{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T18:36:17Z","timestamp":1790361377570,"version":"4.1.0"},"reference-count":98,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2021,3,17]],"date-time":"2021-03-17T00:00:00Z","timestamp":1615939200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Viruses"],"abstract":"<jats:p>The recent SARS-CoV-2 pandemic has brought many questions over the origin of the virus, the threat it poses to animals both in the wild and captivity, and the risks of a permanent viral reservoir developing in animals. Animal experiments have shown that a variety of animals can become infected with the virus. While coronaviruses have been known to infect animals for decades, the true intermediate host of the virus has not been identified, with no cases of SARS-CoV-2 in wild animals. The screening of wild, farmed, and domesticated animals is necessary to help us understand the virus and its origins and prevent future outbreaks of both COVID-19 and other diseases. There is intriguing evidence that farmed mink infections (acquired from humans) have led to infection of other farm workers in turn, with a recent outbreak of a mink variant in humans in Denmark. A thorough examination of the current knowledge and evidence of the ability of SARS-CoV-2 to infect different animal species is therefore vital to evaluate the threat of animal to human transmission and reverse zoonosis.<\/jats:p>","DOI":"10.3390\/v13030494","type":"journal-article","created":{"date-parts":[[2021,3,17]],"date-time":"2021-03-17T21:43:31Z","timestamp":1616017411000},"page":"494","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":76,"title":["SARS-CoV-2 Infections in Animals: Reservoirs for Reverse Zoonosis and Models for Study"],"prefix":"10.3390","volume":"13","author":[{"given":"Tessa","family":"Prince","sequence":"first","affiliation":[{"name":"NIHR Health Protection Unit in Emerging and Zoonotic Infections, Department of Clinical Infection, Microbiology and Immunology, University of Liverpool, Liverpool L69 7TX, UK"},{"name":"Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Liverpool L69 7BE, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shirley L.","family":"Smith","sequence":"additional","affiliation":[{"name":"Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Liverpool L69 7BE, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alan D.","family":"Radford","sequence":"additional","affiliation":[{"name":"Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Liverpool L69 7BE, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tom","family":"Solomon","sequence":"additional","affiliation":[{"name":"NIHR Health Protection Unit in Emerging and Zoonotic Infections, Department of Clinical Infection, Microbiology and Immunology, University of Liverpool, Liverpool L69 7TX, UK"},{"name":"Institute of Infection, Veterinary and Ecological Sciences, University of Liverpool, Liverpool L69 7BE, UK"},{"name":"Walton Centre NHS Foundation Trust, Liverpool L9 7LJ, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7567-7185","authenticated-orcid":false,"given":"Grant L.","family":"Hughes","sequence":"additional","affiliation":[{"name":"NIHR Health Protection Unit in Emerging and Zoonotic Infections, Department of Clinical Infection, Microbiology and Immunology, University of Liverpool, Liverpool L69 7TX, UK"},{"name":"Centre for Neglected Tropical Disease, Departments of Vector Biology and Tropical Disease Biology, Liverpool School of Tropical Medicine, Pembroke Place, Liverpool L3 5QA, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3465-0848","authenticated-orcid":false,"given":"Edward I.","family":"Patterson","sequence":"additional","affiliation":[{"name":"Centre for Neglected Tropical Disease, Departments of Vector Biology and Tropical Disease Biology, Liverpool School of Tropical Medicine, Pembroke Place, Liverpool L3 5QA, UK"},{"name":"Department of Biological Sciences, Brock University, St. Catharines, ON L2S 3A1, Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,3,17]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"270","DOI":"10.1038\/s41586-020-2012-7","article-title":"A pneumonia outbreak associated with a new coronavirus of probable bat origin","volume":"579","author":"Zhou","year":"2020","journal-title":"Nature"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"536","DOI":"10.1038\/s41564-020-0695-z","article-title":"The species Severe acute respiratory syndrome-related coronavirus: Classifying 2019-nCoV and naming it SARS-CoV-2","volume":"5","author":"Gorbalenya","year":"2020","journal-title":"Nat. Microbiol."},{"key":"ref_3","first-page":"325","article-title":"Bats, civets and the emergence of SARS","volume":"315","author":"Wang","year":"2007","journal-title":"Curr. Top. Microbiol. Immunol."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Dudas, G., Carvalho, L.M., Rambaut, A., and Bedford, T. (2018). MERS-CoV spillover at the camel-human interface. Elife, 7.","DOI":"10.7554\/eLife.31257"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1016\/S0140-6736(20)30251-8","article-title":"Genomic characterisation and epidemiology of 2019 novel coronavirus: Implications for virus origins and receptor binding","volume":"395","author":"Lu","year":"2020","journal-title":"Lancet"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"450","DOI":"10.1038\/s41591-020-0820-9","article-title":"The proximal origin of SARS-CoV-2","volume":"26","author":"Andersen","year":"2020","journal-title":"Nat. Med."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"108693","DOI":"10.1016\/j.vetmic.2020.108693","article-title":"Novel human coronavirus (SARS-CoV-2): A lesson from animal coronaviruses","volume":"244","author":"Decaro","year":"2020","journal-title":"Vet. Microbiol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1016\/j.cell.2020.02.052","article-title":"SARS-CoV-2 Cell Entry Depends on ACE2 and TMPRSS2 and Is Blocked by a Clinically Proven Protease Inhibitor","volume":"181","author":"Hoffmann","year":"2020","journal-title":"Cell"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"631","DOI":"10.1002\/path.1570","article-title":"Tissue distribution of ACE2 protein, the functional receptor for SARS coronavirus. A first step in understanding SARS pathogenesis","volume":"203","author":"Hamming","year":"2004","journal-title":"J. Pathol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1128\/JVI.00127-20","article-title":"Receptor Recognition by the Novel Coronavirus from Wuhan: An Analysis Based on Decade-Long Structural Studies of SARS Coronavirus","volume":"94","author":"Wan","year":"2020","journal-title":"J. Virol."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"2196","DOI":"10.1016\/j.cub.2020.05.023","article-title":"A Novel Bat Coronavirus Closely Related to SARS-CoV-2 Contains Natural Insertions at the S1\/S2 Cleavage Site of the Spike Protein","volume":"30","author":"Zhou","year":"2020","journal-title":"Curr. Biol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1002\/jmv.25682","article-title":"Cross-species transmission of the newly identified coronavirus 2019-nCoV","volume":"92","author":"Ji","year":"2020","journal-title":"J. Med. Virol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1346","DOI":"10.1016\/j.cub.2020.03.022","article-title":"Probable Pangolin Origin of SARS-CoV-2 Associated with the COVID-19 Outbreak","volume":"30","author":"Zhang","year":"2020","journal-title":"Curr. Biol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1038\/s41586-020-2313-x","article-title":"Isolation of SARS-CoV-2-related coronavirus from Malayan pangolins","volume":"583","author":"Xiao","year":"2020","journal-title":"Nature"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Chen, D., Sun, J., Zhu, J., Ding, X., Lan, T., Zhu, L., Xiang, R., Ding, P., Wang, H., and Wang, X. (2020). Single-cell screening of SARS-CoV-2 target cells in pets, livestock, poultry and wildlife. bioRxiv.","DOI":"10.1101\/2020.06.13.149690"},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Lee, J., Hughes, T., Lee, M.H., Field, H., Rovie-Ryan, J.J., Sitam, F.T., Sipangkui, S., Nathan, S.K., Ramirez, D., and Kumar, S.V. (2020). No evidence of coronaviruses or other potentially zoonotic viruses in Sunda pangolins (Manis javanica) entering the wildlife trade via Malaysia. bioRxiv.","DOI":"10.1101\/2020.06.19.158717"},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Ulrich, L., Wernike, K., Hoffmann, D., Mettenleiter, T.C., and Ulrich, L. (2020). Experimental infection of cattle with SARS-CoV-2. bioRxiv.","DOI":"10.1101\/2020.08.25.254474"},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Gultom, M., Licheri, M., Laloli, L., Wider, M., Straessle, M., Steiner, S., Kratzel, A., Thao, T.T., Stalder, H., and Portmann, J. (2020). Susceptibility of well-differentiated airway epithelial cell 1 cultures from domestic and wildlife animals to SARS-CoV-2 2 3. bioRxiv.","DOI":"10.1101\/2020.11.10.374587"},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Cahan, E. (2020). COVID-19 hits U.S. mink farms after ripping through Europe. Science.","DOI":"10.1126\/science.abe3870"},{"key":"ref_20","unstructured":"OIE\u2014World Organisation for Animal Health (2020, August 28). Events in Animals. Available online: https:\/\/www.oie.int\/en\/scientific-expertise\/specific-information-and-recommendations\/questions-and-answers-on-2019novel-coronavirus\/events-in-animals\/."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Rabalski, L., Kosinski, M., Smura, T., Aaltonen, K., Kant, R., Szewczyk, B., and Grzybek, M. (2020). Detection and molecular characterisation of SARS-CoV-2 1 in farmed mink (Neovision vision) in Poland 2. bioRxiv.","DOI":"10.1101\/2020.12.24.422670"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2001005","DOI":"10.2807\/1560-7917.ES.2020.25.23.2001005","article-title":"SARS-CoV-2 infection in farmed minks, the Netherlands, April and May 2020","volume":"25","author":"Oreshkova","year":"2020","journal-title":"Eurosurveillance"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1126\/science.abe5901","article-title":"Transmission of SARS-CoV-2 on mink farms between humans and mink and back to humans","volume":"371","author":"Munnink","year":"2020","journal-title":"Science"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1169","DOI":"10.1126\/science.368.6496.1169","article-title":"Coronavirus rips through Dutc mink farms, triggering culls","volume":"368","author":"Enserink","year":"2020","journal-title":"Science"},{"key":"ref_25","unstructured":"Van Dorp, L., Tan, C.C., Lam, S.D., Richard, D., Owen, C., Berchtold, D., Orengo, C., and Balloux, F. (2020). Recurrent mutations in SARS-CoV-2 genomes isolated from mink point to rapid host-adaptation. bioRxiv."},{"key":"ref_26","unstructured":"WHO (2020, November 10). SARS-CoV-2 Mink-Associated Variant Strain\u2014Denmark. Available online: https:\/\/www.who.int\/csr\/don\/06-november-2020-mink-associated-sars-cov2-denmark\/en\/."},{"key":"ref_27","unstructured":"ECDC (2021, January 07). Detection of New SARS-CoV-2 Variants Related to Mink. Eur. Cent. Dis. Prev. Control., Available online: https:\/\/www.ecdc.europa.eu\/sites\/default\/files\/documents\/RRA-SARS-CoV-2-in-mink-12-nov-2020.pdf."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"WHO (2020, November 10). Coronavirus Disease (COVID-19) Outbreak\u2014Mink-Strain of COVID-19 Virus in Denmark. Available online: https:\/\/www.euro.who.int\/en\/health-topics\/health-emergencies\/coronavirus-covid-19\/news\/news\/2020\/11\/mink-strain-of-covid-19-virus-in-denmark.","DOI":"10.46945\/bpj.10.1.03.01"},{"key":"ref_29","unstructured":"(2021, January 07). GOV.UK. New SARS-CoV-2 Variant\u2014GOV.UK, Available online: https:\/\/www.gov.uk\/government\/collections\/new-sars-cov-2-variant."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"340","DOI":"10.1038\/d41586-020-03218-z","article-title":"COVID mink analysis shows mutations are not dangerous\u2014Yet","volume":"587","author":"Mallapaty","year":"2020","journal-title":"Nature"},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Munnink, B.B., Sikkema, R.S., Nieuwenhuijse, D.F., Molenaar, R.J., Munger, E., Molenkamp, R., van der Spek, A., Tolsma, P., Rietveld, A., and Brouwer, M. (2020). Jumping back and forth: Anthropozoonotic and zoonotic transmission of SARS-CoV-2 on mink farms Affiliations. bioRxiv.","DOI":"10.1101\/2020.09.01.277152"},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Mykytyn, A.Z., Lamers, M.M., Okba, N.M., Breugem, T.I., Schipper, D., van den Doel, P.B., van Run, P., van Amerongen, G., de Waal, L., and Koopmans, M.P. (2020). Susceptibility of rabbits to SARS-CoV-2. bioRxiv.","DOI":"10.1101\/2020.08.27.263988"},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Wang, L., Mitchell, P.K., Calle, P.P., Bartlett, S.L., McAloose, D., Killian, M.L., Yuan, F., Fang, Y., Goodman, L.B., and Fredrickson, R. (2020). 2020 Complete Genome Sequence of SARS-CoV-2 in a Tiger from a U.S. Zoological Collection. Microbiol. Resour. Announc., 9.","DOI":"10.1128\/MRA.00468-20"},{"key":"ref_34","unstructured":"USDA (2020, August 12). USDA Statement on the Confirmation of COVID-19 in a Tiger in New York, Available online: https:\/\/www.aphis.usda.gov\/aphis\/newsroom\/news\/sa_by_date\/sa-2020\/ny-zoo-covid-19."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"McAloose, D., Laverack, M., Wang, L., Killian, M.L., Caserta, L.C., Yuan, F., Mitchell, P.K., Queen, K., Mauldin, M.R., and Cronk, B.D. (2020). From people to Panthera: Natural SARS-CoV-2 infection in tigers and lions at the Bronx Zoo. bioRxiv.","DOI":"10.1101\/2020.07.22.213959"},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Bartlett, S.L., Diel, D.G., Wang, L., Zec, S., Laverack, M., Martins, M., Caserta, L.C., Killian, M.L., Terio, K., and Olmstead, C. (2020). SARS-CoV-2 Infection and Longitudinal Fecal Screening In Malayan Tigers (Panthera tigris jacksoni), Amur Tigers (Panthera tigris altaica), And African Lions (Panthera leo krugeri) At The Bronx Zoo, New York, USA. bioRxiv.","DOI":"10.1101\/2020.08.14.250928"},{"key":"ref_37","unstructured":"USDA APHIS (2021, January 13). Confirmation of COVID-19 in Gorillas at a California Zoo, Available online: https:\/\/content.govdelivery.com\/accounts\/USDAAPHIS\/bulletins\/2b5837f\/."},{"key":"ref_38","first-page":"84","article-title":"Surveillance on severe acute respiratory syndrome associated coronavirus in animals at a live animal market of Guangzhou in 2004","volume":"26","author":"Wang","year":"2005","journal-title":"Zhonghua Liu Xing Bing Xue Za Zhi"},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Shi, J., Wen, Z., Zhong, G., Yang, H., Wang, C., Liu, R., He, X., Shuai, L., Sun, Z., and Zhao, Y. (2020). Susceptibility of ferrets, cats, dogs, and different domestic animals to SARS-coronavirus-2. bioRxiv.","DOI":"10.1101\/2020.03.30.015347"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"2322","DOI":"10.1080\/22221751.2020.1833687","article-title":"SARS-CoV-2 infection, disease and transmission in domestic cats Running title: SARS-CoV-2 in domestic cats","volume":"9","author":"Gaudreault","year":"2020","journal-title":"Emerg. Microbes Infect."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"592","DOI":"10.1056\/NEJMc2013400","article-title":"Transmission of SARS-CoV-2 in Domestic Cats","volume":"383","author":"Halfmann","year":"2020","journal-title":"N. Engl. J. Med."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Hamer, S.A., Pauvolid-Corr\u00eaa, A., Zecca, I.B., Davila, E., Auckland, L.D., Roundy, C.M., Tang, W., Torchetti, M., Killian, M.L., and Jenkins-Moore, M. (2020). Natural SARS-CoV-2 infections, including virus isolation, among serially tested cats and dogs in households with confirmed human COVID-19 cases in Texas, USA. bioRxiv.","DOI":"10.1101\/2020.12.08.416339"},{"key":"ref_43","unstructured":"USDA APHIS (2020, August 28). Confirmed Cases of SARS-CoV-2 in Animals in the United States, Available online: https:\/\/www.aphis.usda.gov\/aphis\/ourfocus\/animalhealth\/sa_one_health\/sars-cov-2-animals-us."},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Musso, N., Costantino, A., La Spina, S., Finocchiaro, A., Andronico, F., Stracquadanio, S., Liotta, L., Visalli, R., and Emmanuele, G. (2020). New SARS-CoV-2 Infection Detected in an Italian Pet Cat by RT-qPCR from Deep Pharyngeal Swab. Pathogens, 9.","DOI":"10.3390\/pathogens9090746"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"3071","DOI":"10.3201\/eid2612.202786","article-title":"SARS-CoV-2 in Quarantined Domestic Cats from COVID-19 Households or Close Contacts, Hong Kong, China","volume":"26","author":"Barrs","year":"2020","journal-title":"Emerg. Infect. Dis."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Sailleau, C., Dumarest, M., Vanhomwegen, J., Delaplace, M., Caro, V., Kwasiborski, A., Hourdel, V., Chevaillier, P., Barbarino, A., and Comtet, L. (2020). First detection and genome sequencing of SARS-CoV-2 in an infected cat in France. Transbound. Emerg. Dis.","DOI":"10.22541\/au.158990358.89168563"},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Ruiz-Arrondo, I., Portillo, A., Palomar, A.M., Santib\u00e1\u00f1ez, S., Santib\u00e1\u00f1ez, P., Cervera, C., and Oteo, J.A. (2020). Detection of SARS-CoV-2 in pets living with COVID-19 owners diagnosed during the COVID-19 lockdown in Spain: A case of an asymptomatic cat with SARS-CoV-2 in Europe. Transbound. Emerg. Dis.","DOI":"10.1101\/2020.05.14.20101444"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"24790","DOI":"10.1073\/pnas.2010817117","article-title":"Detection of SARS-CoV-2 in a cat owned by a COVID-19-affected patient in Spain","volume":"117","author":"Puig","year":"2020","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Hosie, M.J., Epifano, I., Herder, V., Orton, R., Stevenson, A., Johnson, N., MacDonald, E., Dunbar, D., McDonald, M., and Howie, F. (2020). Respiratory disease in cats associated with human-to-cat transmission of SARS-CoV-2 in the UK. bioRxiv.","DOI":"10.1101\/2020.09.23.309948"},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Neira, V., Brito, B., Ag\u00fcero, B., Berrios, F., Vald\u00e9s, V., Gutierrez, A., Ariyama, N., Espinoza, P., Retamal, P., and Holmes, E.C. (2020). A household case evidences shorter shedding of SARS-CoV-2 in naturally infected cats 6 compared to their human owners. 7 Affiliations. bioRxiv.","DOI":"10.1101\/2020.10.31.20220608"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"2013","DOI":"10.1080\/22221751.2020.1817796","article-title":"A serological survey of SARS-CoV-2 in cat in Wuhan","volume":"9","author":"Zhang","year":"2020","journal-title":"Emerg. Microbes Infect."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1745","DOI":"10.1111\/tbed.13577","article-title":"Serological survey of SARS-CoV-2 for experimental, domestic, companion and wild animals excludes intermediate hosts of 35 different species of animals","volume":"67","author":"Deng","year":"2020","journal-title":"Transbound. Emerg. Dis."},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Deng, J., Liu, Y., Sun, C., Bai, J., Sun, J., Hao, L., Li, X., and Tian, K. (2020). SARS-CoV-2 Serological Survey of Cats in China before and after the Pandemic. Virol. Sin., 1\u20133.","DOI":"10.1007\/s12250-020-00284-5"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"100164","DOI":"10.1016\/j.onehlt.2020.100164","article-title":"Absence of SARS-CoV-2 infection in cats and dogs in close contact with a cluster of COVID-19 patients in a veterinary campus","volume":"10","author":"Temmam","year":"2020","journal-title":"One Health"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"100192","DOI":"10.1016\/j.onehlt.2020.100192","article-title":"High prevalence of SARS-CoV-2 antibodies in pets from COVID-19+ households","volume":"11","author":"Fritz","year":"2020","journal-title":"One Health"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1038\/s41467-020-20097-0","article-title":"Evidence of exposure to SARS-CoV-2 in cats and dogs from households in Italy","volume":"11","author":"Patterson","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Gaudreault, N.N., Carossino, M., Morozov, I., Trujillo, J.D., Meekins, D.A., Madden, D.W., Cool, K., Artiaga, B.L., McDowell, C., and Bold, D. (2021). Experimentally re-infected cats do not transmit SARS-CoV-2. bioRxiv.","DOI":"10.1101\/2021.01.18.427182"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"776","DOI":"10.1038\/s41586-020-2334-5","article-title":"Infection of dogs with SARS-CoV-2","volume":"586","author":"Sit","year":"2020","journal-title":"Nature"},{"key":"ref_59","unstructured":"Delong, J. (2020, August 17). Dutch Minister Confirms Dog, Three Cats Have Caught Novel Coronavirus|Armenian American Reporter. Available online: https:\/\/www.reporter.am\/dutch-minister-confirms-dog-three-cats-have-caught-novel-coronavirus\/."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Peris\u00e9-Barrios, A.J., Tomeo-Mart\u00edn, B.D., G\u00f3mez-Ochoa, P., Delgado-Bonet, P., Plaza, P., Palau-Concejo, P., Gonz\u00e1lez, J., Ortiz-D\u00edez, G., Mel\u00e9ndez-Lazo, A., and Gentil, M. (2020). Humoral response to SARS-CoV-2 by healthy and sick dogs during COVID-19 pandemic in Spain. bioRxiv.","DOI":"10.1101\/2020.09.22.308023"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1089\/vbz.2020.2650","article-title":"A Critical Needs Assessment for Research in Companion Animals and Livestock Following the Pandemic of COVID-19 in Humans","volume":"20","author":"McNamara","year":"2020","journal-title":"Vector-Borne Zoonotic Dis."},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Fagre, A., Lewis, J., Eckley, M., Zhan, S., Rocha, S.M., Sexton, N.R., Burke, B., Geiss, B., Peersen, O., and Kading, R. (2020). SARS-CoV-2 infection, neuropathogenesis and transmission among deer mice: Implications for reverse zoonosis to New World rodents. bioRxiv.","DOI":"10.1101\/2020.08.07.241810"},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Griffin, B.D., Chan, M., Tailor, N., Mendoza, E.J., Leung, A., Warner, B.M., Duggan, A.T., Moffat, E., He, S., and Garnett, L. (2020). North American deer mice are susceptible to SARS-CoV-2. bioRxiv.","DOI":"10.1101\/2020.07.25.221291"},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"e218","DOI":"10.1016\/S2666-5247(20)30089-6","article-title":"SARS-CoV-2 in fruit bats, ferrets, pigs, and chickens: An experimental transmission study","volume":"1","author":"Schlottau","year":"2020","journal-title":"Lancet Microbe"},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Pickering, B.S., Smith, G., Pinette, M.M., Embury-Hyatt, C., Moffat, E., Marszal, P., and Lewis, C.E. (2020). Susceptibility of domestic swine to experimental infection with SARS-CoV-2. bioRxiv.","DOI":"10.1101\/2020.09.10.288548"},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Suarez, D.L., Pantin-Jackwood, M.J., Swayne, D.E., Lee, S.A., Deblois, S.M., and Spackman, E. (2020). Lack of susceptibility of poultry to SARS-CoV-2 and MERS-CoV. bioRxiv.","DOI":"10.1101\/2020.06.16.154658"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Freuling, C.M., Breithaupt, A., M\u00fcller, T., Sehl, J., Balkema-Buschmann, A., Rissmann, M., Klein, A., Wylezich, C., H\u00f6per, D., and Wernike, K. (2020). 2020 Susceptibility of raccoon dogs for SARS-CoV-2 Title: Susceptibility of raccoon dogs for experimental SARS-CoV-2 infection. bioRxiv.","DOI":"10.1101\/2020.08.19.256800"},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Zhao, Y., Wang, J., Kuang, D., Xu, J., Yang, M., Ma, C., Zhao, S., Li, J., Long, H., and Ding, K. (2020). 2020 Susceptibility of tree shrew to SARS-CoV-2 infection. bioRxiv.","DOI":"10.1101\/2020.04.30.029736"},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Palmer, M.V., Martins, M., Falkenberg, S., Buckley, A.C., Caserta, L.C., Mitchell, P.K., Cassmann, E., Rollins, A., Zylich, N.C., and Renshaw, R.W. (2021). Susceptibility of white-tailed deer (Odocoileus virginianus) to SARS-CoV-2. bioRxiv.","DOI":"10.1101\/2021.01.13.426628"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/s12985-017-0766-9","article-title":"Detection of alpha- and betacoronaviruses in rodents from Yunnan, China","volume":"14","author":"Ge","year":"2017","journal-title":"Virol. J."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"830","DOI":"10.1038\/s41586-020-2312-y","article-title":"The pathogenicity of SARS-CoV-2 in hACE2 transgenic mice","volume":"583","author":"Bao","year":"2020","journal-title":"Nature"},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Clark, J.J., Penrice-Randal, R., Sharma, P., Kipar, A., Dong, X., Davidson, A.D., Williamson, M.K., Matthews, D.A., Turtle, L., and Prince, T. (2020). Sequential infection with influenza A virus followed by severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) leads to more severe disease and encephalitis in a mouse model of COVID-19. bioRxiv.","DOI":"10.1101\/2020.10.13.334532"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.cell.2020.05.027","article-title":"Pathogenesis of SARS-CoV-2 in Transgenic Mice Expressing Human Angiotensin-Converting Enzyme 2","volume":"182","author":"Jiang","year":"2020","journal-title":"Cell"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1070","DOI":"10.1016\/j.cell.2020.09.050","article-title":"A Mouse-adapted SARS-CoV-2 induces Acute Lung Injury (ALI) and mortality in Standard Laboratory Mice","volume":"183","author":"Leist","year":"2020","journal-title":"Cell"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"2428","DOI":"10.1093\/cid\/ciaa644","article-title":"Simulation of the clinical and pathological manifestations of Coronavirus Disease 2019 (COVID-19) in golden Syrian hamster model: Implications for disease pathogenesis and transmissibility","volume":"71","author":"Chan","year":"2020","journal-title":"Clin. Infect. Dis."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Bertzbach, L.D., Vladimirova, D., Dietert, K., Abdelgawad, A., Gruber, A.D., Osterrieder, N., and Trimpert, J. (2020). SARS-CoV-2 infection of Chinese hamsters (Cricetulus griseus) reproduces COVID-19 pneumonia in a well-established small animal model. Transbound. Emerg. Dis.","DOI":"10.1111\/tbed.13837"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1038\/s41467-020-17367-2","article-title":"SARS-CoV-2 is transmitted via contact and via the air between ferrets","volume":"11","author":"Richard","year":"2020","journal-title":"Nat. Commun."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"704","DOI":"10.1016\/j.chom.2020.03.023","article-title":"Infection and Rapid Transmission of SARS-CoV-2 in Ferrets","volume":"27","author":"Kim","year":"2020","journal-title":"Cell Host Microbe"},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Kutter, J.S., de Meulder, D., Bestebroer, T.M., Lexmond, P., Fouchier, R.A., and Herfst, S. (2020). SARS-CoV and SARS-CoV-2 are transmitted through the air between ferrets over more than one meter distance. bioRxiv.","DOI":"10.21203\/rs.3.rs-93725\/v1"},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"Sawatzki, K., Hill, N., Puryear, W., Foss, A., and Stone, J. (2020). Ferrets not infected by SARS-CoV-2 in a high-exposure domestic setting. bioRxiv.","DOI":"10.1101\/2020.08.21.254995"},{"key":"ref_81","unstructured":"Bao, L., Deng, W., Gao, H., Xiao, C., Liu, J., Xue, J., Lv, Q., Liu, J., Yu, P., and Xu, Y. (2020). Reinfection could not occur in SARS-CoV-2 infected rhesus macaques. bioRxiv."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"812","DOI":"10.1126\/science.abc4776","article-title":"SARS-CoV-2 infection protects against rechallenge in rhesus macaques","volume":"369","author":"Chandrashekar","year":"2020","journal-title":"Science"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"1012","DOI":"10.1126\/science.abb7314","article-title":"Comparative pathogenesis of COVID-19, MERS, and SARS in a nonhuman primate model","volume":"368","author":"Rockx","year":"2020","journal-title":"Science"},{"key":"ref_84","doi-asserted-by":"crossref","unstructured":"Woolsey, C., Borisevich, V., Prasad, A.N., Agans, K.N., Deer, D.J., Dobias, N.S., Heymann, J.C., Foster, S.L., Levine, C.B., and Medina, L. (2020). Establishment of an African green monkey model for COVID-19. bioRxiv.","DOI":"10.1101\/2020.05.17.100289"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/s12985-020-01396-w","article-title":"Intranasal exposure of African green monkeys to SARS-CoV-2 results in acute phase pneumonia with shedding and lung injury still present in the early convalescence phase","volume":"17","author":"Cross","year":"2020","journal-title":"Virol. J."},{"key":"ref_86","doi-asserted-by":"crossref","unstructured":"Lu, S., Zhao, Y., Yu, W., Yang, Y., Gao, J., Wang, J., Kuang, D., Yang, M., Yang, J., and Ma, C. (2020). Comparison of SARS-CoV-2 infections among 3 species of non-human primates. bioRxiv.","DOI":"10.1101\/2020.04.08.031807"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"138764","DOI":"10.1016\/j.scitotenv.2020.138764","article-title":"First confirmed detection of SARS-CoV-2 in untreated wastewater in Australia: A proof of concept for the wastewater surveillance of COVID-19 in the community","volume":"728","author":"Ahmed","year":"2020","journal-title":"Sci. Total Environ."},{"key":"ref_88","doi-asserted-by":"crossref","unstructured":"Mathavarajah, S., Stoddart, A.K., Gagnon, G.A., and Dellaire, G. (2020). Pandemic danger to the deep: The risk of marine mammals contracting SARS-CoV-2 from wastewater. bioRxiv.","DOI":"10.1101\/2020.08.13.249904"},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"1795","DOI":"10.3389\/fmicb.2020.01795","article-title":"Coronaviruses in the Sea","volume":"11","author":"Mordecai","year":"2020","journal-title":"Front. Microbiol."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1007\/s12250-020-00251-0","article-title":"SARS-CoV-2 Does Not Replicate in Aedes Mosquito Cells nor Present in Field-Caught Mosquitoes from Wuhan","volume":"35","author":"Xia","year":"2020","journal-title":"Virol. Sin."},{"key":"ref_91","first-page":"1","article-title":"SARS-CoV-2 failure to infect or replicate in mosquitoes: An extreme challenge","volume":"10","author":"Huang","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_92","doi-asserted-by":"crossref","unstructured":"Balaraman, V., Drolet, B.S., Gaudreault, N.N., Wilson, W.C., Owens, J., Bold, D., Swanson, D.A., Jasperson, D.C., Noronha, L.E., and Richt, J.A. (2020). Susceptibility of midge and mosquito vectors to SARS-CoV-2 by natural route of infection. bioRxiv.","DOI":"10.1101\/2020.09.29.317289"},{"key":"ref_93","doi-asserted-by":"crossref","unstructured":"Dai, M., Li, H., Yan, N., Huang, J., Zhao, L., Xu, S., Jiang, S., Pan, C., and Liao, M. (2020). Long-term survival of salmon-attached SARS-CoV-2 at 4 \u00b0C as a potential source of 2 transmission in seafood markets Running Title: Survival of salmon-attached SARS-CoV-2. bioRxiv.","DOI":"10.1101\/2020.09.06.284695"},{"key":"ref_94","unstructured":"(2021, January 18). 2021 COVID-19: Ice Cream Tests Positive for Coronavirus in China|World News|Sky News. Available online: https:\/\/news.sky.com\/story\/covid-19-ice-cream-tests-positive-for-coronavirus-in-china-12188761."},{"key":"ref_95","doi-asserted-by":"crossref","unstructured":"Fisher, D., Reilly, A., Kang, A., Zheng, E., Cook, A.R., and Anderson, D.E. (2020). Seeding of outbreaks of COVID-19 by contaminated fresh and frozen food. bioRxiv.","DOI":"10.1101\/2020.08.17.255166"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"887","DOI":"10.15585\/mmwr.mm6927e2","article-title":"Update: COVID-19 Among Workers in Meat and Poultry Processing Facilities\u2014United States, April\u2013May 2020","volume":"69","author":"Waltenburg","year":"2020","journal-title":"Morb. Mortal. Wkly. Rep."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"260","DOI":"10.1017\/S0950268811000379","article-title":"Multiple norovirus outbreaks linked to imported frozen raspberries","volume":"140","author":"Sarvikivi","year":"2012","journal-title":"Epidemiol. Infect."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"777","DOI":"10.1016\/S0140-6736(21)00424-4","article-title":"Future scenarios for the COVID-19 pandemic","volume":"397","author":"Kegg","year":"2021","journal-title":"Lancet"}],"container-title":["Viruses"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1999-4915\/13\/3\/494\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T05:37:05Z","timestamp":1760161025000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1999-4915\/13\/3\/494"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,3,17]]},"references-count":98,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2021,3]]}},"alternative-id":["v13030494"],"URL":"https:\/\/doi.org\/10.3390\/v13030494","relation":{},"ISSN":["1999-4915"],"issn-type":[{"value":"1999-4915","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,3,17]]}}}