their potential role in transmission should not be underestimated. Future research must be strengthened to clarify their specific role in the ecology of HVs.

The marked heterogeneity in prevalence of HVs in different host species underestimates the critical role of host species in HVs transmission dynamics. Future research should prioritize investigations into HVs transmission mechanisms, as well as host ecology and behavior, to inform the development of targeted intervention strategies. The integration of geographic information systems (GIS) and molecular epidemiological approaches may help clarify the relationships between host species distributions and HVs transmission risk.

In this study, we conducted the first meta-analysis of the combined prevalence of HVs in small mammals in Southeast Asia. Heterogeneity testing revealed high heterogeneity (I² = 95.6%), prompting a subgroup analysis to explore the sources of heterogeneity. Subgroup analysis revealed significant differences in prevalence of HVs across countries and species (p < 0.05), while detection methods showed marginal significance (p = 0.051). Although the difference in prevalence of HVs by methods did not reach the traditional statistical significance threshold, the proximity to the critical value of 0.05 may suggest a potential influence of detection methods on the results. The marginal significance may reflect inadequate subgroup sample sizes, compromising statistical power and obscuring actual intergroup differences. The potential influence of detection methods requires further validation through larger sample sizes or standardized experimental designs.

Although this study is the first to conduct a meta-analysis on prevalence of HVs in Southeast Asia, several limitations should be noted: (1) We only searched English studies and missed non-English and grey literature, which may have left out important data. (2) Nine Southeast Asian countries were included in our study, however, some had very few studies or small samples. This uneven coverage could affect the generalizability of our results to the entire region. (3) This meta-analysis revealed significant heterogeneity among the studies. (4) The study detected significant publication bias (p < 0.001), which may affect the estimation of the overall positive rate. (5) Although different detection methods can affect the results of prevalence of HVs, the subgroup analysis by country in this study did not distinguish between these methods. (6) Given the high diversity of HVs subtypes and the complexity of viral strains carried by hosts (e.g., rodents), methods in this study poses certain challenges for the simultaneous differentiation and identification of multiple HVs subtypes—particularly in the overlapping regions of the Indian Ocean periphery and Southeast Asia, where viral strain coexistence is highly prevalent [49].

A significant result of Egger’s test and the asymmetry was observed in the funnel plot suggest the possibility of publication bias in the included literatures. Publication bias typically arises when studies with statistically significant or ‘positive’ findings (e.g., higher prevalence rates) are more likely to be published than those with non-significant or ‘negative’ results. This could imply that our original pooled prevalence estimates of 6.07% may be slightly overestimated. However, the trim-and-fill analysis indicated that any potential overestimation was relatively limited in magnitude, and the overarching conclusion—that HVs are present in small mammals across Southeast Asia—remained robust following adjustment. Nevertheless, readers are advised to interpret the absolute prevalence with this potential limitation in consideration.

To address current knowledge gaps, we recommend: (1) enhanced longitudinal surveillance and molecular epidemiological studies in under-represented regions (e.g., Cambodia, Laos) and key host species, particularly in highly endemic areas; (2) implementation of multicenter longitudinal studies incorporating molecular characterization to improve predictive accuracy and intervention efficacy. Research priorities should emphasize HVs ecology, spatiotemporal distribution patterns, and determinants of host population dynamics to inform evidence-based control strategies.

Supporting information

S1 Fig. Subgroup Forest Map by Country.

https://doi.org/10.1371/journal.pntd.0014075.s001

(TIF)

S2 Fig. Subgroup Forest Plot by Methods.

https://doi.org/10.1371/journal.pntd.0014075.s002

(TIF)

S3 Fig. Subgroup Forest Map by Specie.

https://doi.org/10.1371/journal.pntd.0014075.s003

(TIF)

S4 Fig. Subgroup Forest Map by Publication year.

https://doi.org/10.1371/journal.pntd.0014075.s004

(TIF)

S5 Fig. Subgroup Forest Map by Sample size.

https://doi.org/10.1371/journal.pntd.0014075.s005

(TIF)

S6 Fig. Publication bias funnel plot of the prevalence of Hantavirus carried by small mammals in Southeast Asia.

https://doi.org/10.1371/journal.pntd.0014075.s006

(TIF)

S7 Fig. Subgroup Forest Map by Specie and Genu.

Complete detection data were available for 1,093 rodents, but species-specific sample sizes were not reported according to Griffiths et al. (2022). Species-level prevalence was estimated using overall detection rates and the species distribution among 1,126 rodents (990 Rattus norvegicu, 136 Rattus rattus).

https://doi.org/10.1371/journal.pntd.0014075.s007

(TIF)

S8 Fig. Heat map showing the prevalence of Hantavirus in different small mammals in Southeast Asia.

Bandicota indica (A), Rattus norvegicus (B), Rattus rattus (C), Rattus hosaensis (D), Suncus murinus (E), Rattus exulans (F), Rattus tiomanicus (G), Bandicota savilei (H), Mus caroli (I), Rattus argentiventer (J), Rattus losea (K), Mus cervicolor (L), Rattus tanezumi (M), and Hipposideros bats (N). All geographic data processing and visualization were performed with R software (version 4.5.0), using base map data from Natural Earth (public domain, https://www.naturalearthdata.com/) retrieved via the rnaturalearth and rnaturalearthdata package.

https://doi.org/10.1371/journal.pntd.0014075.s008

(TIF)

S1 Table. Characteristics of articles included in the Meta-analysis. ^a Indirect immunofluorescence assay. ^b Enzyme-Linked immunosorbent assay. ^c Reverse transcription-polymerase chain reaction. ^d Cross-Sectional Study.

https://doi.org/10.1371/journal.pntd.0014075.s009

(DOC)

S2 Table. Literature Quality Evaluation.

https://doi.org/10.1371/journal.pntd.0014075.s010

(DOC)

S1 File. PRISMA 2020 Checklist.

From: Page MJ, McKenzie JE, Bossuyt PM, Boutron I, Hoffmann TC, Mulrow CD, et al. The PRISMA 2020 statement: an updated guideline for reporting systematic reviews. BMJ 2021;372:n71. https://doi.org/10.1136/bmj.n71. This work is licensed under CC BY 4.0. To view a copy of this license, visit https://creativecommons.org/licenses/by/4.0/deed.en.

https://doi.org/10.1371/journal.pntd.0014075.s011

(DOCX)

Acknowledgments

We thank the China Public Health Science Data Center for the support of the Hainan hemorrhagic fever data. In addition, we thank Professor Lu Wang from the Chinese Centre for Disease Control and Prevention for her suggestions on this article.

References

1. Wang Q-W, Tao L, Lu S-Y, Zhu C-Q, Ai L, Luo Y, et al. Genetic and hosts characterization of hantaviruses in port areas in Hainan Province, P. R. China. PLoS One. 2022;17(3):e0264859. pmid:35239751
- View Article
- PubMed/NCBI
- Google Scholar
2. Truong TT, Yoshimatsu K, Araki K, Lee B-H, Nakamura I, Endo R, et al. Molecular epidemiological and serological studies of hantavirus infection in northern Vietnam. J Vet Med Sci. 2009;71(10):1357–63. pmid:19887743
- View Article
- PubMed/NCBI
- Google Scholar
3. Arai S, Aoki K, Sơn NT, Tú VT, Kikuchi F, Kinoshita G, et al. Đakrông virus, a novel mobatvirus (Hantaviridae) harbored by the Stoliczka’s Asian trident bat (Aselliscus stoliczkanus) in Vietnam. Sci Rep. 2019;9(1):10239. pmid:31308502
- View Article
- PubMed/NCBI
- Google Scholar
4. Thorp L, Fullerton L, Whitesell A, Dehority W. Hantavirus Pulmonary Syndrome: 1993-2018. Pediatrics. 2023;151(4):e2022059352.
- View Article
- Google Scholar
5. Blasdell K, Cosson JF, Chaval Y, Herbreteau V, Douangboupha B, Jittapalapong S, et al. Rodent-borne hantaviruses in Cambodia, Lao PDR, and Thailand. Ecohealth. 2011;8(4):432–43. pmid:22124701
- View Article
- PubMed/NCBI
- Google Scholar
6. Sehgal A, Mehta S, Sahay K, Martynova E, Rizvanov A, Baranwal M. Hemorrhagic Fever with Renal Syndrome in Asia: History, Pathogenesis, Diagnosis, Treatment, and Prevention. Viruses. 2023;15(2):561.
- View Article
- Google Scholar
7. Johansson P, Yap G, Low H-T, Siew C-C, Kek R, Ng L-C, et al. Molecular characterization of two hantavirus strains from different rattus species in Singapore. Virol J. 2010;7:15. pmid:20096099
- View Article
- PubMed/NCBI
- Google Scholar
8. Wang X, Shen W, Qin Y, Ying L, Li H, Lu J, et al. A case-control study on the risk factors for hemorrhagic fever with renal syndrome. BMC Infect Dis. 2020;20(1):103. pmid:32019494
- View Article
- PubMed/NCBI
- Google Scholar
9. Su F, Liu Y, Ling F, Zhang R, Wang Z, Sun J. Epidemiology of hemorrhagic fever with renal syndrome and host surveillance in Zhejiang province, China, 1990-2021. Viruses. 2024;16(1):145.
- View Article
- Google Scholar
10. Chen Y-M, Hu S-J, Lin X-D, Tian J-H, Lv J-X, Wang M-R, et al. Host traits shape virome composition and virus transmission in wild small mammals. Cell. 2023;186(21):4662–4675.e12. pmid:37734372
- View Article
- PubMed/NCBI
- Google Scholar
11. Jonsson CB, Figueiredo LTM, Vapalahti O. A global perspective on hantavirus ecology, epidemiology, and disease. Clin Microbiol Rev. 2010;23(2):412–41. pmid:20375360
- View Article
- PubMed/NCBI
- Google Scholar
12. Findlater A, Bogoch II. Human Mobility and the Global Spread of Infectious Diseases: A Focus on Air Travel. Trends Parasitol. 2018;34(9):772–83. pmid:30049602
- View Article
- PubMed/NCBI
- Google Scholar
13. Song M, Wang B, Liu J, Gratz N. Insect vectors and rodents arriving in China aboard international transport. J Travel Med. 2003;10(4):241–4. pmid:12946302
- View Article
- PubMed/NCBI
- Google Scholar
14. Moher D, Liberati A, Tetzlaff J, Altman DG, PRISMA Group. Preferred reporting items for systematic reviews and meta-analyses: the PRISMA statement. PLoS Med. 2009;6(7):e1000097. pmid:19621072
- View Article
- PubMed/NCBI
- Google Scholar
15. Peng T-R, Lin H-H, Yang L-J, Huang Y-Y, Wu T-W, Chao Y-C. The impact of inflammatory bowel disease on dementia risk: a current systematic review and meta-analysis. Sci Rep. 2025;15(1):12852. pmid:40229330
- View Article
- PubMed/NCBI
- Google Scholar
16. Abate SM, Checkol YA, Mantefardo B. Global prevalence and determinants of mortality among patients with COVID-19: A systematic review and meta-analysis. Ann Med Surg (Lond). 2021;64:102204. pmid:33692899
- View Article
- PubMed/NCBI
- Google Scholar
17. Begg CB, Mazumdar M. Operating characteristics of a rank correlation test for publication bias. Biometrics. 1994;50(4):1088–101. pmid:7786990
- View Article
- PubMed/NCBI
- Google Scholar
18. Elwell MR, Ward GS, Tingpalapong M, LeDuc JW. Serologic evidence of hantaan-like virus in rodents and man in Thailand. Southeast Asian Journal of Tropical Medicine and Public Health. 1985;16(3):349–54.
- View Article
- Google Scholar
19. Tantivanich S, Ayuthaya PI, Usawattanakul W, Imphand P. Hantaanvirus among urban rats from a slum area in Bangkok. The Southeast Asian Journal of Tropical Medicine and Public Health. 1992;23(3):504–9.
- View Article
- Google Scholar
20. Ibrahim IN, Sudomo M, Morita C, Uemura S, Muramatsu Y, Ueno H, et al. Seroepidemiological survey of wild rats for Seoul virus in Indonesia. Jpn J Med Sci Biol. 1996;49(2):69–74. pmid:8806040
- View Article
- PubMed/NCBI
- Google Scholar
21. Praseno P, Suwarso S. Isolation and presumptive serological characterization of hantavirus from wild rat (Bandicota indica). Med J Indones. 1998;7(3):115.
- View Article
- Google Scholar
22. Nitatpattana N, Chauvancy G, Dardaine J, Poblap T, Jumronsawat K, Tangkanakul W, et al. Serological study of hantavirus in the rodent population of Nakhon Pathom and Nakhon Ratchasima Provinces Thailand. Southeast Asian J Trop Med Public Health. 2000;31(2):277–82. pmid:11127326
- View Article
- PubMed/NCBI
- Google Scholar
23. Lam SK, Chua KB, Myshrall T, Devi S, Zainal D, Afifi SA. Serological evidence of hantavirus infections in Malaysia. The Southeast Asian Journal of Tropical Medicine and Public Health. 2001;32(4):809–13.
- View Article
- Google Scholar
24. Nitatpattana N, Henrich T, Palabodeewat S, Tangkanakul W, Poonsuksombat D, Chauvancy G, et al. Hantaan virus antibody prevalence in rodent populations of several provinces of northeastern Thailand. Trop Med Int Health. 2002;7(10):840–5. pmid:12358618
- View Article
- PubMed/NCBI
- Google Scholar
25. Reynes J-M, Soares J-L, Hüe T, Bouloy M, Sun S, Kruy SL, et al. Evidence of the presence of Seoul virus in Cambodia. Microbes Infect. 2003;5(9):769–73. pmid:12850202
- View Article
- PubMed/NCBI
- Google Scholar
26. Hugot J-P, Plyusnina A, Herbreteau V, Nemirov K, Laakkonen J, Lundkvist A, et al. Genetic analysis of Thailand hantavirus in Bandicota indica trapped in Thailand. Virol J. 2006;3:72. pmid:16953877
- View Article
- PubMed/NCBI
- Google Scholar
27. Pattamadilok S, Lee B-H, Kumperasart S, Yoshimatsu K, Okumura M, Nakamura I, et al. Geographical distribution of hantaviruses in Thailand and potential human health significance of Thailand virus. Am J Trop Med Hyg. 2006;75(5):994–1002. pmid:17124002
- View Article
- PubMed/NCBI
- Google Scholar
28. Kosasih H, Ibrahim IN, Wicaksana R, Alisjahbana B, Hoo Y, Yo IH, et al. Evidence of human hantavirus infection and zoonotic investigation of hantavirus prevalence in rodents in western Java, Indonesia. Vector Borne Zoonotic Dis. 2011;11(6):709–13. pmid:21142967
- View Article
- PubMed/NCBI
- Google Scholar
29. Luan VD, Yoshimatsu K, Endo R, Taruishi M, Huong VT, Dat DT, et al. Studies on hantavirus infection in small mammals captured in southern and central highland area of Vietnam. J Vet Med Sci. 2012;74(9):1155–62. pmid:22673720
- View Article
- PubMed/NCBI
- Google Scholar
30. Ibrahim I, Shimizu K, Yoshimatsu K, Yunianto A, Salwati E, Yasuda SP. Epidemiology of hantavirus infection in Thousand Islands regency of Jakarta, Indonesia. Jpn J Infect Dis. 2013;75(8):1003–8.
- View Article
- Google Scholar
31. Koma T, Yoshimatsu K, Yasuda SP, Li T, Amada T, Shimizu K. A survey of rodent-borne pathogens carried by wild Rattus spp. in Northern Vietnam. Epidemiol Infect. 2013;141(9):1876–84.
- View Article
- Google Scholar
32. Van Cuong N, Carrique-Mas J, Vo Be H, An NN, Tue NT, Anh NL, et al. Rodents and risk in the Mekong Delta of Vietnam: seroprevalence of selected zoonotic viruses in rodents and humans. Vector Borne Zoonotic Dis. 2015;15(1):65–72. pmid:25629782
- View Article
- PubMed/NCBI
- Google Scholar
33. Arai S, Taniguchi S, Aoki K, Yoshikawa Y, Kyuwa S, Tanaka-Taya K, et al. Molecular phylogeny of a genetically divergent hantavirus harbored by the Geoffroy’s rousette (Rousettus amplexicaudatus), a frugivorous bat species in the Philippines. Infect Genet Evol. 2016;45:26–32. pmid:27516187
- View Article
- PubMed/NCBI
- Google Scholar
34. Hamdan NES, Ng YL, Lee WB, Tan CS, Khan FAA, Chong YL. Rodent Species Distribution and Hantavirus Seroprevalence in Residential and Forested areas of Sarawak, Malaysia. Trop Life Sci Res. 2017;28(1):151–9. pmid:28228923
- View Article
- PubMed/NCBI
- Google Scholar
35. Arai S, Kikuchi F, Bawm S, Sơn NT, Lin KS, Tú VT, et al. Molecular Phylogeny of Mobatviruses (Hantaviridae) in Myanmar and Vietnam. Viruses. 2019;11(3):228. pmid:30866403
- View Article
- PubMed/NCBI
- Google Scholar
36. Zana B, Kemenesi G, Buzás D, Csorba G, Görföl T, Khan FAA, et al. Molecular Identification of a Novel Hantavirus in Malaysian Bronze Tube-Nosed Bats (Murina aenea). Viruses. 2019;11(10):887. pmid:31546677
- View Article
- PubMed/NCBI
- Google Scholar
37. Kikuchi F, Aoki K, Ohdachi SD, Tsuchiya K, Motokawa M, Jogahara T, et al. Genetic Diversity and Phylogeography of Thottapalayam thottimvirus (Hantaviridae) in Asian House Shrew (Suncus murinus) in Eurasia. Front Cell Infect Microbiol. 2020;10:438. pmid:32974220
- View Article
- PubMed/NCBI
- Google Scholar
38. Kikuchi F, Senoo K, Arai S, Tsuchiya K, Sơn NT, Motokawa M, et al. Rodent-Borne Orthohantaviruses in Vietnam, Madagascar and Japan. Viruses. 2021;13(7):1343. pmid:34372549
- View Article
- PubMed/NCBI
- Google Scholar
39. Griffiths J, Yeo HL, Yap G, Mailepessov D, Johansson P, Low HT, et al. Survey of rodent-borne pathogens in Singapore reveals the circulation of Leptospira spp., Seoul hantavirus, and Rickettsia typhi. Scientific Reports. 2022;12(1):2692.
- View Article
- Google Scholar
40. Susanti HN, Noor SM, Dharmayanti NLPI, Randusari P, Shimizu K, Hirayama K, et al. Serological Surveillance of Zoonotic Pathogens in Rats in Markets in Bogor, Indonesia. Vector Borne Zoonotic Dis. 2022;22(11):568–70. pmid:36322894
- View Article
- PubMed/NCBI
- Google Scholar
41. Miura K, Chambers J, Takahashi N, Nuradji H, Dharmayanti NI, Randusari P. Coinfection with Orthohantavirus and Leptospira spp. in Rats Collected from Markets in Indonesia. Vector-Borne and Zoonotic Diseases. 2025;25(1):43–8.
- View Article
- Google Scholar
42. Sun Q, Liu Y, Han Y, Liu W, Cao X, Li B. Rodent ecology and etiological investigation in China: results from vector biology surveillance - Shandong Province, China, 2012-2022. China CDC Wkly. 2024;6(36):911–7.
- View Article
- Google Scholar
43. Guterres A, de Lemos ERS. Hantaviruses and a neglected environmental determinant. One Health. 2018;5:27–33. pmid:29911161
- View Article
- PubMed/NCBI
- Google Scholar
44. Hofmann J, Weiss S, Kuhns M, Zinke A, Heinsberger H, Kruger DH. Importation of Human Seoul Virus Infection to Germany from Indonesia. Emerg Infect Dis. 2018;24(6):1099–102. pmid:29774860
- View Article
- PubMed/NCBI
- Google Scholar
45. Zhang XY, Wang YH, Wei XM, Li C, Shen YY, Xu YF. Characterization of rodent viromes and epidemiology of key viruses in China. Acta Theriologica Sinica. 2025;:1–13.
- View Article
- Google Scholar
46. Heldstab SA. Latitude, life history and sexual size dimorphism correlate with reproductive seasonality in rodents. Mammal Review. 2021;51(2):256–71.
- View Article
- Google Scholar
47. Obando-Rico CJ, Valencia-Grajales YF, Bonilla-Aldana DK. Prevalence of orthohantavirus in rodents: A systematic review and meta-analysis. Travel Med Infect Dis. 2023;51:102504. pmid:36402291
- View Article
- PubMed/NCBI
- Google Scholar
48. Villarreal JA, Schlegel WM, Prange HD. Thermal environment affects morphological and behavioral development of Rattus norvegicus. Physiol Behav. 2007;91(1):26–35. pmid:17341426
- View Article
- PubMed/NCBI
- Google Scholar
49. Lokupathirage SMW, Muthusinghe DS, Shimizu K, Nishigami K, Noda K, Tsuda Y, et al. Serological Evidence of Thailand Orthohantavirus or Antigenically Related Virus Infection Among Rodents in a Chronic Kidney Disease of Unknown Etiology Endemic Area, Girandurukotte, Sri Lanka. Vector Borne Zoonotic Dis. 2019;19(11):859–66. pmid:31339833
- View Article
- PubMed/NCBI
- Google Scholar

[ref1] 1. Wang Q-W, Tao L, Lu S-Y, Zhu C-Q, Ai L, Luo Y, et al. Genetic and hosts characterization of hantaviruses in port areas in Hainan Province, P. R. China. PLoS One. 2022;17(3):e0264859. pmid:35239751
View Article
PubMed/NCBI
Google Scholar

[2] View Article

[3] PubMed/NCBI

[4] Google Scholar

[ref2] 2. Truong TT, Yoshimatsu K, Araki K, Lee B-H, Nakamura I, Endo R, et al. Molecular epidemiological and serological studies of hantavirus infection in northern Vietnam. J Vet Med Sci. 2009;71(10):1357–63. pmid:19887743
View Article
PubMed/NCBI
Google Scholar

[6] View Article

[7] PubMed/NCBI

[8] Google Scholar

[ref3] 3. Arai S, Aoki K, Sơn NT, Tú VT, Kikuchi F, Kinoshita G, et al. Đakrông virus, a novel mobatvirus (Hantaviridae) harbored by the Stoliczka’s Asian trident bat (Aselliscus stoliczkanus) in Vietnam. Sci Rep. 2019;9(1):10239. pmid:31308502
View Article
PubMed/NCBI
Google Scholar

[10] View Article

[11] PubMed/NCBI

[12] Google Scholar

[ref4] 4. Thorp L, Fullerton L, Whitesell A, Dehority W. Hantavirus Pulmonary Syndrome: 1993-2018. Pediatrics. 2023;151(4):e2022059352.
View Article
Google Scholar

[14] View Article

[15] Google Scholar

[ref5] 5. Blasdell K, Cosson JF, Chaval Y, Herbreteau V, Douangboupha B, Jittapalapong S, et al. Rodent-borne hantaviruses in Cambodia, Lao PDR, and Thailand. Ecohealth. 2011;8(4):432–43. pmid:22124701
View Article
PubMed/NCBI
Google Scholar

[17] View Article

[18] PubMed/NCBI

[19] Google Scholar

[ref6] 6. Sehgal A, Mehta S, Sahay K, Martynova E, Rizvanov A, Baranwal M. Hemorrhagic Fever with Renal Syndrome in Asia: History, Pathogenesis, Diagnosis, Treatment, and Prevention. Viruses. 2023;15(2):561.
View Article
Google Scholar

[21] View Article

[22] Google Scholar

[ref7] 7. Johansson P, Yap G, Low H-T, Siew C-C, Kek R, Ng L-C, et al. Molecular characterization of two hantavirus strains from different rattus species in Singapore. Virol J. 2010;7:15. pmid:20096099
View Article
PubMed/NCBI
Google Scholar

[24] View Article

[25] PubMed/NCBI

[26] Google Scholar

[ref8] 8. Wang X, Shen W, Qin Y, Ying L, Li H, Lu J, et al. A case-control study on the risk factors for hemorrhagic fever with renal syndrome. BMC Infect Dis. 2020;20(1):103. pmid:32019494
View Article
PubMed/NCBI
Google Scholar

[28] View Article

[29] PubMed/NCBI

[30] Google Scholar

[ref9] 9. Su F, Liu Y, Ling F, Zhang R, Wang Z, Sun J. Epidemiology of hemorrhagic fever with renal syndrome and host surveillance in Zhejiang province, China, 1990-2021. Viruses. 2024;16(1):145.
View Article
Google Scholar

[32] View Article

[33] Google Scholar

[ref10] 10. Chen Y-M, Hu S-J, Lin X-D, Tian J-H, Lv J-X, Wang M-R, et al. Host traits shape virome composition and virus transmission in wild small mammals. Cell. 2023;186(21):4662–4675.e12. pmid:37734372
View Article
PubMed/NCBI
Google Scholar

[35] View Article

[36] PubMed/NCBI

[37] Google Scholar

[ref11] 11. Jonsson CB, Figueiredo LTM, Vapalahti O. A global perspective on hantavirus ecology, epidemiology, and disease. Clin Microbiol Rev. 2010;23(2):412–41. pmid:20375360
View Article
PubMed/NCBI
Google Scholar

[39] View Article

[40] PubMed/NCBI

[41] Google Scholar

[ref12] 12. Findlater A, Bogoch II. Human Mobility and the Global Spread of Infectious Diseases: A Focus on Air Travel. Trends Parasitol. 2018;34(9):772–83. pmid:30049602
View Article
PubMed/NCBI
Google Scholar

[43] View Article

[44] PubMed/NCBI

[45] Google Scholar

[ref13] 13. Song M, Wang B, Liu J, Gratz N. Insect vectors and rodents arriving in China aboard international transport. J Travel Med. 2003;10(4):241–4. pmid:12946302
View Article
PubMed/NCBI
Google Scholar

[47] View Article

[48] PubMed/NCBI

[49] Google Scholar

[ref14] 14. Moher D, Liberati A, Tetzlaff J, Altman DG, PRISMA Group. Preferred reporting items for systematic reviews and meta-analyses: the PRISMA statement. PLoS Med. 2009;6(7):e1000097. pmid:19621072
View Article
PubMed/NCBI
Google Scholar

[51] View Article

[52] PubMed/NCBI

[53] Google Scholar

[ref15] 15. Peng T-R, Lin H-H, Yang L-J, Huang Y-Y, Wu T-W, Chao Y-C. The impact of inflammatory bowel disease on dementia risk: a current systematic review and meta-analysis. Sci Rep. 2025;15(1):12852. pmid:40229330
View Article
PubMed/NCBI
Google Scholar

[55] View Article

[56] PubMed/NCBI

[57] Google Scholar

[ref16] 16. Abate SM, Checkol YA, Mantefardo B. Global prevalence and determinants of mortality among patients with COVID-19: A systematic review and meta-analysis. Ann Med Surg (Lond). 2021;64:102204. pmid:33692899
View Article
PubMed/NCBI
Google Scholar

[59] View Article

[60] PubMed/NCBI

[61] Google Scholar

[ref17] 17. Begg CB, Mazumdar M. Operating characteristics of a rank correlation test for publication bias. Biometrics. 1994;50(4):1088–101. pmid:7786990
View Article
PubMed/NCBI
Google Scholar

[63] View Article

[64] PubMed/NCBI

[65] Google Scholar

[ref18] 18. Elwell MR, Ward GS, Tingpalapong M, LeDuc JW. Serologic evidence of hantaan-like virus in rodents and man in Thailand. Southeast Asian Journal of Tropical Medicine and Public Health. 1985;16(3):349–54.
View Article
Google Scholar

[67] View Article

[68] Google Scholar

[ref19] 19. Tantivanich S, Ayuthaya PI, Usawattanakul W, Imphand P. Hantaanvirus among urban rats from a slum area in Bangkok. The Southeast Asian Journal of Tropical Medicine and Public Health. 1992;23(3):504–9.
View Article
Google Scholar

[70] View Article

[71] Google Scholar

[ref20] 20. Ibrahim IN, Sudomo M, Morita C, Uemura S, Muramatsu Y, Ueno H, et al. Seroepidemiological survey of wild rats for Seoul virus in Indonesia. Jpn J Med Sci Biol. 1996;49(2):69–74. pmid:8806040
View Article
PubMed/NCBI
Google Scholar

[73] View Article

[74] PubMed/NCBI

[75] Google Scholar

[ref21] 21. Praseno P, Suwarso S. Isolation and presumptive serological characterization of hantavirus from wild rat (Bandicota indica). Med J Indones. 1998;7(3):115.
View Article
Google Scholar

[77] View Article

[78] Google Scholar

[ref22] 22. Nitatpattana N, Chauvancy G, Dardaine J, Poblap T, Jumronsawat K, Tangkanakul W, et al. Serological study of hantavirus in the rodent population of Nakhon Pathom and Nakhon Ratchasima Provinces Thailand. Southeast Asian J Trop Med Public Health. 2000;31(2):277–82. pmid:11127326
View Article
PubMed/NCBI
Google Scholar

[80] View Article

[81] PubMed/NCBI

[82] Google Scholar

[ref23] 23. Lam SK, Chua KB, Myshrall T, Devi S, Zainal D, Afifi SA. Serological evidence of hantavirus infections in Malaysia. The Southeast Asian Journal of Tropical Medicine and Public Health. 2001;32(4):809–13.
View Article
Google Scholar

[84] View Article

[85] Google Scholar

[ref24] 24. Nitatpattana N, Henrich T, Palabodeewat S, Tangkanakul W, Poonsuksombat D, Chauvancy G, et al. Hantaan virus antibody prevalence in rodent populations of several provinces of northeastern Thailand. Trop Med Int Health. 2002;7(10):840–5. pmid:12358618
View Article
PubMed/NCBI
Google Scholar

[87] View Article

[88] PubMed/NCBI

[89] Google Scholar

[ref25] 25. Reynes J-M, Soares J-L, Hüe T, Bouloy M, Sun S, Kruy SL, et al. Evidence of the presence of Seoul virus in Cambodia. Microbes Infect. 2003;5(9):769–73. pmid:12850202
View Article
PubMed/NCBI
Google Scholar

[91] View Article

[92] PubMed/NCBI

[93] Google Scholar

[ref26] 26. Hugot J-P, Plyusnina A, Herbreteau V, Nemirov K, Laakkonen J, Lundkvist A, et al. Genetic analysis of Thailand hantavirus in Bandicota indica trapped in Thailand. Virol J. 2006;3:72. pmid:16953877
View Article
PubMed/NCBI
Google Scholar

[95] View Article

[96] PubMed/NCBI

[97] Google Scholar

[ref27] 27. Pattamadilok S, Lee B-H, Kumperasart S, Yoshimatsu K, Okumura M, Nakamura I, et al. Geographical distribution of hantaviruses in Thailand and potential human health significance of Thailand virus. Am J Trop Med Hyg. 2006;75(5):994–1002. pmid:17124002
View Article
PubMed/NCBI
Google Scholar

[99] View Article

[100] PubMed/NCBI

[101] Google Scholar

[ref28] 28. Kosasih H, Ibrahim IN, Wicaksana R, Alisjahbana B, Hoo Y, Yo IH, et al. Evidence of human hantavirus infection and zoonotic investigation of hantavirus prevalence in rodents in western Java, Indonesia. Vector Borne Zoonotic Dis. 2011;11(6):709–13. pmid:21142967
View Article
PubMed/NCBI
Google Scholar

[103] View Article

[104] PubMed/NCBI

[105] Google Scholar

[ref29] 29. Luan VD, Yoshimatsu K, Endo R, Taruishi M, Huong VT, Dat DT, et al. Studies on hantavirus infection in small mammals captured in southern and central highland area of Vietnam. J Vet Med Sci. 2012;74(9):1155–62. pmid:22673720
View Article
PubMed/NCBI
Google Scholar

[107] View Article

[108] PubMed/NCBI

[109] Google Scholar

[ref30] 30. Ibrahim I, Shimizu K, Yoshimatsu K, Yunianto A, Salwati E, Yasuda SP. Epidemiology of hantavirus infection in Thousand Islands regency of Jakarta, Indonesia. Jpn J Infect Dis. 2013;75(8):1003–8.
View Article
Google Scholar

[111] View Article

[112] Google Scholar

[ref31] 31. Koma T, Yoshimatsu K, Yasuda SP, Li T, Amada T, Shimizu K. A survey of rodent-borne pathogens carried by wild Rattus spp. in Northern Vietnam. Epidemiol Infect. 2013;141(9):1876–84.
View Article
Google Scholar

[114] View Article

[115] Google Scholar

[ref32] 32. Van Cuong N, Carrique-Mas J, Vo Be H, An NN, Tue NT, Anh NL, et al. Rodents and risk in the Mekong Delta of Vietnam: seroprevalence of selected zoonotic viruses in rodents and humans. Vector Borne Zoonotic Dis. 2015;15(1):65–72. pmid:25629782
View Article
PubMed/NCBI
Google Scholar

[117] View Article

[118] PubMed/NCBI

[119] Google Scholar

[ref33] 33. Arai S, Taniguchi S, Aoki K, Yoshikawa Y, Kyuwa S, Tanaka-Taya K, et al. Molecular phylogeny of a genetically divergent hantavirus harbored by the Geoffroy’s rousette (Rousettus amplexicaudatus), a frugivorous bat species in the Philippines. Infect Genet Evol. 2016;45:26–32. pmid:27516187
View Article
PubMed/NCBI
Google Scholar

[121] View Article

[122] PubMed/NCBI

[123] Google Scholar

[ref34] 34. Hamdan NES, Ng YL, Lee WB, Tan CS, Khan FAA, Chong YL. Rodent Species Distribution and Hantavirus Seroprevalence in Residential and Forested areas of Sarawak, Malaysia. Trop Life Sci Res. 2017;28(1):151–9. pmid:28228923
View Article
PubMed/NCBI
Google Scholar

[125] View Article

[126] PubMed/NCBI

[127] Google Scholar

[ref35] 35. Arai S, Kikuchi F, Bawm S, Sơn NT, Lin KS, Tú VT, et al. Molecular Phylogeny of Mobatviruses (Hantaviridae) in Myanmar and Vietnam. Viruses. 2019;11(3):228. pmid:30866403
View Article
PubMed/NCBI
Google Scholar

[129] View Article

[130] PubMed/NCBI

[131] Google Scholar

[ref36] 36. Zana B, Kemenesi G, Buzás D, Csorba G, Görföl T, Khan FAA, et al. Molecular Identification of a Novel Hantavirus in Malaysian Bronze Tube-Nosed Bats (Murina aenea). Viruses. 2019;11(10):887. pmid:31546677
View Article
PubMed/NCBI
Google Scholar

[133] View Article

[134] PubMed/NCBI

[135] Google Scholar

[ref37] 37. Kikuchi F, Aoki K, Ohdachi SD, Tsuchiya K, Motokawa M, Jogahara T, et al. Genetic Diversity and Phylogeography of Thottapalayam thottimvirus (Hantaviridae) in Asian House Shrew (Suncus murinus) in Eurasia. Front Cell Infect Microbiol. 2020;10:438. pmid:32974220
View Article
PubMed/NCBI
Google Scholar

[137] View Article

[138] PubMed/NCBI

[139] Google Scholar

[ref38] 38. Kikuchi F, Senoo K, Arai S, Tsuchiya K, Sơn NT, Motokawa M, et al. Rodent-Borne Orthohantaviruses in Vietnam, Madagascar and Japan. Viruses. 2021;13(7):1343. pmid:34372549
View Article
PubMed/NCBI
Google Scholar

[141] View Article

[142] PubMed/NCBI

[143] Google Scholar

[ref39] 39. Griffiths J, Yeo HL, Yap G, Mailepessov D, Johansson P, Low HT, et al. Survey of rodent-borne pathogens in Singapore reveals the circulation of Leptospira spp., Seoul hantavirus, and Rickettsia typhi. Scientific Reports. 2022;12(1):2692.
View Article
Google Scholar

[145] View Article

[146] Google Scholar

[ref40] 40. Susanti HN, Noor SM, Dharmayanti NLPI, Randusari P, Shimizu K, Hirayama K, et al. Serological Surveillance of Zoonotic Pathogens in Rats in Markets in Bogor, Indonesia. Vector Borne Zoonotic Dis. 2022;22(11):568–70. pmid:36322894
View Article
PubMed/NCBI
Google Scholar

[148] View Article

[149] PubMed/NCBI

[150] Google Scholar

[ref41] 41. Miura K, Chambers J, Takahashi N, Nuradji H, Dharmayanti NI, Randusari P. Coinfection with Orthohantavirus and Leptospira spp. in Rats Collected from Markets in Indonesia. Vector-Borne and Zoonotic Diseases. 2025;25(1):43–8.
View Article
Google Scholar

[152] View Article

[153] Google Scholar

[ref42] 42. Sun Q, Liu Y, Han Y, Liu W, Cao X, Li B. Rodent ecology and etiological investigation in China: results from vector biology surveillance - Shandong Province, China, 2012-2022. China CDC Wkly. 2024;6(36):911–7.
View Article
Google Scholar

[155] View Article

[156] Google Scholar

[ref43] 43. Guterres A, de Lemos ERS. Hantaviruses and a neglected environmental determinant. One Health. 2018;5:27–33. pmid:29911161
View Article
PubMed/NCBI
Google Scholar

[158] View Article

[159] PubMed/NCBI

[160] Google Scholar

[ref44] 44. Hofmann J, Weiss S, Kuhns M, Zinke A, Heinsberger H, Kruger DH. Importation of Human Seoul Virus Infection to Germany from Indonesia. Emerg Infect Dis. 2018;24(6):1099–102. pmid:29774860
View Article
PubMed/NCBI
Google Scholar

[162] View Article

[163] PubMed/NCBI

[164] Google Scholar

[ref45] 45. Zhang XY, Wang YH, Wei XM, Li C, Shen YY, Xu YF. Characterization of rodent viromes and epidemiology of key viruses in China. Acta Theriologica Sinica. 2025;:1–13.
View Article
Google Scholar

[166] View Article

[167] Google Scholar

[ref46] 46. Heldstab SA. Latitude, life history and sexual size dimorphism correlate with reproductive seasonality in rodents. Mammal Review. 2021;51(2):256–71.
View Article
Google Scholar

[169] View Article

[170] Google Scholar

[ref47] 47. Obando-Rico CJ, Valencia-Grajales YF, Bonilla-Aldana DK. Prevalence of orthohantavirus in rodents: A systematic review and meta-analysis. Travel Med Infect Dis. 2023;51:102504. pmid:36402291
View Article
PubMed/NCBI
Google Scholar

[172] View Article

[173] PubMed/NCBI

[174] Google Scholar

[ref48] 48. Villarreal JA, Schlegel WM, Prange HD. Thermal environment affects morphological and behavioral development of Rattus norvegicus. Physiol Behav. 2007;91(1):26–35. pmid:17341426
View Article
PubMed/NCBI
Google Scholar

[176] View Article

[177] PubMed/NCBI

[178] Google Scholar

[ref49] 49. Lokupathirage SMW, Muthusinghe DS, Shimizu K, Nishigami K, Noda K, Tsuda Y, et al. Serological Evidence of Thailand Orthohantavirus or Antigenically Related Virus Infection Among Rodents in a Chronic Kidney Disease of Unknown Etiology Endemic Area, Girandurukotte, Sri Lanka. Vector Borne Zoonotic Dis. 2019;19(11):859–66. pmid:31339833
View Article
PubMed/NCBI
Google Scholar

[180] View Article

[181] PubMed/NCBI

[182] Google Scholar

Figures

Abstract

Background

Methods

Results

Conclusions

Author summary

Introduction

Materials and methods

Protocol and registration

Search strategy

Literature inclusion and exclusion criteria

Literature screening and data extraction

Quality assessment

Statistical analysis

Results

Database search

Study characteristics

Meta-analysis

Prevalence of HVs in small mammals in Southeast Asia.

Subgroup analysis of countries.

Subgroup analysis of detection methods.

Subgroup analysis of species.

Subgroup analysis of publication years.

Subgroup analysis of year of Sample size.

Sensitivity analysis and publication bias

Discussion

Supporting information

S1 Fig. Subgroup Forest Map by Country.

S2 Fig. Subgroup Forest Plot by Methods.

S3 Fig. Subgroup Forest Map by Specie.

S4 Fig. Subgroup Forest Map by Publication year.

S5 Fig. Subgroup Forest Map by Sample size.

S6 Fig. Publication bias funnel plot of the prevalence of Hantavirus carried by small mammals in Southeast Asia.

S7 Fig. Subgroup Forest Map by Specie and Genu.

S8 Fig. Heat map showing the prevalence of Hantavirus in different small mammals in Southeast Asia.

S1 Table. Characteristics of articles included in the Meta-analysis. a Indirect immunofluorescence assay. b Enzyme-Linked immunosorbent assay. c Reverse transcription-polymerase chain reaction. d Cross-Sectional Study.

S2 Table. Literature Quality Evaluation.

S1 File. PRISMA 2020 Checklist.

Acknowledgments

References

S1 Table. Characteristics of articles included in the Meta-analysis. ^a Indirect immunofluorescence assay. ^b Enzyme-Linked immunosorbent assay. ^c Reverse transcription-polymerase chain reaction. ^d Cross-Sectional Study.