ABSTRACT Proposed mechanisms of zoonotic virus spillover often posit that wildlife transmission and amplification precede human outbreaks. Between 2006 and 2012, the palm _Raphia

farinifera_, a rich source of dietary minerals for wildlife, was nearly extirpated from Budongo Forest, Uganda. Since then, chimpanzees, black-and-white colobus, and red duiker were observed

feeding on bat guano, a behavior not previously observed. Here we show that guano consumption may be a response to dietary mineral scarcity and may expose wildlife to bat-borne viruses.

Videos from 2017–2019 recorded 839 instances of guano consumption by the aforementioned species. Nutritional analysis of the guano revealed high concentrations of sodium, potassium,

HUMAN-GREAT APE ENTERIC EUKARYOTIC VIROMES IN CENTRAL AFRICAN FOREST THAN IN A EUROPEAN ZOO: A ONE HEALTH ANALYSIS Article Open access 21 June 2023 LAND-USE CHANGE AND THE LIVESTOCK

REVOLUTION INCREASE THE RISK OF ZOONOTIC CORONAVIRUS TRANSMISSION FROM RHINOLOPHID BATS Article 31 May 2021 PATHOGENS AND PLANETARY CHANGE Article 15 January 2025 INTRODUCTION Spillover of

viruses from wildlife to humans is often thought to be preceded by viral transmission and amplification among wildlife. For example, human ebolavirus outbreaks in Africa follow sylvatic

transmission cycles in non-human primates and ungulates, with humans likely becoming infected through contact with carcasses1,2,3. Similarly, epidemiological data and analyses of inferred

viral genomic recombination suggest that approximately half of human-infecting coronaviruses underwent transmission from wildlife reservoirs to humans through intermediary hosts4,5. Despite

the high social and economic costs of zoonoses6, the mechanisms underlying such antecedent virus transmission within animals remain poorly understood. Budongo Forest Reserve, western Uganda,

contains approximately 482 km2 of medium-altitude, semi-deciduous forest7 and is located in the Albertine Rift, a region of exceptional biodiversity and endemism8. Until approximately 2008,

the swamp forests of Budongo contained _Raphia farinifera_, a palm that, when decaying, provided a high-quality source of essential dietary minerals to wildlife9. Between 2006 and 2012,

tobacco farming increased markedly in the area due to rising international demand and incentives from tobacco companies with longstanding operations in Uganda10. As a result, local farmers

nearly extirpated _R. farinifera_ because of its usefulness for making strings on which to dry tobacco leaves9,11. Budongo’s eastern chimpanzees (_Pan troglodytes schweinfurthii_) altered

their feeding behavior in response to this loss of a primary source of dietary minerals, increasingly consuming alternative sources such as clay, termite mounds, and the decaying pith of

other tree species9. In 2017, we observed a never-before documented behavior by several species of wildlife in Budongo, including chimpanzees: the consumption of bat guano. Here we present

the results of an investigation as to whether this behavior could be an adaptation to dietary mineral scarcity, analogous to what has been documented for similar behaviors at this site9. We

also investigate whether guano consumption could be an ecological mechanism whereby wildlife such as chimpanzees might be exposed to bat-borne viruses. We document high frequencies of guano

consumption by three species of wildlife in Budongo, high concentrations of essential dietary minerals in the guano, and diverse bat-borne viruses in the guano, including a novel

betacoronavirus within the _Hibecovirus_ subgenus. These results illustrate how remote upstream forces can induce unanticipated causal chains that alter wildlife ecology and behavior, one

result of which may be to increase virus spillover risk. RESULTS FIELD STUDIES On June 25, 2017, we first observed chimpanzees consuming bat guano from under a large, hollow tree

(_Mildbraediodendron excelsum_) in which a colony of Noack’s roundleaf bat (_Hipposideros ruber_) was roosting (Fig. 1). Using a trail camera, we captured video images of chimpanzees,

black-and-white colobus (_Colobus guereza occidentalis_) and red duiker (_Cephalophus natalensis_) repeatedly consuming guano from beneath the tree (Fig. 1). Animals consumed the guano

directly, and not incidentally (e.g. from consumption of adjacent contaminated clay and water), as evidenced by the clearly visible selection of the guano itself during all instances and the

presence of excavations and characteristic hand prints in the guano after animals had left (Supplementary Fig. 1a). We recorded 92 separate instances of guano consumption by chimpanzees on

71 different days, with between 1 and 13 chimpanzees per instance. Chimpanzees removed and ate guano with their hands (Fig. 1b, Supplementary Videos 1 and 2), or they drank adjacent water

using a leaf sponge (folded leaves used to collect water12; Supplementary Video 3). Cameras captured black-and-white colobus feeding on guano on 65 occasions during 56 different days, with

between 1 and 9 individuals per instance. These primates ate guano directly (Fig. 1c; Supplementary Video 4). Cameras captured solitary red duikers on 682 occasions on 210 different days.

Duikers either licked guano directly or drank adjacent water next to the pile (Fig. 1d; Supplementary Video 5). On one occasion, we observed a ~ 2 m human-modified pole, suggesting that

local people had also visited this tree, perhaps to collect guano (Supplementary Fig. 1b). DIETARY MINERAL ANALYSES Nutritional analyses revealed that the guano contained concentrations of

magnesium, phosphorus and potassium higher than in any other recorded dietary source of minerals at Budongo (Table 1). The guano also contained concentrations of sodium approximately equal

to that of decaying _Cleistopholis patens_ wood, the primary alternative source of dietary sodium for chimpanzees subsequent to the loss of _R. farinifera_9 (Table 1). Concentrations of

calcium, manganese and iron were within ranges of other sources at Budongo (Table 1). VIRUS IDENTIFICATION AND CHARACTERIZATION Metagenomic analyses of the bat guano revealed 27 novel

putative eukaryotic viruses with 30.2–92.7% amino acid identity to known viruses of 12 families (_Coronaviridae_, _Dicistroviridae_, _Hepeviridae_, _Iflaviridae_, _Nodaviridae_,

_Parvoviridae_, _Picobirnaviridae_, _Picornaviridae_, _Permutotetraviridae_, _Polycipiviridae_, _Reoviridae_ and _Totiviridae_) and to 7 currently unclassified viruses (Supplementary Table 

1). Individual guano samples analyzed contained an average of 14.5 viruses (standard deviation 3.6) that varied in prevalence from 9 to 100% among samples and in abundance over approximately

4 orders of magnitude, with arthropod-infecting viruses generally most prevalent and most abundant, consistent with the insectivorous diet of _H. ruber_ (Supplementary Fig. 2). Sequences

corresponding to a novel betacoronavirus (_Coronaviridae_: _Betacoronavirus_) were present in 6 samples (55%) (Supplementary Fig. 2). Due to the public health significance of

betacoronaviruses, we intensively sequenced this virus, Buhirugu virus 1 (BHRGV-1), using the sample with the most abundant reads (sample 9 in Supplementary Fig. 2) and succeeded in

obtaining 15,433 bases of the orf1a/b polyprotein gene and 3,181 bases of the spike protein gene (GenBank OP199247). Phylogenetic analyses (Fig. 2) show BHRGV-1 to be a novel member of the

subgenus _Hibecovirus_, approximately equidistant from Bat Hp-betacoronavirus and Zaria bat coronavirus13,14. BHRGV-1 and the other hibecoviruses form a well-supported clade most closely

related to viruses of the separate subgenus _Sarbecovirus_, which contains SARS-CoV and SARS-CoV-2 (Fig. 2)15. To investigate the potential host range of BHRGV-1, we conducted predicted

protein structure and in silico docking analysis of the BHRGV-1 spike protein (Supplementary Fig. 3) and the angiotensin II (ACE2) receptors of humans and the other mammals observed

consuming bat guano (Supplementary Fig. 4). In cases where the ACE2 nucleotide sequence of a particular animal was not available, we used sequences from a closely related species. The

Ramachandran scores for BGHRV-1 S and the various species of ACE2 within the energetically favored region of the protein ranged from 95 to 98% (Supplementary Table 3). Docking analyses of

the BHRGV-1 spike protein indicate non-permissive binding interactions between BHRGV-1 S and ACE2 receptors in all sequences analyzed (Supplementary Table 4), implying that ACE2 may not be

the _Hibecovirus_ receptor. To investigate other potential receptors, we repeated this analysis on Aminopeptidase-N, Dipeptidyl peptidase 4, and CEACAM116, with similar results indicating

non-permissive binding interactions between BHRGV-1 S and each of these molecules (Supplementary Table 4, Supplementary Figs. 5–7 and Supplementary Tables 2–4). Finally, we examined whether

BHRGV-1 has a predicted hemagglutinin-esterase region, which would implicate use of O-acetylated sialic acids as a receptor17, but we found no evidence of such a region by scanning for

matches in the InterPro protein signature databases18,19. DISCUSSION Minerals are essential for physiological functioning, growth, reproduction and immunity20. Minerals are also often

limiting in the core diets of wild animals21. Some cave-dwelling invertebrates, fish and salamanders consume bat guano to obtain minerals in their nutrient-limited subterranean

environments22. However, to our knowledge, guano ingestion by forest-dwelling mammals has not previously been reported. Bat guano also contains nutrients critical to plant growth, such as

nitrogen, phosphate, and potassium, making guano an efficient and widely used fertilizer23. This may explain why people appear to have visited the same tree where we documented guano

consumption by wildlife. We note that another betacoronavirus has been described in bat guano collected as fertilizer in Thailand24, and that harvesting bat guano for this purpose is a

widespread but underappreciated practice that may increase pandemic risk25. Our results suggest that guano consumption by Budongo wildlife may be a behavioral adaptation to mineral scarcity.

This inference is supported by a decades-long body of evidence showing that wildlife in Budono have responded to the disappearance of _R. farinifera_ by seeking alternative mineral

sources9,26,27. The guano contained concentrations of potassium, magnesium, sodium and phosphorus equal to or in excess of concentrations in other dietary sources. Past studies have shown

that consumption of alternative sources of minerals by Budongo chimpanzees began with the disappearance of _R. farinifera_9,26,27. Black-and-white colobus and duiker have not been as

intensively studied in Budongo, so it is unknown whether guano consumption is also a new behavior for these animals. Black-and-white colobus frequently consume soil, clay, aquatic plants and

even cement, demonstrating extreme dietary plasticity with respect to mineral acquisition28. Guano consumption also appears to expose wildlife to bat-associated viruses, to the extent that

the sequences we obtained represent infectious viruses. BHRGV-1 is a member of the subgenus _Hibecovirus_, which contains viruses that primarily infect bats of the genus _Hipposideros_ but

have also been documented in the bat genera _Macronycteris_, _Nycteris_, and _Rhinolophus_29, and _Hibecovirus_ is sister taxon to the subgenus _Sarbecovirus_, which contains SARS-CoV and

SARS-CoV-2. Predicted protein structure analysis of the BHRGV-1 S protein indicates the highest structural similarity to the SARS-CoV S protein (Supplementary Fig. 3 and Supplementary Table 

2). However, the binding affinity of the BHRGV-1 S protein for all ACE2 proteins examined (including those of _H. armiger_, a close relative of _H. ruber_) is low (Supplementary Fig. 4 and

Supplementary Tables 3 and 4), indicating that ACE2 is likely not a viable receptor for host cell entry in these mammals. We obtained similar results for the putative alternative receptors

Aminopeptidase-N, Dipeptidyl peptidase 4, and CEACAM116 (Supplementary Figs. 5–7 and Supplementary Tables 3 and 4), and we found no evidence that BHRGV-1 has a predicted

hemagglutinin-esterase region that might bind O-acetylated sialic acids, as has been shown for other coronaviruses17. If none of these molecules are, in fact, receptors for BHRGV-1 and other

hibecoviruses, this would merit further study, especially for predicting the host range and zoonotic potential of coronaviruses in subgenera other than _Sarbecovirus_. Coronavirus

infections of wildlife have not been documented in Budongo to date. Examining local wildlife for evidence of BHRGV-1 or similar viruses in feces could yield additional information about the

breadth of species that might have been exposed to bat-borne coronaviruses. However, multiple outbreaks of respiratory disease in the chimpanzees of Budongo have been observed, with the

causes remaining undiagnosed. Respiratory disease outbreaks in other chimpanzee populations in Uganda have resulted from cross-species transmission of viruses from humans30,31, and human

betacoronavirus OC43 can infect wild chimpanzees and cause clinical disease32. Intriguingly, chimpanzees, black-and-white colobus, and red duiker have all been implicated in ebolavirus

outbreaks in Central and West Africa1,33,34. The natural history of the ebolaviruses is poorly understood, but multi-host models of sylvatic ebolavirus transmission posit that outbreaks

occur when primates and ungulates become infected by bats and serve as amplifying hosts2,3,35. Similarly, many bat-borne coronaviruses have emerged in humans after transmission through

intermediary hosts4,5. Mechanisms of virus transmission from bats to other wildlife in nature remain poorly understood, although consumption of fruit contaminated by bats35,36 and contact

with viruses shed into the environment37,38 have been hypothesized. Our data suggest another plausible ecological mechanism for exposure of wildlife to bat-associated viruses: consumption of

bat guano as a source of dietary minerals. Tropical forest plants and soils are mineral-poor39. Depletion of primary sources of minerals such as _R. farinifera_ could create conditions that

favor guano consumption as a “fallback” mineral source. Infectious disease emergence is often attributed to drivers such as land conversion, hunting, urbanization, climate change, and

agricultural intensification40, but the ecological mechanisms whereby these drivers lead to cross-species pathogen transmission remain imprecisely understood. Our results provide an

illustration of how these mechanisms might follow elaborate causal chains. In Budongo, international demand for tobacco caused local selective deforestation and loss of a primary source of

dietary minerals, which led to fallback consumption of guano by wildlife and exposure of wildlife to bat-associated viruses, including a congeneric of the pandemic SARS coronaviruses.

Mathematical tools for representing causal chains and causal networks are becoming widespread in epidemiology41 and might prove useful for assessing how environmental and social drivers

ultimately lead to zoonotic transmission. This understanding, in turn, could lead to improved precision in the application of tools for preventing pandemics. For example, compared to the

costs of a pandemic, the costs of offering local farmers substitutes to _R. farinifera_ for making strings to dry tobacco leaves would likely be trivial6,11. In general, we suggest that

understanding causal chains and identifying their “breakable” links holds promise for illuminating disease ecology and improving zoonoses prevention. METHODS FIELD STUDIES The study took

place in the Budongo Forest Reserve, Uganda. We first observed chimpanzees in the habituated Waibira community42,43 feeding on bat guano in a hollow tree on the 25th of June, 2017, even

though chimpanzees had regularly been observed since 2011. On the 5th of July 2017, we installed a trail camera (Bushnell Trophy Cam, model 119,774) with the following default settings: 10 s

video length and interval, auto sensor level, low night vision shutter, and 24 h camera mode. We mounted the camera to a tree 6 m from the tree atop the guano at a height of 1 m. Image

capture occurred between the 6th of July and the 18th of October 2017 and again between the 14th of September 2018 and the 28th of April 2019. We analyzed the resulting 14,567 10 s video

recordings (40.46 camera-hours in total) for the presence and number of animals feeding on the guano. We defined an “instance” as a set of sequential video recordings of animals of a given

species feeding on the guano, separated from the previous instance by at least 30 min during which no individuals of the same species were recorded. We collected guano samples from this

hollow tree monthly from 13th September 2018 to 29th April 2019 (Supplementary Fig. 2) and divided them for mineral content analysis and virus identification. We oven-dried samples for

mineral content analysis (approximately 50 g) and stored and shipped them to the USA at ambient temperature. We placed samples for molecular analysis (approximately 0.9 ml) in 1.8 ml sterile

cryovials, mixed them thoroughly with an equal volume of RNAlater Stabilization Solution (Thermo Fisher, Waltham, MA, USA), and stored them cold in the field and in liquid nitrogen prior to

and during shipment to the USA. All animal use was strictly non-invasive and observational. The study protocol was reviewed and approved by the Uganda Wildlife Authority and the Uganda

National Council for Science and Technology, and was in compliance with the guidelines of the Animal Welfare and Ethical Review Body of the University of Stirling and all applicable

regulations governing the protection of animals and research. We have complied with all relevant ethical regulations for animal use. The species of animals were _Pan troglodytes

schweinfurthii_, _Hipposideros ruber_, _Colobus guereza occidentalis_, and _Cephalophus natalensis_, and all were wild-type and of undetermined sex and age. DIETARY MINERAL ANALYSES Prior to

analysis, we inactivated samples with ultraviolet radiation and oven-dried them at 105 °C for 48 h. We then digested samples using the MARS 6 Microwave Digestion System (CEM Corporation,

Matthews, NC, USA) and analyzed them for Ca, P, Mg, K, Na, Fe, Zn, Cu, Mn, and Mo on an iCAP 6300 inductively coupled plasma radial spectrometer (Thermo Fisher, Waltham, MA, USA)44.

METAGENOMIC ANALYSES We processed guano samples for virus identification using metagenomic methods31,45. Briefly, we added 200 μl of guano+RNAlater to 800 μl of Hanks’ balanced salt solution

and homogenized them in PowerBead Tubes (Qiagen, Hilden, Germany) containing 2.38 mm metal beads. We then treated the homogenate with nucleases to reduce unencapsidated nucleic acids46. We

used the QIAmp MinElute Virus Spin Kit (Qiagen, Hilden, Germany) to isolate total nucleic acids, and we and converted RNA to double-stranded cDNA using the SuperScript double-stranded cDNA

Synthesis Kit (Invitrogen, Carlsbad, CA, USA). We cleaned cDNA using Agencourt AmpureXP beads (Beckman Coulter, Brea, CA, USA) and synthesized DNA libraries using the Nextera XT DNA sample

preparation kit (Illumina, San Diego, CA, USA). We sequenced libraries on an Illumina MiSeq instrument using 600 cycle v3 MiSeq Reagent Kits. We trimmed resulting sequences at a Phred

quality score <30, discarded reads <50 bp, and removed sequences matching host genomes and known contaminants. We thereby sequenced samples to a mean depth after quality and length

trimming and host genome subtraction of 1.9 M reads (standard error 0.2 M reads), ranging from 1.4 to 3.5 M reads per sample. DATA PROCESSING, BIOINFORMATICS, STATISTICS, AND REPRODUCIBILITY

We subjected sequence reads to de novo assembly using SPAdes 3.13.047, discarded resulting contiguous sequences (contigs) <500 nucleotides, and used cd-hit48 to remove redundant contigs

(90% similarity threshold). We compared remaining contigs to custom databases of representative virus protein sequences and to the NCBI non-redundant protein sequence database using

blastx49. We ran and analyzed blank samples in parallel to ensure that cross contamination had not occurred. To investigate BHRGV-1 in greater detail, we queried the initial 15,433 bp contig

representing this virus against the National Center for Biotechnology Information (NCBI) nucleotide database using blastn in BLAST + 50,51. Based on this analysis, we chose two reference

sequences for downstream comparisons: bat Hp-betacoronavirus/Zhejiang2013 (NCBI accession ID NC_025217.1) with 74.5% nucleotide sequence identity to BHRGV-1 and Zaria bat coronavirus strain

ZBCoV (Genbank: HQ166910.1) with 76.4% nucleotide sequence identity to BHRGV-1. We then mapped sequence reads from the sample containing the highest concentration of this virus (sample 9,

Fig. S2) against each reference separately using CLC Genomics Workbench (Qiagen, Hilden, Germany), specifying a minimum length fraction of 0.5 and a minimum similarity fraction of 0.8. We

extended mapped regions in CLC Genomics Workbench using iterative mapping of previously unmapped reads to consensus sequences extracted from each prior iteration. We collected reads thus

identified, assembled them de novo, and aligned the resulting contigs to both references to create a genome scaffold (each reference was useful for assembling different regions of the

BHRGV-1 genome). We then mapped contigs and reads that did not form contigs against the genome scaffold to create a final draft 32,594 bp sequence of the BHRGV-1 ORF1ab polyprotein and spike

protein (S) genes. We constructed phylogenetic trees of the BHRGV-1 ORF1ab polyprotein and spike protein (S) genes using PhyML52 with smart model selection53 (GTR + I models selected in

both cases) and 1000 bootstrap replicates to assess statistical confidence in clades. We modeled 3D-structures of the receptor binding domain (RBD) of BHRGV-1 S and the putative receptor

proteins ACE2, Aminopeptidase-N, Dipeptidyl peptidase 4, and CEACAM116 from selected animal species using Modeler 10.254. In cases for which a particular species lacked an available

representative sequence, we chose the closest phylogenetic relative of that species for which a representative sequence was available: _Hipposideros armiger_ (great roundleaf bat) in place

of _Hipposideros ruber_ (Noack’s roundleaf bat); _Capra hircus_ (goat) in place of _Cephalophus natalensis_ (red duiker); and _Colobus angolensis palliates_ (Angolan black-and-white colobus)

in place of _Colobus guereza occidentalis_ (black-and-white colobus) (Table S2). We modeled the BHRGV-1 S protein using the well characterized SARS-CoV spike fusion protein (PDB ID: 2BEZ)

as a homologous structural template. Similarly, we used the human ACE2 (PDB ID: 1R42), Aminopeptidase-N (PDB ID: 5LHD), Dipeptidyl peptidase 4 (PDB ID: 2QT9), and CEACAM1 (PDB ID: 4QXW)

proteins as homologous structural templates in the other species analyzed (Table S2). We evaluated the quality of the resulting models using the GA341 score55, DOPE (Discrete Optimized

Protein Energy) method scores56, and the SWISS-MODEL structure assessment server57. We then refined structures with the lowest DOPE scores using molecular dynamics (MD) simulations and

further analyzed them for quality using Ramachandran Plot and MolProbity in SWISS-MODEL (_49_) (Table S3). We used HDOCK58 to model putative receptor/BGHRV-1 binding complexes. HDOCK maps

receptor and ligand protein molecules onto grids, then “docks” two molecules using a hierarchical approach based on fast Fourier transformation. To minimize bias, we applied the

template-free docking method with structures generated by Modeler 10.2. We optimized the final docked protein complexes using the AMBER99SB-ILDN force field in GROMACS59. Specifically,

docked complexes were immersed in a truncated octahedron box of TIP3P water molecules. The solvated box was further neutralized with Na + or Cl − counter ions using the tleap program. We

used Particle Mesh Ewald (PME) to calculate long-range electrostatic interactions, with a cut-off distance for long-range van der Waals (VDW) energy term of 12.0 Å, and the system minimized

without restraints. We applied 2500 cycles of steepest descent minimization followed by 5000 cycles of conjugate gradient minimization. We initiated MD simulations by heating each system in

the NVT ensemble from 0 to 300 K for 50 ps using a Langevin thermostat with a coupling coefficient of 1.0/ps and a force constant of 2.0 kcal/mol·Å2 on the complex. We ran the MD simulation

for 100 ns at a constant temperature of 300 K in the NPT ensemble with periodic boundary conditions for each system. During the MD procedure, we applied the SHAKE algorithm was to all

covalent bonds involving hydrogen atoms, with a time step of 2 fs. We calculated free energies of binding for all simulated docked structures using the molecular mechanics Poisson Boltzmann

surface area (MM-PBSA) tool in GROMACS 202260 (Table S4). To examine BGHRV-1 for a predicted hemagglutinin-esterase region, we searched for matches in the InterPro protein signature

databases using InterProScan 5.65-97.018,19. DATA AVAILABILITY All raw sequence reads were deposited in the NIH National Center for Biotechnology Information (NCBI) Sequence Read Achieve

under BioProject PRJNA1087330 (accession numbers SAMN40440184-SAMN40440195). All assembled virus genome sequences were deposited in NCBI GenBank under accession numbers OP199247 and

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54, 1951–1962 (2014). Article  CAS  PubMed  Google Scholar  Download references ACKNOWLEDGEMENTS We are grateful to the Uganda Wildlife Authority and the Uganda National Council for Science

and Technology for kindly granting permission to conduct field studies. We also thank the staff and management of Budongo Conservation Field Station for logistic support, and in particular

Jacob Ariyo, Vicent Kiiza, Stephen Mugisha and Charles Rabu for assisting with field data collection. This work was supported in part through the European Research Council, grant agreement

number 679787 ((to C. C.) and 802179) (to C.H.), the Royal Zoological Society of Scotland (to V.R.), the Armed Forces Health Surveillance Division (AFHSD), Global Emerging Infections

Surveillance (GEIS) Branch, ProMIS ID P0167_22_NM (to K.A.B-L.), Navy WUN A1417 (to K.A.B-L.), and the University of Wisconsin-Madison John D. MacArthur Research Chair (to T.L.G.). The views

expressed in this article are those of the authors and do not necessarily reflect the official policy or position of the Department of Defense, Department of the Navy, nor the U.S.

Government. Several of the authors are U.S. Government employees. This work was prepared as part of their official duties. Title 17 U.S.C. § 105 provides that ‘Copyright protection under

this title is not available for any work of the United States Government.’ Title 17 U.S.C. §101 defines a U.S. Government work as a work prepared by a military service member or employee of

the U.S. Government as part of that person’s official duties. AUTHOR INFORMATION Author notes * These authors contributed equally: Pawel Fedurek, Caroline Asiimwe. AUTHORS AND AFFILIATIONS *

Division of Psychology, Faculty of Natural Sciences, University of Stirling, Stirling, FK9 4LA, UK Pawel Fedurek * Budongo Conservation Field Station, PO Box 362, Masindi, Uganda Pawel

Fedurek, Caroline Asiimwe, Walter J. Akankwasa, Vernon Reynolds, Catherine Hobaiter, Geoffrey Muhanguzi & Klaus Zuberbühler * Biological Defense Research Directorate, Naval Medical

Research Command, Fort Detrick, MD, 21702, USA Gregory K. Rice, Regina Z. Cer, Andrew J. Bennett & Kimberly A. Bishop-Lilly * Leidos, 1750 Presidents St, Reston, VA, 20190, USA Gregory

K. Rice & Andrew J. Bennett * School of Anthropology, University of Oxford, 51/53 Banbury Road, Oxford, OX2 6PE, UK Vernon Reynolds * School of Psychology and Neuroscience, University of

St Andrews; St Mary’s Quad, South Street, St Andrews, KY16 9JP, UK Catherine Hobaiter & Klaus Zuberbühler * Department of Zoology, Entomology & Fisheries Sciences, Makerere

University, PO Box 7062, Kampala, Uganda Robert Kityo * Institute of Biology, University of Neuchâtel, Rue Emile-Argand 11, CH-2000, Neuchâtel, Switzerland Klaus Zuberbühler * Max Planck

Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103, Leipzig, Germany Catherine Crockford * Institut des Sciences Cognitives, 67 Bd Pinel, 69500, Bron, France Catherine

Crockford * Department of Anthropology, Hunter College of the City University of New York, 695 Park Avenue, New York, NY, 10065, USA Jessica M. Rothman * School of Veterinary Medicine,

Department of Pathobiological Sciences, University of Wisconsin-Madison, 1656 Linden Drive, Madison, WI, USA Tony L. Goldberg Authors * Pawel Fedurek View author publications You can also

search for this author inPubMed Google Scholar * Caroline Asiimwe View author publications You can also search for this author inPubMed Google Scholar * Gregory K. Rice View author

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Reynolds View author publications You can also search for this author inPubMed Google Scholar * Catherine Hobaiter View author publications You can also search for this author inPubMed 

Google Scholar * Robert Kityo View author publications You can also search for this author inPubMed Google Scholar * Geoffrey Muhanguzi View author publications You can also search for this

author inPubMed Google Scholar * Klaus Zuberbühler View author publications You can also search for this author inPubMed Google Scholar * Catherine Crockford View author publications You can

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Google Scholar CONTRIBUTIONS P.F., C.A., and T.L.G. contributed to the study conception and design. P.F., C.A., W.J.A., V.R., C.H., R. K., G.M., K.Z. and C.C. contributed to sample and data

collection in the field. P.F., J.M.R., G.K.R., R.Z.C. A.J.B., K.B.L. and T.L.G. conducted laboratory analyses. G.K.R., R.Z.C., A.J.B., K.B.L. and T.L.G. performed statistical analyses and

interpretations. P.F. and T.L.G. wrote the initial manuscript, and all authors read, edited and approved the final manuscript. CORRESPONDING AUTHOR Correspondence to Tony L. Goldberg. ETHICS

DECLARATIONS COMPETING INTERESTS The authors declare no competing interests. ETHICS APPROVAL The study protocol was reviewed and approved by the Uganda Wildlife Authority and the Uganda

National Council for Science and Technology and was in compliance with the guidelines of the Animal Welfare and Ethical Review Body of the University of Stirling and all applicable

regulations governing the protection of animals and research. We have complied with all relevant ethical regulations for animal use. PEER REVIEW PEER REVIEW INFORMATION _Communications

Biology_ thanks Daniel Becker, Katharina C. Wollenberg Valero and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. Primary Handling Editor: Tobias

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permissions ABOUT THIS ARTICLE CITE THIS ARTICLE Fedurek, P., Asiimwe, C., Rice, G.K. _et al._ Selective deforestation and exposure of African wildlife to bat-borne viruses. _Commun Biol_ 7,

470 (2024). https://doi.org/10.1038/s42003-024-06139-z Download citation * Received: 23 May 2023 * Accepted: 02 April 2024 * Published: 22 April 2024 * DOI:

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