Abstract
The repeated emergence of pandemic viruses underscores the linkages between land-use change and wildlife disease, and urban-adapted wildlife are of special interest due to their close proximity to humans. However, viral diversity within urban-adapted species and their zoonotic potential remain largely unexplored. Here we compiled a dataset of documented records spanning from 1574 to 2023 on red foxes, raccoons, raccoon dogs, masked palm civets, European hedgehogs, European shrews, wild boars and their viruses, covering 116 countries. These urban-adapted mammals host 286 virus species spanning 24 orders and 38 families, 14 of which are potentially high risk for human infection. Raccoon dogs had increased viral positivity in urban habitats compared to raccoons, wild boars and red foxes. Many viruses in urban-adapted species were phylogenetically related to those found in humans, and our data suggest possible viral spillback. These results highlight zoonotic risks associated with urban-adapted species and suggest enhanced surveillance to mitigate future outbreaks.
This is a preview of subscription content, access via your institution
Access options
Access Nature and 54 other Nature Portfolio journals
Get Nature+, our best-value online-access subscription
$32.99 / 30 days
cancel any time
Subscribe to this journal
Receive 12 digital issues and online access to articles
$119.00 per year
only $9.92 per issue
Buy this article
- Purchase on SpringerLink
- Instant access to the full article PDF.
USD 39.95
Prices may be subject to local taxes which are calculated during checkout






Similar content being viewed by others
Data availability
The primary dataset is available on GitHub at https://github.com/Viraldata-host/Virus (ref. 80). The sequences of seven urban-adapted mammal species associated viruses analysed in this study are available in GenBank (https://www.ncbi.nlm.nih.gov/nuccore/) under accession numbers shown in Source Data Fig. 6. All animal silhouette images used in Figs. 1, 2, 4, Extended Data Fig. 4 and Supplementary Figs. 4 and 15 were sourced from the public-domain database Phylopic (https://www.phylopic.org/). These images are made available under either the CC0 1.0 Universal Public Domain Dedication or the Public Domain Mark 1.0, which permits unrestricted use, sharing and adaptation. Phylogenetic tree data for Fig. 6 and Extended Data Figs. 1–3 are available in Source Data Fig. 6 and Source Data Extended Data Figs. 1–3. Host–virus associations were compiled from PubMed, China National Knowledge Infrastructure (CNKI), EID2, VIRION and GenBank, with detailed information available in Source Data Fig. 3. Geographic distributions of seven urban-adapted mammal species and associated viruses were compiled from PubMed, CNKI and GBIF (https://www.gbif.org/), with further details provided in Source Data Fig. 2. All source data needed to fully replicate and evaluate the analyses are provided as Source Data Figs. 2–6 and Source Data Extended Data Figs. 1–4. Source data are provided with this paper.
References
Magle, S. B., Hunt, V. M., Vernon, M. & Crooks, K. R. Urban wildlife research: past, present, and future. Biol. Conserv. 155, 23–32 (2012).
Santini, L. et al. One strategy does not fit all: determinants of urban adaptation in mammals. Ecol. Lett. 22, 365–376 (2019).
Ma, D. et al. Global expansion of human–wildlife overlap in the 21st century. Sci. Adv. 10, eadp7706 (2024).
Jones, K. E. et al. Global trends in emerging infectious diseases. Nature 451, 990–993 (2008).
Olival, K. J. et al. Host and viral traits predict zoonotic spillover from mammals. Nature 546, 646–650 (2017).
Plowright, R. K. et al. Land use-induced spillover: a call to action to safeguard environmental, animal, and human health. Lancet Planet. Health 5, e237–e245 (2021).
Betke, B. A., Gottdenker, N. L., Meyers, L. A. & Becker, D. J. Ecological and evolutionary characteristics of anthropogenic roosting ability in bats of the world. iScience 27, 110369 (2024).
Ecke, F. et al. Population fluctuations and synanthropy explain transmission risk in rodent-borne zoonoses. Nat. Commun. 13, 7532 (2022).
Xiao, X., Newman, C., Buesching, C. D., Macdonald, D. W. & Zhou, Z.-M. Animal sales from Wuhan wet markets immediately prior to the COVID-19 pandemic. Sci. Rep. 11, 11898 (2021).
Zhao, J., Cui, W. & Tian, B. The potential intermediate hosts for SARS-CoV-2. Front. Microbiol. 11, 580137 (2020).
Hassell, J. M., Begon, M., Ward, M. J. & Fèvre, E. M. Urbanization and disease emergence: dynamics at the wildlife–livestock–human interface. Trends Ecol. Evol. 32, 55–67 (2017).
Bradley, C. A. & Altizer, S. Urbanization and the ecology of wildlife diseases. Trends Ecol. Evol. 22, 95–102 (2007).
Blasdell, K. R. et al. Rats and the city: implications of urbanization on zoonotic disease risk in Southeast Asia. Proc. Natl Acad. Sci. USA 119, e2112341119 (2022).
Feng, A. et al. Transmission of SARS-CoV-2 in free-ranging white-tailed deer in the United States. Nat. Commun. 14, 4078 (2023).
Fagre, A. C. et al. Assessing the risk of human-to-wildlife pathogen transmission for conservation and public health. Ecol. Lett. 25, 1534–1549 (2022).
McBride, D. S. et al. Accelerated evolution of SARS-CoV-2 in free-ranging white-tailed deer. Nat. Commun. 14, 5105 (2023).
Rasche, A. et al. Highly diversified shrew hepatitis B viruses corroborate ancient origins and divergent infection patterns of mammalian hepadnaviruses. Proc. Natl Acad. Sci. USA 116, 17007–17012 (2019).
Goethert, H. K., Mather, T. N., Johnson, R. W. & Telford, S. R. Incrimination of shrews as a reservoir for Powassan virus. Commun. Biol. 4, 1319 (2021).
Campbell, S. J. et al. Red fox viromes in urban and rural landscapes. Virus Evol. 6, veaa065 (2020).
Luca, B. et al. Highly pathogenic avian influenza H5N1 virus infections in wild red foxes (Vulpes vulpes) show neurotropism and adaptive virus mutations. Microbiol. Spectr. 11, e02867-22 (2023).
Fisher, A. M. et al. The ecology of viruses in urban rodents with a focus on SARS-CoV-2. Emerg. Microbes Infect. 12, 2217940 (2023).
Schotte, U. et al. Phylogeny and spatiotemporal dynamics of hepatitis E virus infections in wild boar and deer from six areas of Germany during 2013–2017. Transbound. Emerg. Dis. 69, e1992–e2005 (2022).
Chang, W.-S. et al. Metatranscriptomic analysis of virus diversity in urban wild birds with paretic disease. J. Virol. https://doi.org/10.1128/jvi.00606-20 (2020).
Denstedt, E. et al. Detection of African swine fever virus in free-ranging wild boar in Southeast Asia. Transbound. Emerg. Dis. 68, 2669–2675 (2021).
Song, T. et al. First detection and phylogenetic analysis of porcine circovirus type 2 in raccoon dogs. BMC Vet. Res. 15, 107 (2019).
Albery, G. F. et al. Urban-adapted mammal species have more known pathogens. Nat. Ecol. Evol. 6, 794–801 (2022).
Werner, C. S. & Nunn, C. L. Effect of urban habitat use on parasitism in mammals: a meta-analysis. Proc. Biol. Sci. 287, 20200397 (2020).
Chen, Y.-M. et al. Host traits shape virome composition and virus transmission in wild small mammals. Cell 186, 4662–4675.e12 (2023).
Williams, S. H. et al. Viral diversity of house mice in New York City. mBio 9, e01354-17 (2018).
Snow, N. P., Jarzyna, M. A. & VerCauteren, K. C. Interpreting and predicting the spread of invasive wild pigs. J. Appl. Ecol. 54, 2022–2032 (2017).
Jackowiak, M. et al. Colonization of Warsaw by the red fox Vulpes vulpes in the years 1976–2019. Sci. Rep. 11, 13931 (2021).
Fischer, M. L. et al. Assessing and predicting the spread of non-native raccoons in Germany using hunting bag data and dispersal weighted models. Biol. Invasions 18, 57–71 (2016).
Myśliwy, I., Perec-Matysiak, A. & Hildebrand, J. Invasive raccoon (Procyon lotor) and raccoon dog (Nyctereutes procyonoides) as potential reservoirs of tick-borne pathogens: data review from native and introduced areas. Parasit. Vectors 15, 126 (2022).
Rijks, J. M. et al. Highly pathogenic avian influenza A(H5N1) virus in wild red foxes, the Netherlands, 2021. Emerg. Infect. Dis. 27, 2960–2962 (2021).
Zhao, C. et al. Hedgehogs as amplifying hosts of severe fever with thrombocytopenia syndrome virus, China. Emerg. Infect. Dis. 28, 2491 (2022).
Goldberg, A. R. et al. Widespread exposure to SARS-CoV-2 in wildlife communities. Nat. Commun. 15, 6210 (2024).
Pathogens Prioritization: A Scientific Framework for Epidemic and Pandemic Research Preparedness (World Health Organization, 2024).
Pastoret, P. P. & Brochier, B. Epidemiology and control of fox rabies in Europe. Vaccine 17, 1750–1754 (1999).
Jin, M. et al. Norovirus outbreak surveillance, China, 2016–2018. Emerg. Infect. Dis. 26, 437 (2020).
Xue-Jie, Y. et al. Fever with thrombocytopenia associated with a novel bunyavirus in China. N. Engl. J. Med. 364, 1523–1532 (2024).
Jackson, R. T., Webala, P. W., Ogola, J. G., Lunn, T. J. & Forbes, K. M. Roost selection by synanthropic bats in rural Kenya: implications for human–wildlife conflict and zoonotic pathogen spillover. R. Soc. Open Sci. 10, 230578 (2023).
Balasubramaniam, K. N. et al. Impact of joint interactions with humans and social interactions with conspecifics on the risk of zooanthroponotic outbreaks among wildlife populations. Sci. Rep. 12, 11600 (2022).
Becker, D. J., Streicker, D. G. & Altizer, S. Linking anthropogenic resources to wildlife–pathogen dynamics: a review and meta-analysis. Ecol. Lett. 18, 483–495 (2015).
Altizer, S. et al. Food for contagion: synthesis and future directions for studying host–parasite responses to resource shifts in anthropogenic environments. Phil. Trans. R. Soc. B Biol. Sci. 373, 20170102 (2018).
Qi, X. et al. Molecular characterization of highly pathogenic H5N1 avian influenza A viruses isolated from raccoon dogs in China. PLoS ONE 4, e4682 (2009).
Martin, B. E. et al. Feral swine in the United States have been exposed to both avian and swine influenza A viruses. Appl. Environ. Microbiol. 83, e01346-17 (2017).
Schäffr, J. R. et al. Origin of the pandemic 1957 H2 influenza A virus and the persistence of its possible progenitors in the avian reservoir. Virology 194, 781–788 (1993).
Kawaoka, Y., Krauss, S. & Webster, R. G. Avian-to-human transmission of the PB1 gene of influenza A viruses in the 1957 and 1968 pandemics. J. Virol. 63, 4603–4608 (1989).
Bean, W. J. et al. Evolution of the H3 influenza virus hemagglutinin from human and nonhuman hosts. J. Virol. 66, 1129–1138 (1992).
Olival, K. J. et al. Possibility for reverse zoonotic transmission of SARS-CoV-2 to free-ranging wildlife: a case study of bats. PLoS Pathog. 16, e1008758 (2020).
Pickering, B. et al. Divergent SARS-CoV-2 variant emerges in white-tailed deer with deer-to-human transmission. Nat. Microbiol. 7, 2011–2024 (2022).
Murray, M. H. et al. City sicker? A meta-analysis of wildlife health and urbanization. Front. Ecol. Environ. 17, 575–583 (2019).
Wang, Y. et al. Behavioral plasticity of raccoon dogs (Nyctereutes procyonoides) provides new insights for urban wildlife management in metropolis Shanghai, China. Environ. Res. Lett. 19, 104063 (2024).
Süld, K. et al. An invasive vector of zoonotic disease sustained by anthropogenic resources: the raccoon dog in northern Europe. PLoS ONE 9, e96358 (2014).
Plowright, R. K., Becker, D. J., McCallum, H. & Manlove, K. R. Sampling to elucidate the dynamics of infections in reservoir hosts. Phil. Trans. R. Soc. Lond. B Biol. Sci. 374, 20180336 (2019).
Firth, C. et al. Detection of zoonotic pathogens and characterization of novel viruses carried by commensal Rattus norvegicus in New York City. mBio 5, e01933-14 (2014).
Wu, Z. et al. Decoding the RNA viromes in rodent lungs provides new insight into the origin and evolutionary patterns of rodent-borne pathogens in mainland Southeast Asia. Microbiome 9, 18 (2021).
Hause, B. M., Nelson, E. A. & Christopher-Hennings, J. North American big brown bats (Eptesicus fuscus) harbor an exogenous deltaretrovirus. mSphere 5, e00902-20 (2020).
Hause, B. M., Nelson, E. & Christopher-Hennings, J. Eptesicus fuscus orthorubulavirus, a close relative of human parainfluenza virus 4, discovered in a bat in South Dakota. Microbiol. Spectr. 9, e0093021 (2021).
Streicker, D. G., Franka, R., Jackson, F. R. & Rupprecht, C. E. Anthropogenic roost switching and rabies virus dynamics in house-roosting big brown bats. Vector Borne Zoonotic Dis. 13, 498–504 (2013).
Elsmo, E. J. et al. Highly pathogenic avian influenza A(H5N1) virus clade 2.3.4.4b infections in wild terrestrial mammals, United States, 2022. Emerg. Infect. Dis. 29, 2451–2460 (2023).
Malmlov, A., Breck, S., Fry, T. & Duncan, C. Serologic survey for cross-species pathogens in urban coyotes (Canis latrans), Colorado, USA. J. Wildl. Dis. 50, 946–950 (2014).
GBIF Occurrence Download (GBIF.org, 2024); https://www.gbif.org/zh/occurrence/search
Page, M. J. et al. The PRISMA 2020 statement: an updated guideline for reporting systematic reviews. BMJ 372, n71 (2021).
Wardeh, M., Risley, C., McIntyre, M. K., Setzkorn, C. & Baylis, M. Database of host–pathogen and related species interactions, and their global distribution. Sci. Data 2, 150049 (2015).
Carlson, C. J. et al. The Global Virome in One Network (VIRION): an atlas of vertebrate–virus associations. mBio 13, e0298521 (2022).
Gibb, R. et al. Zoonotic host diversity increases in human-dominated ecosystems. Nature 584, 398–402 (2020).
Wood, S. N. Generalized Additive Models: An Introduction with R (Chapman and Hall/CRC, 2017).
Simpson, G. L. Modelling palaeoecological time series using generalised additive models. Front. Ecol. Evol. 6, 149 (2018).
Viechtbauer, W. Conducting meta-analyses in R with the metafor package. J. Stat. Softw. 36, 1–48 (2010).
Sterne, J. A. C. et al. Recommendations for examining and interpreting funnel plot asymmetry in meta-analyses of randomised controlled trials. BMJ 343, d4002 (2011).
Stanley, T. D. & Doucouliagos, H. Meta-regression approximations to reduce publication selection bias. Res. Synth. Methods 5, 60–78 (2014).
Katoh, K. & Standley, D. M. MAFFT Multiple Sequence Alignment Software Version 7: improvements in performance and usability. Mol. Biol. Evol. 30, 772–780 (2013).
Nguyen, L.-T., Schmidt, H. A., von Haeseler, A. & Minh, B. Q. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Mol. Biol. Evol. 32, 268–274 (2015).
Xu, S. et al. Ggtree: a serialized data object for visualization of a phylogenetic tree and annotation data. iMeta 1, e56 (2022).
Wickham, H. ggplot2: Elegant Graphics for Data Analysis (Springer, 2016).
Shannon, P. et al. Cytoscape: a software environment for integrated models of biomolecular interaction networks. Genome Res. 13, 2498–2504 (2003).
Bonacich, P. & Lloyd, P. Eigenvector-like measures of centrality for asymmetric relations. Soc. Networks 23, 191–201 (2001).
Wells, K., Morand, S., Wardeh, M. & Baylis, M. Distinct spread of DNA and RNA viruses among mammals amid prominent role of domestic species. Glob. Ecol. Biogeogr. 29, 470–481 (2020).
Wei, X. et al. An open database of seven urban-adapted mammal species virus surveillance. GitHub https://github.com/Viraldata-host/Virus (2026).
Acknowledgements
This study was supported by the National Natural Science Foundation of China (32470561, Y.X.; 32271605, Z.Y.X.H.) and Taishan Scholars Project (tsqn202306003, Y.X.). D.J.B. was supported by the National Science Foundation (BII 2213854). The funders had no role in the conceptualization and design of the study, data collection, analysis, decision to publish or preparation of this manuscript.
Author information
Authors and Affiliations
Contributions
Y.X. and F.W. conceptualized the initial hypothesis and conceived and designed the study. X.W., H.L., S.L., Y.W., J.L. and L.H. collected the data and conducted the data analyses. X.W., H.L. and D.J.B. performed the statistical analyses. X.W. and S.L. carried out phylogenetic analyses and interpretation. X.W., S.L. and H.L. created and prepared the figures and tables. Y.X., D.J.B., Z.Y.X.H., X.W. and H.L. wrote the first draft of the manuscript. X.W., H.L., Z.Y.X.H., S.L., Y.W., J.L., L.H., Y.L., D.J.B., F.W. and Y.X. contributed substantially to data acquisition, interpretation, and revision and editing of the manuscript.
Corresponding authors
Ethics declarations
Competing interests
The authors declare no competing interests.
Peer review
Peer review information
Nature Microbiology thanks the anonymous reviewers for their contribution to the peer review of this work. Peer reviewer reports are available.
Additional information
Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Extended data
Extended Data Fig. 1 Phylogenetic tree of SFTSV showing potential bidirectional transmission between urban-adapted wildlife and humans.
The trees were midpoint-rooted, with the scale bar representing the number of nucleotide substitutions per site. Only support values > 80% are shown. Virus names include GenBank accession numbers, host species, sampling location, and sampling year.
Extended Data Fig. 2 Additional phylogenetic trees illustrating clustering between viruses from urban-adapted species and human viruses.
The trees were midpoint-rooted, with the scale bar representing the number of nucleotide substitutions per site. Only support values > 80% are shown. Virus names include GenBank accession numbers, host species, sampling location, and sampling year.
Extended Data Fig. 3 Additional phylogenetic trees of viruses from urban-adapted species clustering with human and domestic animal viruses.
The trees were midpoint-rooted, with the scale bar representing the number of nucleotide substitutions per site. Only support values > 80% are shown. Virus names include GenBank accession numbers, host species, sampling location, and sampling year.
Extended Data Fig. 4 Virus transmission among seven urban-adapted mammal species, domestic animals, and humans.
a) The human-associated virus-sharing network. Node size indicates the number of ‘human-associated viruses’ carried by each urban-adapted species. The edge thickness represents the number of shared viruses between two hosts. b) Boxplot showing the number of shared ‘human-associated viruses’ between each urban-adapted species and domestic animals. The sample size for each group is n = 21. c) Boxplot showing the number of shared viruses between each domestic animal and urban-adapted species. The sample size for each group is n = 7. The lollipop chart displays the eigenvector centrality of each domestic animal species, with circle color representing the eigenvector centrality and the y-axis representing the weighted eigenvector centrality. In b and c, the boxes represent the interquartile range (IQR), which spans from the first to the third quartiles. The lines outside the boxes represent values within 1.5 times the IQR. The horizontal line within each box marks the median, while the rhombus indicates the mean.
Supplementary information
Supplementary Information (download PDF )
Supplementary Discussion and Figs. 1–19.
Supplementary Tables (download XLSX )
Supplementary Tables 1–8.
Source data
41564_2026_2311_MOESM5_ESM.zip (download ZIP )
Source Data Fig. 2 Statistical source data. Source Data Fig. 3 Statistical source data. Source Data Fig. 4 Statistical source data. Source Data Fig. 5 Statistical source data. Source Data Fig. 6 Unprocessed phylogenetic tree. Source Data Extended Data Fig. 1 Unprocessed phylogenetic tree. Source Data Extended Data Fig. 2 Unprocessed phylogenetic tree. Source Data Extended Data Fig. 3 Unprocessed phylogenetic tree. Source Data Extended Data Fig. 4 Statistical source data.
Rights and permissions
Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.
About this article
Cite this article
Wei, X., Li, H., Huang, Z.Y.X. et al. A global-scale assessment of zoonotic virus diversity and spillover potential in urban-adapted mammal species. Nat Microbiol (2026). https://doi.org/10.1038/s41564-026-02311-9
Received:
Accepted:
Published:
Version of record:
DOI: https://doi.org/10.1038/s41564-026-02311-9


