Molecular Detection of Coronavirus and Paramyxovirus in Phyllostimid Bat Communities From Ecuador’s Northwestern Chocoan Forests | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Molecular Detection of Coronavirus and Paramyxovirus in Phyllostimid Bat Communities From Ecuador’s Northwestern Chocoan Forests Camila Acosta-López, Jonathan Bustamante, Maria Mercedes Gavilanez This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4030611/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Bats are important reservoirs for pathogenic viruses. Amidst rising human-wildlife interactions, the risk of potential spillover of zoonotic diseases escalates. We aimed to identify Coronavirus and Paramyxovirus in bat saliva samples for species inhabiting Ecuador’s northwestern Choco region. Sampling was undertaken at FCAT reserve in the Esmeraldas province in November 2022. A total of 123 samples from individuals appertaining 20 species, were analyzed using RT-PCR. We recorded 28 positive cases for paramyxovirus, 18 for coronavirus and four co-infections. These findings underscore the necessity of enhancing zoonotic surveillance and establishing viral diversity monitoring strategies. Figures Figure 1 Full Text Bats are classified as the fourth largest group of mammals globally, with their habitat spanning every continent except Antarctica [1]. Ecuador is distinguished as the fourth richest country in bat biodiversity, hosting 171 different species [2]. Serving as reservoirs for various pathogens, bats notably harbor coronaviruses and paramyxoviruses. About 16% of species carry coronaviruses — evidenced by the discovery of 4,000 unique coronavirus sequences. These viruses are linked to significant zoonotic viruses, such as SARS-CoV, SARS-CoV2, and MERS-CoV [3]. Paramyxoviruses, also widespread among bat populations, with 657 unique sequences identified, underline their role as progenitors of numerous diseases affecting humans and animals [4]. The surfacing of zoonotic viral diseases could be driven by factors such as human population growth, urbanization, and agricultural practices. These factors intensify human-animal interactions, elevating the likelihood of zoonosis [5]. In 2002, the outbreak of SARS-CoV-1, originating from bats and transmitted through civet cats to humans, resulted in 8,098 infections and 774 fatalities, marking a mortality rate of 9.6% [6]. Considering the recent pandemic, a consensus among epidemiologists’ points to SARS-CoV-2 originating from a market trading wild animals for human consumption in Wuhan, China. The prevailing hypothesis is that bats, the initial hosts, transmitted the virus to pangolins, which in turn infected humans [7, 8]. Human-bat interactions can also enhance paramyxovirus transmission. A study in Cameroon found antibodies against paramyxoviruses, like Nipah and Hendra, in seven people [9]. In Malaysia's Tioman Island, the Tioman virus, a member of the Paramyxoviridae family affecting bats, pigs, and humans with flu-like symptoms, was studied. Antibodies against the Tioman virus were found in 5 of the 169 individuals tested, indicating its presence and potential health impact [10]. While zoonotic diseases pose a substantial threat, Ecuador lacks data on them. A review covering 25 studies with 9,371 subjects highlighted Brazil, Mexico, US, Jamaica, Peru, Bolivia, and Argentina for coronavirus prevalence in bats, but Ecuador remains unstudied [11]. A similar scene occurs for paramyxovirus, indicating a critical gap in understanding and managing potential zoonotic risks within the country [12–16]. This study's objective was to assess the presence of these viruses in bats from the Fundación para la Conservación de los Andes Tropicales (FCAT) in Northwestern Ecuador. During November 2022, bats were systematically trapped through mist nets placed along known bat flight and feeding paths over ten nights at the FCAT Reserve in Esmeraldas province in three differently forested areas (Figure 1). Individuals were subsequently identified to species level with the Guide to the Mammals of Ecuador [17]. A buccal swab sample was collected from all captured bats using sterile polystyrene swabs. Swabs were preserved in 500 uL of DNA Shield (Ecogen) until analysis. For viral RNA extraction the AccuPrep Dx Viral RNA Extraction kit (Bioneer) was used, following manufacturer's instructions. A total of 33 samples were analyzed with a two-step RT-PCR and 90 of them with a one-step RT-PCR. For the reverse transcription was performed at 50°C for 15 minutes and 85°C for 5 minutes with the following protocol: 10 μL of nuclease-free water, 4 μL of 5X RT Buffer, 2 μL of RNA, and 1 μL of reverse transcriptase. Primers for the helicase gene were used for coronavirus while primers the RNA-dependent RNA polymerase gene (RdRp) were used for paramyxovirus. For the PCR reagents were added as follows: 10X Buffer, 100 pmol/μL of each primer, 1,5 μM MgCl2, 10μM dNTPs, 1 μL (1X) Fastgen Taq DNA polimerase and 2 μL of cDNA. For coronavirus the PCR began with a 95°C denaturation – 15 min, followed by 45 cycles of denaturation 95°C – 1 min, 48°C – 1 min and 72°C – 1 min, with a final extension of 10 minutes [18]. For paramyxovirus the PCR conditions were initial denaturation 60°C – 15 min, 98°C denaturation – 30 sec, followed by 40 cycles of denaturation 98°C – 10 sec, 50°C – 30 sec and 72°C for 30 sec, concluding with a 72°C extension of 7 minutes [19]. For the 90 samples processed through one step RT-PCR; 25 μL of 2X One-Step RT-PCR Buffer, 4 μL of RT-PCR Enzyme mix, 2.5 μL of each primer (100pmol/μL), 5 μL of RNA, and 10 μL of nuclease-free water, were used. The temperature conditions were the same as those described previously for PCR. The resultant amplicon fragments were analyzed via 2% agarose gel electrophoresis. To determine the presence of coronavirus the expected fragment was 591 bp, whilst for paramyxovirus, the expected fragment size was 580 bp. In the study, 123 bats spanning 20 species were captured (see Table 1). Analysis revealed that 28 samples were positive for paramyxovirus, and 18 samples were positive for coronavirus. A total of 42 bats were infected with coronavirus and/or paramyxovirus, involving 10 species across 8 genera (see Table 2). Coinfections were found in 4 individuals. These include: Carollia perspicilata , Vampyrodes major , Artibeus lituratus and Dermanura rosenbergi . In examining the species captured, the forest zone boasted the greatest diversity and number of individuals, succeeded by the reforestation zone, and lastly, the agricultural zone (Table 1). This observation aligns with ecological principles. Research comparing bat biodiversity in untouched forest zones to that in fragmented areas revealed a direct correlation between the extent of forest coverage and the diversity and abundance of bat species [20]. Moreover, research in Colombia evaluated the species composition and richness of bats in three different sites within Nariño's tropical dry forest. From a total of 60 bats, the highest individual count was observed in the preserved dry spiny scrub. In contrast, areas subjected to disturbance showed reduced species richness. This aligns with the outcomes of the current study [21]. In a study carried out in Peru, the species identified as most prevalent were Carollia perspicillata , Carollia brevicauda , and Artibeus planirostris . Conversely, in our study, the most numerous species identified were C. perspicillata , C. brevicauda , and Dermanura rava . The overlap of two out of three predominant species in both studies can be attributed to their shared habitat preferences. C. brevicauda and C. perspicillata have a wide distribution, thriving in both the moist and arid forests located on the eastern and western slopes of the Andes encompassing a large altitudinal range [22] . Moreover, these forest environments abound with key food sources for C. perspicillata and C. brevicauda [23]. Specifically, in northwestern Ecuador, within the FCAT region, bats consume Piper spikes, which are prevalent in this area. The diet of bats in this part of Ecuador is diversified to include Cecropia concolor , Ficus eximia , Piper aduncum , Piper longistylosum , Piper tuberculatum , and Solanum sp . The widespread availability of these plant species ensures their presence in both the Peruvian study and this research. Furthermore, Carollia bats play a crucial role as seed dispersers for these plants, establishing a mutualistic relationship [24]. In the context of coronavirus and paramyxovirus detection using PCR, a study in Sao Paulo, Brazil, collected fecal samples from bats in 73 municipalities to screen for coronavirus. From the 305 samples collected, 9 samples were identified as coronavirus positive from five species: Cynomops abrasus , Cynomops planirostris , Desmodus rotundus , Glossophaga soricina , and Platyrrhinus lineatus . In contrast, the coronavirus positive species in our study were Artibeus lituratus , C. brevicauda , C. perspicillata , Dermanura rava , Dermanura rosenbergi , Vampyrodes major , and Sturnira luisi (Table 2.). It is notable that the species identified as positive in our investigation do not overlap with those found in the Brazilian study [25]. From the current study, two positive species are also distributed in the Sao Paulo area: A. lituratus and C. perspicillata [26]. However, these two species present in both regions, were not found to be infected in Brazil. This discrepancy can be attributed to environmental variations affecting each group, including differences in predator presence, trophic interactions, coinfections, and exposure to pathogens. As a result, the likelihood of identical bat species being infected with coronavirus across different regions is minimal [27]. A separate investigation into paramyxovirus in French Guiana's bat population found 103 individuals positive for the virus. The detection rates varied with the biological fluid examined; D. rotundus , for example, tested positive for paramyxovirus only in kidney and urine samples, not in saliva, lung, or blood [28]. Contrary to this, the current research identified positive paramyxovirus cases in saliva samples from A. lituratus , C. brevicauda , C. perspicillata , D. rava , D. rosenbergi , Lonchophylla concava , S. luisi , V. major , Platyrrhinus vittatus, and Vampyriscus nymphaea . Similarly, a study focused on identifying coronavirus in North American bats through RT-PCR, revealed that all 22 saliva samples tested were negative. Conversely, 6 out of 28 fecal samples were positive for the virus [29]. In comparison, the current study found coronavirus in 18 saliva samples from a total of 123 tested. This variation could be due to each virus's specific replication and excretion mechanisms [28]. Furthermore, in the second study the number of analyzed saliva samples was relatively low compared to the current research. Bats' social behavior, particularly their use of communal feeding areas, facilitates the transmission of different viruses among individuals and across colonies of the same species [29]. These communal habits extend to resting sites, where bats also dispose of their waste. Since evidence suggests that bat feces may harbor paramyxovirus and coronavirus, it's plausible to deduce that the communal lifestyle contributes to the spread of infections [30]. Consequently, the species found in abundance in this study, C. brevicauda , C. perspicillata , D. rosenbergi , and D. rava , also exhibited the highest instances of coronavirus and paramyxovirus infections. Prevalence of coronavirus was 14.63%, and 22.76% for paramyxovirus. These values are considered high in comparison with other studies, where significantly lower values were recorded with larger sample sizes. For example, a study conducted in Italy, analyzing bat saliva and feces, found 12% and 1% prevalence for coronavirus and paramyxovirus respectively, out of 302 captured bats [14]. In Costa Rica, from 421 fecal samples, only 4 of them were positive for coronavirus [31]. Similarly, in Peru, from 436 monitored individuals, a prevalence of 10.3% was recorded for paramyxovirus [15]. Due to these high values, meticulous epidemiological surveillance must be maintained in bats to detect coronavirus, paramyxovirus, and other viruses in general. This is crucial for several reasons, one of which being the variety of viruses harbored by bats. Some of the other viral families found include adenovirus, astrovirus, bornavirus, circovirus, herpesvirus, parvovirus, polyomavirus, and rhabdovirus [32]. Another crucial reason is that it could allow us to identify potential sites of zoonotic outbreaks. This enables the possibility for taking preemptive measures to prevent the spread of such diseases [33]. A total of 4 coinfections were recorded. A systematic review noted that out of 725 bats with coinfections, only 4 individuals had a coinfection with only two viruses, namely paramyxovirus and coronavirus. These findings imply that the prevalence of coinfections in the current study is also high, in relation to the number of samples [32]. This phenomenon could be attributed to peak viral proliferation phases, resulting in higher viral shedding [34]. Additionally, it might be explained by the possibility that the viral strains infecting these individuals possess molecular mechanisms that hinder the action of interferons, which are crucial for bats in regulating viral replication [29]. The likelihood of coinfections is also affected by the phylogenetic relationships among bat species. The ability of a virus to infect is contingent upon overcoming physiological and molecular defenses. Viruses are more likely to infect two bat species that are phylogenetically close, as they may share similar defenses. For example, the SARS CX1 coronavirus is evolutionarily akin to SARS-CoV-2, and both can infect identical bat species because the receptor-binding domain of their spike proteins only differs by five amino acids [35]. In this study, the bulk of positive cases were identified in C. brevicauda with 15 positive samples, followed by C. perspicillata with 10, and D. rosenbergi with 7 positive samples. Notably, among these, two cases of coinfections were detected: one in C. perspicillata and another in D. rosenbergi . This observation of coinfections may be attributed to the dense populations of the species, as well as to the phylogenetic distance between the mentioned species [35]. Findings of this study highlight a significant prevalence of coronavirus and paramyxovirus among the bat populations examined. This marks the first documentation in Ecuador of various bat species harboring infections of coronavirus and paramyxovirus. Given these findings, it underscores the imperative need for further research within Ecuador to track the viral diversity of coronavirus and paramyxovirus, particularly in regions proximate to human communities. Declarations FUNDING This work was supported by Dirección de Investigación, Universidad Central del Ecuador seed grant cifFuce-cv-fcb-1. COMPETING INTERESTS The authors have no relevant financial or non-financial interests to disclose. AUTHOR CONTRIBUTIONS Camila Acosta-López and María Mercedes Gavilanez contributed to the study conception and design. Material preparation and data collection were performed by Jonathan Bustamante. Data analyses were performed by Camila Acosta-López, María Mercedes Gavilanez and Jonathan Bustamante. The first draft of the manuscript was written by Jonathan Bustamante, all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript. DATA AVAILABILITY The datasets generated and analyzed during the current study are available from the corresponding author on reasonable request. ETHICS APPROVAL This study was performed in line with the criteria stablished in the manual for recommendations for the use and handling of wild mammals. Since the samples were collected through noninvasive techniques, no administration of pain killers or chemical agents was needed. Approval was granted by the Comisión de Investigación de la Facultad de Medicina Veterinaria y Zootecnia from Universidad Central del Ecuador. References Banerjee A, Kulcsar K, Misra V, et al (2019) Bats and coronaviruses. Viruses 11:7–9. https://doi.org/10.3390/v11010041 Méndez-Rodríguez A, Horta P, Zarza H, et al (2024) Surveying Bat-Hosted Adenoviruses and Herpesviruses: A Comprehensive Analysis. Diversity (Basel) 16 Anthony SJ, Johnson CK, Greig DJ, et al (2017) Global patterns in coronavirus diversity. Virus Evol 3:. https://doi.org/10.1093/ve/vex012 Hayman DTS (2016) Bats as Viral Reservoirs. Annu Rev Virol 3:77–99 Mcmahon B, Morand S, Gray J (2018) Ecosystem change and zoonoses in the Anthropocene. Zoonoses Public Health 755–765 Wang J-T, Chang S-C (2004) Severe acute respiratory syndrome. Curr Opin Infect Dis 17:143–148. https://doi.org/10.1097/00001432-200404000-00013 Lau SKP, Luk HKH, Wong ACP, et al (2020) Possible Bat Origin of Severe Acute Respiratory Syndrome Coronavirus 2. Emerg Infect Dis 26:1542–1547. https://doi.org/10.3201/eid2607.200092 Xiao K, Zhai J, Feng Y, et al (2020) Isolation of SARS-CoV-2-related coronavirus from Malayan pangolins. Nature 583:286–289. https://doi.org/10.1038/s41586-020-2313-x Thibault PA, Watkinson RE, Moreira-Soto A, et al (2017) Zoonotic Potential of Emerging Paramyxoviruses. In: Advances in Virus Research. pp 1–55 Koon CY, Crameri G, Wang L, et al (2007) Serological evidence of possible human infection with Tioman virus, a newly described paramyxovirus of bat origin. Journal of Infectious Diseases 196:884–886. https://doi.org/10.1086/520817 Hernández-Aguilar I, Lorenzo C, Santos-Moreno A, et al (2021) Coronaviruses in bats: A review for the americas. Viruses 13 Johnson RI, Tachedjian M, Clayton BA, et al (2019) Characterization of teviot virus, an Australian bat-borne paramyxovirus. Journal of General Virology 100:403–413. https://doi.org/10.1099/jgv.0.001214 Markotter W, Geldenhuys M, Van Vuren PJ, et al (2019) Paramyxo- And coronaviruses in Rwandan bats. Trop Med Infect Dis 4:. https://doi.org/10.3390/tropicalmed4030099 Rizzo F, Edenborough KM, Toffoli R, et al (2017) Coronavirus and paramyxovirus in bats from Northwest Italy. BMC Vet Res 13:. https://doi.org/10.1186/s12917-017-1307-x Segovia H K, Salmon Mulanovich G, Ghersi BM, et al (2016) Prevalence of paramyxoviruses in bats in six areas of madre de dios and puno, Peru with two levels of anthropogenic disturbance. Revista de Investigaciones Veterinarias del Peru 27:241–251. https://doi.org/10.15381/rivep.v27i2.11640 Berto A, Anh PH, Carrique-Mas JJ, et al (2018) Detection of potentially novel paramyxovirus and coronavirus viral RNA in bats and rats in the Mekong Delta region of southern Viet Nam. Zoonoses Public Health 65:30–42. https://doi.org/10.1111/zph.12362 Tirira DG, Brito J, Burneo SF, et al (2021) ASOCIACIÓN ECUATORIANA DE MASTOZOOLOGÍA MAMÍFEROS DEL ECUADOR: LISTA OFICIAL ACTUALIZADA DE ESPECIES MAMMALS OF ECUADOR: OFFICIAL UPDATED SPECIES CHECKLIST ASOCIACIÓN ECUATORIANA DE MASTOZOOLOGÍA Poon LLM, Peiris JSM (2008) Detection of group 1 coronaviruses in bats using universal coronavirus reverse transcription polymerase chain reactions. Methods in Molecular Biology 454:13–26. https://doi.org/10.1007/978-1-59745-181-9_2 Tong S, Chern SWW, Li Y, et al (2008) Sensitive and broadly reactive reverse transcription-PCR assays to detect novel paramyxoviruses. J Clin Microbiol 46:2652–2658. https://doi.org/10.1128/JCM.00192-08 Vleut I, Levy-Tacher SI, Galindo-González J, et al (2012) Tropical rain-forest matrix quality affects bat assemblage structure in secondary forest patches. J Mammal 93:1469–1479. https://doi.org/10.1644/12-MAMM-A-005.1 Cabrera-Ojeda C, Noguera-Urbano EA, Calderón-Leytón JJ, Paí CF (2016) Ecología de murciélagos en el bosque seco tropical de Nariño (Colombia) y algunos comentarios sobre su conservación. Rev Peru Biol 23:27–34. https://doi.org/10.15381/rpb.v23i1.11830 Ruelas D (2017) Diferenciación morfológica de Carollia brevicauda y C. perspicillata (Chiroptera: Phyllostomidae) de Perú y Ecuador. Rev Peru Biol 24:363–382. https://doi.org/10.15381/rpb.v24i4.14063 Lou S, Yurrita CL (2005) Análisis de nicho alimentario en la comunidad de murciélagos frugívoros de Yaxhá, Petén, Guatemala. Acta Zool Mex 21:83–94 Burneo S, Tiria D (2015) PLAN DE ACCIÓN PARA LA CONSERVACIÓN DE LOS MURCIÉLAGOS DEL ECUADOR. Quito Asano KM, Hora AS, Scheffer KC, et al (2016) Alphacoronavirus in urban Molossidae and Phyllostomidae bats, Brazil. Virol J 13:. https://doi.org/10.1186/s12985-016-0569-4 Dos NR, Adriano R, Peracchi L, et al Morcegos do Brasil Woo PCY, Lau SKP, Li KSM, et al (2006) Molecular diversity of coronaviruses in bats. Virology 351:180–187. https://doi.org/10.1016/j.virol.2006.02.041 Darcissac E, Donato D, de Thoisy B, et al (2021) Paramyxovirus circulation in bat species from French Guiana. Infection, Genetics and Evolution 90:104769. https://doi.org/10.1016/j.meegid.2021.104769 Dominguez SR, O’Shea TJ, Oko LM, Holmes K V. (2007) Detection of group 1 coronaviruses in bats in North America. Emerg Infect Dis 13:1295–1300 Kuzmin I V., Bozick B, Guagliardo SA, et al (2011) Bats, emerging infectious diseases, and the rabies paradigm revisited. Emerg Health Threats J 4 Li L, Victoria JG, Wang C, et al (2010) Bat Guano Virome: Predominance of Dietary Viruses from Insects and Plants plus Novel Mammalian Viruses. J Virol 84:6955–6965. https://doi.org/10.1128/jvi.00501-10 Moreira-Soto A, Taylor-Castillo L, Vargas-Vargas N, et al (2015) Neotropical Bats from Costa Rica harbour Diverse Coronaviruses. Zoonoses Public Health 62:501–505. https://doi.org/10.1111/zph.12181 Kaufman EJ Two (or more) viruses in one bat: a systematic quantitative Two (or more) viruses in one bat: a systematic quantitative literature review of viral coinfection in bats literature review of viral coinfection in bats Heeney JL (2006) Zoonotic viral diseases and the frontier of early diagnosis, control and prevention. J Intern Med 260:399–408 Davy CM, Donaldson ME, Subudhi S, et al (2018) White-nose syndrome is associated with increased replication of a naturally persisting coronaviruses in bats. Sci Rep 8:. https://doi.org/10.1038/s41598-018-33975-x Wang J, Pan Y fei, Yang L fen, et al (2023) Individual bat virome analysis reveals co-infection and spillover among bats and virus zoonotic potential. Nat Commun 14:. https://doi.org/10.1038/s41467-023-39835-1 Tables Table 1. Number and species of bats found in the different habitats. SPECIES AGRICULTURAL ZONE FOREST REGION REFORESTATION SITE 1 Carollia perspicillata 18 10 2 Carollia brevicauda 2 10 16 3 Carollia castanea 7 4 Glossophaga soricina 1 2 5 Dermanura rosenbergi 2 1 2 6 Dermanura rava 2 25 4 7 Loncophyla concava 1 8 Artibeus lituratus 1 9 Artibeus aequatorialis 1 10 Sturnira luisi 1 2 11 Micronycteris megalotis 1 12 Vampyriscus nymphaea 4 13 Tonatia saurophila 1 14 Phylostomus discolor 2 15 Phyllostomus elongatus 1 16 Platyrrhinus vittatus 1 17 Vampyrum spectrum 1 18 Vampyressa thyone 1 19 Gardnerycteris crenulatum 2 20 Vampyrodes major 1 ABUNDANCE (IND) 10 78 35 RICHNESS (SP) 7 16 7 Table 2. Species found positive for coronavirus and/or paramyxovirus, together with the number of total samples collected for that species. Species Coronavirus positive samples/Total samples per species (%) Paramyxovirus positive samples/Total samples per species (%) Artibeus lituratus 1/1 (100%) 1/1 (100%) Carollia brevicauda 7/36(19.44%) 8/36 (22.22%) Carollia perspicillata 2/35(5.71%) 8/35(22.85) Dermanura rava 1/13 (7.69%) 4/13(30.76%) Dermanura rosenbergi 5/14(35.71%) 2/14(14.28%) Loncophylla concava 0/1 (0%) 1/1(100%) Platyrrhinus vittatus 0/1(0%) 1/1 (100%) Sturnira luisi 1/3(33.33%) 1/3(33.33%) Vampyrodes major 1/1(100%) 1/1(100%) Vampyriscus nymphaea 0/2(0%) 1/2(50%) Total 18/123 (14.63%) 28/123 (22.76%) Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-4030611","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":305818460,"identity":"8f3a60dc-3301-45e3-a37d-d2e618859136","order_by":0,"name":"Camila Acosta-López","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA1UlEQVRIiWNgGAWjYBADGX4IzUyshgQGHskGkrUYHCBWC3/74YcPf/6w4zG+kfzwAUOFdWIDe/MDvFokzqQZG/MkJPOY3UgzNmA4k57YwHPMAK8WA4YcNmmGBGaglhw2Cca2w4kNEjn4HWbA/4ZN8kdCPY/xjBz2H4z/gFrk3xDQIgE0nCfhMA+IwcDYALKFB78WiRvPgH5JO84jceaZsUTCsXTjNp40/H7h709++PCHTbUcf3vyww8faqxl+9kPP8BvDQpIAGI2EtSPglEwCkbBKMABAOhMPWB5t0qAAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0001-6555-4627","institution":"Universidad Central del Ecuador","correspondingAuthor":true,"prefix":"","firstName":"Camila","middleName":"","lastName":"Acosta-López","suffix":""},{"id":305818461,"identity":"ec2e99bc-9325-4123-b435-caa5afee99f8","order_by":1,"name":"Jonathan Bustamante","email":"","orcid":"","institution":"Universidad Central del Ecuador","correspondingAuthor":false,"prefix":"","firstName":"Jonathan","middleName":"","lastName":"Bustamante","suffix":""},{"id":305818462,"identity":"41b13c79-b69e-439f-b155-f95ac1d906b4","order_by":2,"name":"Maria Mercedes Gavilanez","email":"","orcid":"","institution":"Universidad Central del Ecuador","correspondingAuthor":false,"prefix":"","firstName":"Maria","middleName":"Mercedes","lastName":"Gavilanez","suffix":""}],"badges":[],"createdAt":"2024-03-07 18:28:13","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4030611/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4030611/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":57943558,"identity":"d5b4a764-bb34-4977-b607-bced0eceb3f3","added_by":"auto","created_at":"2024-06-07 19:06:20","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":91047,"visible":true,"origin":"","legend":"\u003cp\u003eMap of the sampling sites in FCAT. The dot in the Esmeraldas province shows the site where FCAT is located, dots in the map show the sampling sites, A = Agriculture, R = regeneration and F = Forest. All sites registered positive samples.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-4030611/v1/925495449b712defad69ce50.png"},{"id":57943664,"identity":"83f0f029-9fa4-4b71-9fef-665b82acc85f","added_by":"auto","created_at":"2024-06-07 19:06:35","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":467457,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4030611/v1/7547d082-170e-4a06-98a0-79a79bb4255b.pdf"}],"financialInterests":"","formattedTitle":"\u003cp\u003eMolecular Detection of Coronavirus and Paramyxovirus in Phyllostimid Bat Communities From Ecuador’s Northwestern Chocoan Forests\u003c/p\u003e","fulltext":[{"header":"Full Text","content":"\u003cp\u003eBats are classified as the fourth largest group of mammals globally, with their habitat spanning every continent except Antarctica\u0026nbsp;[1]. Ecuador is distinguished as the fourth richest country in bat biodiversity, hosting 171 different species\u0026nbsp;[2]. Serving as reservoirs for various pathogens, bats notably harbor coronaviruses and paramyxoviruses. About 16% of species carry coronaviruses \u0026mdash; evidenced by the discovery of 4,000 unique coronavirus sequences. These viruses are linked to significant zoonotic viruses, such as SARS-CoV, SARS-CoV2, and MERS-CoV\u0026nbsp;[3]. Paramyxoviruses, also widespread among bat populations, with 657 unique sequences identified, underline their role as progenitors of numerous diseases affecting humans and animals\u0026nbsp;[4].\u003c/p\u003e\n\u003cp\u003eThe surfacing of zoonotic viral diseases could be driven by factors such as human population growth, urbanization, and agricultural practices. These factors intensify human-animal interactions, elevating the likelihood of zoonosis\u0026nbsp;[5]. In 2002, the outbreak of SARS-CoV-1, originating from bats and transmitted through civet cats to humans, resulted in 8,098 infections and 774 fatalities, marking a mortality rate of 9.6%\u0026nbsp;[6]. Considering the recent pandemic, a consensus among epidemiologists\u0026rsquo; points to SARS-CoV-2 originating from a market trading wild animals for human consumption in Wuhan, China. The prevailing hypothesis is that bats, the initial hosts, transmitted the virus to pangolins, which in turn infected humans\u0026nbsp;[7, 8].\u003c/p\u003e\n\u003cp\u003eHuman-bat interactions can also enhance paramyxovirus transmission. A study in Cameroon found antibodies against paramyxoviruses, like Nipah and Hendra, in seven people\u0026nbsp;[9]. In Malaysia\u0026apos;s Tioman Island, the Tioman virus, a member of the Paramyxoviridae family affecting bats, pigs, and humans with flu-like symptoms, was studied. Antibodies against the Tioman virus were found in 5 of the 169 individuals tested, indicating its presence and potential health impact\u0026nbsp;[10]. While zoonotic diseases pose a substantial threat, Ecuador lacks data on them. A review covering 25 studies with 9,371 subjects highlighted Brazil, Mexico, US, Jamaica, Peru, Bolivia, and Argentina for coronavirus prevalence in bats, but Ecuador remains unstudied\u0026nbsp;[11]. A similar scene occurs for paramyxovirus, indicating a critical gap in understanding and managing potential zoonotic risks within the country\u0026nbsp;[12\u0026ndash;16]. This study\u0026apos;s objective was to assess the presence of these viruses in bats from the Fundaci\u0026oacute;n para la Conservaci\u0026oacute;n de los Andes Tropicales (FCAT) in Northwestern Ecuador.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eDuring November 2022, bats were systematically trapped through mist nets placed along known bat flight and feeding paths over ten nights at the FCAT Reserve in Esmeraldas province in three differently forested areas (Figure 1). Individuals were subsequently identified to species level with the Guide to the Mammals of Ecuador\u0026nbsp;[17]. A buccal swab sample was collected from all captured bats using sterile polystyrene swabs. Swabs were preserved in 500 uL of DNA Shield (Ecogen) until analysis.\u003c/p\u003e\n\u003cp\u003eFor viral RNA extraction the AccuPrep Dx Viral RNA Extraction kit (Bioneer) was used, following manufacturer\u0026apos;s instructions. A total of 33 samples were analyzed with a two-step RT-PCR and 90 of them with a one-step RT-PCR. For the reverse transcription was performed at 50\u0026deg;C for 15 minutes and 85\u0026deg;C for 5 minutes with the following protocol: 10\u0026nbsp;\u0026mu;L of nuclease-free water, 4\u0026nbsp;\u0026mu;L of 5X RT Buffer, 2\u0026nbsp;\u0026mu;L of RNA, and 1\u0026nbsp;\u0026mu;L of reverse transcriptase. Primers for the helicase gene were used for coronavirus while primers the RNA-dependent RNA polymerase gene (RdRp) were used for paramyxovirus.\u003c/p\u003e\n\u003cp\u003eFor the PCR reagents were added as follows:\u0026nbsp;10X Buffer, 100 pmol/\u0026mu;L of each primer, 1,5\u0026nbsp;\u0026mu;M MgCl2, 10\u0026mu;M dNTPs, 1\u0026nbsp;\u0026mu;L (1X) Fastgen Taq DNA polimerase and 2\u0026nbsp;\u0026mu;L of cDNA. For coronavirus\u0026nbsp;the PCR began with a 95\u0026deg;C denaturation \u0026ndash; 15 min, followed by 45 cycles of denaturation 95\u0026deg;C \u0026ndash; 1 min, 48\u0026deg;C \u0026ndash; 1 min and 72\u0026deg;C \u0026ndash; 1 min, with a final extension of 10 minutes\u0026nbsp;[18]. For paramyxovirus the PCR conditions were initial denaturation 60\u0026deg;C \u0026ndash; 15 min, 98\u0026deg;C denaturation \u0026ndash; 30 sec, followed by 40 cycles of denaturation 98\u0026deg;C \u0026ndash; 10 sec, 50\u0026deg;C \u0026ndash; 30 sec and 72\u0026deg;C for 30 sec, concluding with a 72\u0026deg;C extension of 7 minutes\u0026nbsp;[19].\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFor the 90 samples processed through one step RT-PCR; 25\u0026nbsp;\u0026mu;L of 2X One-Step RT-PCR Buffer, 4\u0026nbsp;\u0026mu;L of RT-PCR Enzyme mix, 2.5\u0026nbsp;\u0026mu;L of each primer (100pmol/\u0026mu;L), 5\u0026nbsp;\u0026mu;L of RNA, and 10\u0026nbsp;\u0026mu;L of nuclease-free water, were used. The temperature conditions were the same as those described previously for PCR. The resultant amplicon fragments were analyzed via 2% agarose gel electrophoresis. To determine the presence of coronavirus the expected fragment was 591 bp, whilst for paramyxovirus, the expected fragment size was 580 bp.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eIn the study, 123 bats spanning 20 species were captured (see Table 1).\u0026nbsp;Analysis revealed that 28 samples were positive for paramyxovirus, and 18 samples were positive for coronavirus. A total of 42 bats were infected with coronavirus and/or paramyxovirus, involving 10 species across 8 genera (see Table 2).\u003c/p\u003e\n\u003cp\u003eCoinfections were found in 4 individuals. These include: \u003cem\u003eCarollia perspicilata\u003c/em\u003e, \u003cem\u003eVampyrodes major\u003c/em\u003e, \u003cem\u003eArtibeus lituratus\u003c/em\u003e and \u003cem\u003eDermanura rosenbergi\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eIn examining the species captured, the forest zone boasted the greatest diversity and number of individuals, succeeded by the reforestation zone, and lastly, the agricultural zone (Table 1). This observation aligns with ecological principles. Research comparing bat biodiversity in untouched forest zones to that in fragmented areas revealed a direct correlation between the extent of forest coverage and the diversity and abundance of bat species\u0026nbsp;[20]. Moreover, research in Colombia evaluated the species composition and richness of bats in three different sites within Nari\u0026ntilde;o\u0026apos;s tropical dry forest. From a total of 60 bats, the highest individual count was observed in the preserved dry spiny scrub. In contrast, areas subjected to disturbance showed reduced species richness. This aligns with the outcomes of the current study\u0026nbsp;[21].\u003c/p\u003e\n\u003cp\u003eIn a study carried out in Peru, the species identified as most prevalent were \u003cem\u003eCarollia perspicillata\u003c/em\u003e, \u003cem\u003eCarollia brevicauda\u003c/em\u003e, and \u003cem\u003eArtibeus planirostris\u003c/em\u003e. Conversely, in our study, the most numerous species identified were \u003cem\u003eC. perspicillata\u003c/em\u003e, \u003cem\u003eC. brevicauda\u003c/em\u003e, and \u003cem\u003eDermanura rava\u003c/em\u003e. The overlap of two out of three predominant species in both studies can be attributed to their shared habitat preferences. \u003cem\u003eC. brevicauda\u003c/em\u003e and \u003cem\u003eC. perspicillata\u003c/em\u003e have a wide distribution, thriving in both the moist and arid forests located on the eastern and western slopes of the Andes encompassing a large altitudinal range\u0026nbsp;\u003cspan lang=\"EN-US\"\u003e[22]\u003c/span\u003e.\u0026nbsp;Moreover, these forest environments abound with key food sources for \u003cem\u003eC. perspicillata\u003c/em\u003e and \u003cem\u003eC. brevicauda\u003c/em\u003e [23]. Specifically, in northwestern Ecuador, within the FCAT region, bats consume \u003cem\u003ePiper\u003c/em\u003e spikes, which are prevalent in this area. The diet of bats in this part of Ecuador is diversified to include \u003cem\u003eCecropia concolor\u003c/em\u003e, \u003cem\u003eFicus eximia\u003c/em\u003e, \u003cem\u003ePiper aduncum\u003c/em\u003e, \u003cem\u003ePiper longistylosum\u003c/em\u003e, \u003cem\u003ePiper tuberculatum\u003c/em\u003e, and \u003cem\u003eSolanum sp\u003c/em\u003e. The widespread availability of these plant species ensures their presence in both the Peruvian study and this research. Furthermore, \u003cem\u003eCarollia\u003c/em\u003e bats play a crucial role as seed dispersers for these plants, establishing a mutualistic relationship\u0026nbsp;[24].\u003c/p\u003e\n\u003cp\u003eIn the context of coronavirus and paramyxovirus detection using PCR, a study in Sao Paulo, Brazil, collected fecal samples from bats in 73 municipalities to screen for coronavirus. From the 305 samples collected, 9 samples were identified as coronavirus positive from five species: \u003cem\u003eCynomops abrasus\u003c/em\u003e, \u003cem\u003eCynomops planirostris\u003c/em\u003e, \u003cem\u003eDesmodus rotundus\u003c/em\u003e, \u003cem\u003eGlossophaga soricina\u003c/em\u003e, and \u003cem\u003ePlatyrrhinus lineatus\u003c/em\u003e. In contrast, the coronavirus positive species in our study were \u003cem\u003eArtibeus lituratus\u003c/em\u003e, \u003cem\u003eC. brevicauda\u003c/em\u003e, \u003cem\u003eC. perspicillata\u003c/em\u003e, \u003cem\u003eDermanura rava\u003c/em\u003e, \u003cem\u003eDermanura rosenbergi\u003c/em\u003e, \u003cem\u003eVampyrodes major\u003c/em\u003e, and \u003cem\u003eSturnira luisi\u0026nbsp;\u003c/em\u003e(Table 2.). It is notable that the species identified as positive in our investigation do not overlap with those found in the Brazilian study\u0026nbsp;[25]. From the current study, two positive species are also distributed in the Sao Paulo area: \u003cem\u003eA. lituratus\u003c/em\u003e and \u003cem\u003eC. perspicillata\u003c/em\u003e [26]. However, these two species present in both regions, were not found to be infected in Brazil. This discrepancy can be attributed to environmental variations affecting each group, including differences in predator presence, trophic interactions, coinfections, and exposure to pathogens. As a result, the likelihood of identical bat species being infected with coronavirus across different regions is minimal\u0026nbsp;[27].\u003c/p\u003e\n\u003cp\u003eA separate investigation into paramyxovirus in French Guiana\u0026apos;s bat population found 103 individuals positive for the virus. The detection rates varied with the biological fluid examined; \u003cem\u003eD. rotundus\u003c/em\u003e, for example, tested positive for paramyxovirus only in kidney and urine samples, not in saliva, lung, or blood\u0026nbsp;[28]. Contrary to this, the current research identified positive paramyxovirus cases in saliva samples from \u003cem\u003eA. lituratus\u003c/em\u003e, \u003cem\u003eC. brevicauda\u003c/em\u003e, \u003cem\u003eC. perspicillata\u003c/em\u003e, \u003cem\u003eD. rava\u003c/em\u003e, \u003cem\u003eD. rosenbergi\u003c/em\u003e, \u003cem\u003eLonchophylla concava\u003c/em\u003e, \u003cem\u003eS. luisi\u003c/em\u003e, \u003cem\u003eV. major\u003c/em\u003e, \u003cem\u003ePlatyrrhinus vittatus,\u003c/em\u003e and \u003cem\u003eVampyriscus nymphaea\u003c/em\u003e. Similarly, a study focused on identifying coronavirus in North American bats through RT-PCR, revealed that all 22 saliva samples tested were negative. Conversely, 6 out of 28 fecal samples were positive for the virus\u0026nbsp;[29]. In comparison, the current study found coronavirus in 18 saliva samples from a total of 123 tested. This variation could be due to each virus\u0026apos;s specific replication and excretion mechanisms\u0026nbsp;[28]. Furthermore, in the second study the number of analyzed saliva samples was relatively low compared to the current research.\u003c/p\u003e\n\u003cp\u003eBats\u0026apos; social behavior, particularly their use of communal feeding areas, facilitates the transmission of different viruses among individuals and across colonies of the same species\u0026nbsp;[29]. These communal habits extend to resting sites, where bats also dispose of their waste. Since evidence suggests that bat feces may harbor paramyxovirus and coronavirus, it\u0026apos;s plausible to deduce that the communal lifestyle contributes to the spread of infections\u0026nbsp;[30]. Consequently, the species found in abundance in this study, \u003cem\u003eC. brevicauda\u003c/em\u003e, \u003cem\u003eC. perspicillata\u003c/em\u003e, \u003cem\u003eD. rosenbergi\u003c/em\u003e, and \u003cem\u003eD. rava\u003c/em\u003e, also exhibited the highest instances of coronavirus and paramyxovirus infections.\u003c/p\u003e\n\u003cp\u003ePrevalence of coronavirus was 14.63%, and 22.76% for paramyxovirus. These values are considered high in comparison with other studies, where significantly lower values were recorded with larger sample sizes. For example, a study conducted in Italy, analyzing bat saliva and feces, found 12% and 1% prevalence for coronavirus and paramyxovirus respectively, out of 302 captured bats\u0026nbsp;[14]. In Costa Rica, from 421 fecal samples, only 4 of them were positive for coronavirus\u0026nbsp;[31].\u0026nbsp;Similarly, in Peru, from 436 monitored individuals, a prevalence of 10.3% was recorded for paramyxovirus\u0026nbsp;[15].\u003c/p\u003e\n\u003cp\u003eDue to these high values, meticulous epidemiological surveillance must be maintained in bats to detect coronavirus, paramyxovirus, and other viruses in general. This is crucial for several reasons, one of which being the variety of viruses harbored by bats. Some of the other viral families found include adenovirus, astrovirus, bornavirus, circovirus, herpesvirus, parvovirus, polyomavirus, and rhabdovirus\u0026nbsp;[32]. Another crucial reason is that it could allow us to identify potential sites of zoonotic outbreaks. This enables the possibility for taking preemptive measures to prevent the spread of such diseases\u0026nbsp;[33].\u003c/p\u003e\n\u003cp\u003eA total of 4 coinfections were recorded. A systematic review noted that out of 725 bats with coinfections, only 4 individuals had a coinfection with only two viruses, namely paramyxovirus and coronavirus. These findings imply that the prevalence of coinfections in the current study is also high, in relation to the number of samples\u0026nbsp;[32]. This phenomenon could be attributed to peak viral proliferation phases, resulting in higher viral shedding\u0026nbsp;[34]. Additionally, it might be explained by the possibility that the viral strains infecting these individuals possess molecular mechanisms that hinder the action of interferons, which are crucial for bats in regulating viral replication\u0026nbsp;[29]. The likelihood of coinfections is also affected by the phylogenetic relationships among bat species. The ability of a virus to infect is contingent upon overcoming physiological and molecular defenses. Viruses are more likely to infect two bat species that are phylogenetically close, as they may share similar defenses. For example, the SARS CX1 coronavirus is evolutionarily akin to SARS-CoV-2, and both can infect identical bat species because the receptor-binding domain of their spike proteins only differs by five amino acids\u0026nbsp;[35].\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn this study, the bulk of positive cases were identified in \u003cem\u003eC. brevicauda\u003c/em\u003e with 15 positive samples, followed by \u003cem\u003eC. perspicillata\u003c/em\u003e with 10, and \u003cem\u003eD. rosenbergi\u003c/em\u003e with 7 positive samples. Notably, among these, two cases of coinfections were detected: one in \u003cem\u003eC. perspicillata\u003c/em\u003e and another in \u003cem\u003eD. rosenbergi\u003c/em\u003e. This observation of coinfections may be attributed to the dense populations of the species, as well as to the phylogenetic distance between the mentioned species [35]. Findings of this study highlight a significant prevalence of coronavirus and paramyxovirus among the bat populations examined. This marks the first documentation in Ecuador of various bat species harboring infections of coronavirus and paramyxovirus. Given these findings, it underscores the imperative need for further research within Ecuador to track the viral diversity of coronavirus and paramyxovirus, particularly in regions proximate to human communities.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eFUNDING\u003c/p\u003e\n\u003cp\u003eThis work was supported by Direcci\u0026oacute;n de Investigaci\u0026oacute;n, Universidad Central del Ecuador seed grant cifFuce-cv-fcb-1.\u003c/p\u003e\n\u003cp\u003eCOMPETING INTERESTS\u003c/p\u003e\n\u003cp\u003eThe authors have no relevant financial or non-financial interests to disclose.\u003c/p\u003e\n\u003cp\u003eAUTHOR CONTRIBUTIONS\u003c/p\u003e\n\u003cp\u003eCamila Acosta-L\u0026oacute;pez and Mar\u0026iacute;a Mercedes Gavilanez contributed to the study conception and design. Material preparation and data collection were performed by Jonathan Bustamante. Data analyses were performed by Camila Acosta-L\u0026oacute;pez, Mar\u0026iacute;a Mercedes Gavilanez and Jonathan Bustamante. The first draft of the manuscript was written by Jonathan Bustamante, all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003eDATA AVAILABILITY\u003c/p\u003e\n\u003cp\u003eThe datasets generated and analyzed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e\n\u003cp\u003eETHICS APPROVAL\u003c/p\u003e\n\u003cp\u003eThis study was performed in line with the criteria stablished in the manual for recommendations for the use and handling of wild mammals. Since the samples were collected through noninvasive techniques, no administration of pain killers or chemical agents was needed. Approval was granted by the Comisi\u0026oacute;n de Investigaci\u0026oacute;n de la Facultad de Medicina Veterinaria y Zootecnia from Universidad Central del Ecuador.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eBanerjee A, Kulcsar K, Misra V, et al (2019) Bats and coronaviruses. Viruses 11:7\u0026ndash;9. https://doi.org/10.3390/v11010041\u003c/li\u003e\n\u003cli\u003eM\u0026eacute;ndez-Rodr\u0026iacute;guez A, Horta P, Zarza H, et al (2024) Surveying Bat-Hosted Adenoviruses and Herpesviruses: A Comprehensive Analysis. Diversity (Basel) 16\u003c/li\u003e\n\u003cli\u003eAnthony SJ, Johnson CK, Greig DJ, et al (2017) Global patterns in coronavirus diversity. Virus Evol 3:. https://doi.org/10.1093/ve/vex012\u003c/li\u003e\n\u003cli\u003eHayman DTS (2016) Bats as Viral Reservoirs. Annu Rev Virol 3:77\u0026ndash;99\u003c/li\u003e\n\u003cli\u003eMcmahon B, Morand S, Gray J (2018) Ecosystem change and zoonoses in the Anthropocene. Zoonoses Public Health 755\u0026ndash;765\u003c/li\u003e\n\u003cli\u003eWang J-T, Chang S-C (2004) Severe acute respiratory syndrome. Curr Opin Infect Dis 17:143\u0026ndash;148. https://doi.org/10.1097/00001432-200404000-00013\u003c/li\u003e\n\u003cli\u003eLau SKP, Luk HKH, Wong ACP, et al (2020) Possible Bat Origin of Severe Acute Respiratory Syndrome Coronavirus 2. Emerg Infect Dis 26:1542\u0026ndash;1547. https://doi.org/10.3201/eid2607.200092\u003c/li\u003e\n\u003cli\u003eXiao K, Zhai J, Feng Y, et al (2020) Isolation of SARS-CoV-2-related coronavirus from Malayan pangolins. Nature 583:286\u0026ndash;289. https://doi.org/10.1038/s41586-020-2313-x\u003c/li\u003e\n\u003cli\u003eThibault PA, Watkinson RE, Moreira-Soto A, et al (2017) Zoonotic Potential of Emerging Paramyxoviruses. In: Advances in Virus Research. pp 1\u0026ndash;55\u003c/li\u003e\n\u003cli\u003eKoon CY, Crameri G, Wang L, et al (2007) Serological evidence of possible human infection with Tioman virus, a newly described paramyxovirus of bat origin. Journal of Infectious Diseases 196:884\u0026ndash;886. https://doi.org/10.1086/520817\u003c/li\u003e\n\u003cli\u003eHern\u0026aacute;ndez-Aguilar I, Lorenzo C, Santos-Moreno A, et al (2021) Coronaviruses in bats: A review for the americas. Viruses 13\u003c/li\u003e\n\u003cli\u003eJohnson RI, Tachedjian M, Clayton BA, et al (2019) Characterization of teviot virus, an Australian bat-borne paramyxovirus. Journal of General Virology 100:403\u0026ndash;413. https://doi.org/10.1099/jgv.0.001214\u003c/li\u003e\n\u003cli\u003eMarkotter W, Geldenhuys M, Van Vuren PJ, et al (2019) Paramyxo- And coronaviruses in Rwandan bats. Trop Med Infect Dis 4:. https://doi.org/10.3390/tropicalmed4030099\u003c/li\u003e\n\u003cli\u003eRizzo F, Edenborough KM, Toffoli R, et al (2017) Coronavirus and paramyxovirus in bats from Northwest Italy. BMC Vet Res 13:. https://doi.org/10.1186/s12917-017-1307-x\u003c/li\u003e\n\u003cli\u003eSegovia H K, Salmon Mulanovich G, Ghersi BM, et al (2016) Prevalence of paramyxoviruses in bats in six areas of madre de dios and puno, Peru with two levels of anthropogenic disturbance. Revista de Investigaciones Veterinarias del Peru 27:241\u0026ndash;251. https://doi.org/10.15381/rivep.v27i2.11640\u003c/li\u003e\n\u003cli\u003eBerto A, Anh PH, Carrique-Mas JJ, et al (2018) Detection of potentially novel paramyxovirus and coronavirus viral RNA in bats and rats in the Mekong Delta region of southern Viet Nam. Zoonoses Public Health 65:30\u0026ndash;42. https://doi.org/10.1111/zph.12362\u003c/li\u003e\n\u003cli\u003eTirira DG, Brito J, Burneo SF, et al (2021) ASOCIACI\u0026Oacute;N ECUATORIANA DE MASTOZOOLOG\u0026Iacute;A MAM\u0026Iacute;FEROS DEL ECUADOR: LISTA OFICIAL ACTUALIZADA DE ESPECIES MAMMALS OF ECUADOR: OFFICIAL UPDATED SPECIES CHECKLIST ASOCIACI\u0026Oacute;N ECUATORIANA DE MASTOZOOLOG\u0026Iacute;A\u003c/li\u003e\n\u003cli\u003ePoon LLM, Peiris JSM (2008) Detection of group 1 coronaviruses in bats using universal coronavirus reverse transcription polymerase chain reactions. Methods in Molecular Biology 454:13\u0026ndash;26. https://doi.org/10.1007/978-1-59745-181-9_2\u003c/li\u003e\n\u003cli\u003eTong S, Chern SWW, Li Y, et al (2008) Sensitive and broadly reactive reverse transcription-PCR assays to detect novel paramyxoviruses. J Clin Microbiol 46:2652\u0026ndash;2658. https://doi.org/10.1128/JCM.00192-08\u003c/li\u003e\n\u003cli\u003eVleut I, Levy-Tacher SI, Galindo-Gonz\u0026aacute;lez J, et al (2012) Tropical rain-forest matrix quality affects bat assemblage structure in secondary forest patches. J Mammal 93:1469\u0026ndash;1479. https://doi.org/10.1644/12-MAMM-A-005.1\u003c/li\u003e\n\u003cli\u003eCabrera-Ojeda C, Noguera-Urbano EA, Calder\u0026oacute;n-Leyt\u0026oacute;n JJ, Pa\u0026iacute; CF (2016) Ecolog\u0026iacute;a de murci\u0026eacute;lagos en el bosque seco tropical de Nari\u0026ntilde;o (Colombia) y algunos comentarios sobre su conservaci\u0026oacute;n. Rev Peru Biol 23:27\u0026ndash;34. https://doi.org/10.15381/rpb.v23i1.11830\u003c/li\u003e\n\u003cli\u003eRuelas D (2017) Diferenciaci\u0026oacute;n morfol\u0026oacute;gica de Carollia brevicauda y C. perspicillata (Chiroptera: Phyllostomidae) de Per\u0026uacute; y Ecuador. Rev Peru Biol 24:363\u0026ndash;382. https://doi.org/10.15381/rpb.v24i4.14063\u003c/li\u003e\n\u003cli\u003eLou S, Yurrita CL (2005) An\u0026aacute;lisis de nicho alimentario en la comunidad de murci\u0026eacute;lagos frug\u0026iacute;voros de Yaxh\u0026aacute;, Pet\u0026eacute;n, Guatemala. Acta Zool Mex 21:83\u0026ndash;94\u003c/li\u003e\n\u003cli\u003eBurneo S, Tiria D (2015) PLAN DE ACCI\u0026Oacute;N PARA LA CONSERVACI\u0026Oacute;N DE LOS MURCI\u0026Eacute;LAGOS DEL ECUADOR. Quito\u003c/li\u003e\n\u003cli\u003eAsano KM, Hora AS, Scheffer KC, et al (2016) Alphacoronavirus in urban Molossidae and Phyllostomidae bats, Brazil. Virol J 13:. https://doi.org/10.1186/s12985-016-0569-4\u003c/li\u003e\n\u003cli\u003eDos NR, Adriano R, Peracchi L, et al Morcegos do Brasil\u003c/li\u003e\n\u003cli\u003eWoo PCY, Lau SKP, Li KSM, et al (2006) Molecular diversity of coronaviruses in bats. Virology 351:180\u0026ndash;187. https://doi.org/10.1016/j.virol.2006.02.041\u003c/li\u003e\n\u003cli\u003eDarcissac E, Donato D, de Thoisy B, et al (2021) Paramyxovirus circulation in bat species from French Guiana. Infection, Genetics and Evolution 90:104769. https://doi.org/10.1016/j.meegid.2021.104769\u003c/li\u003e\n\u003cli\u003eDominguez SR, O\u0026rsquo;Shea TJ, Oko LM, Holmes K V. (2007) Detection of group 1 coronaviruses in bats in North America. Emerg Infect Dis 13:1295\u0026ndash;1300\u003c/li\u003e\n\u003cli\u003eKuzmin I V., Bozick B, Guagliardo SA, et al (2011) Bats, emerging infectious diseases, and the rabies paradigm revisited. Emerg Health Threats J 4\u003c/li\u003e\n\u003cli\u003eLi L, Victoria JG, Wang C, et al (2010) Bat Guano Virome: Predominance of Dietary Viruses from Insects and Plants plus Novel Mammalian Viruses. J Virol 84:6955\u0026ndash;6965. https://doi.org/10.1128/jvi.00501-10\u003c/li\u003e\n\u003cli\u003eMoreira-Soto A, Taylor-Castillo L, Vargas-Vargas N, et al (2015) Neotropical Bats from Costa Rica harbour Diverse Coronaviruses. Zoonoses Public Health 62:501\u0026ndash;505. https://doi.org/10.1111/zph.12181\u003c/li\u003e\n\u003cli\u003eKaufman EJ Two (or more) viruses in one bat: a systematic quantitative Two (or more) viruses in one bat: a systematic quantitative literature review of viral coinfection in bats literature review of viral coinfection in bats\u003c/li\u003e\n\u003cli\u003eHeeney JL (2006) Zoonotic viral diseases and the frontier of early diagnosis, control and prevention. J Intern Med 260:399\u0026ndash;408\u003c/li\u003e\n\u003cli\u003eDavy CM, Donaldson ME, Subudhi S, et al (2018) White-nose syndrome is associated with increased replication of a naturally persisting coronaviruses in bats. Sci Rep 8:. https://doi.org/10.1038/s41598-018-33975-x\u003c/li\u003e\n\u003cli\u003eWang J, Pan Y fei, Yang L fen, et al (2023) Individual bat virome analysis reveals co-infection and spillover among bats and virus zoonotic potential. Nat Commun 14:. https://doi.org/10.1038/s41467-023-39835-1\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003e\u003cstrong\u003eTable 1.\u003c/strong\u003e Number and species of bats found in the different habitats.\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"100%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eSPECIES\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eAGRICULTURAL\u003c/strong\u003e \u003cstrong\u003eZONE\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eFOREST REGION\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eREFORESTATION SITE\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e1\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eCarollia perspicillata\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e2\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eCarollia brevicauda\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e3\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eCarollia castanea\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e4\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eGlossophaga soricina\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e5\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eDermanura rosenbergi\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e6\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eDermanura rava\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e7\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eLoncophyla concava\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e8\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eArtibeus lituratus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e9\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eArtibeus aequatorialis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e10\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eSturnira luisi\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.122448979591836%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e11\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"55.10204081632653%\" colspan=\"2\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eMicronycteris megalotis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.26530612244898%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"25.510204081632654%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e12\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eVampyriscus nymphaea\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e13\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eTonatia saurophila\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e14\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePhylostomus discolor\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e15\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePhyllostomus elongatus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e16\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePlatyrrhinus vittatus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e17\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eVampyrum spectrum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e18\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eVampyressa thyone\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.122448979591836%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e19\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"55.10204081632653%\" colspan=\"2\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eGardnerycteris crenulatum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.26530612244898%\" valign=\"top\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.510204081632654%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e20\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eVampyrodes major\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eABUNDANCE (IND)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e78\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\n \u003cp\u003e35\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"6.185567010309279%\" valign=\"top\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd width=\"30.927835051546392%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eRICHNESS (SP)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"23.711340206185568%\" valign=\"top\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"13.402061855670103%\" valign=\"top\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"25.77319587628866%\" valign=\"top\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eTable 2.\u0026nbsp;Species found positive for coronavirus and/or paramyxovirus, together with the number of total samples collected for that species.\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"100%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003eSpecies\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eCoronavirus positive samples/Total samples per species (%)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eParamyxovirus positive samples/Total samples per species (%)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eArtibeus lituratus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e1/1 (100%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e1/1 (100%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eCarollia brevicauda\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e7/36(19.44%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e8/36 (22.22%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eCarollia perspicillata\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e2/35(5.71%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e8/35(22.85)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eDermanura rava\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e1/13 (7.69%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e4/13(30.76%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eDermanura rosenbergi\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e5/14(35.71%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e2/14(14.28%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eLoncophylla concava\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e0/1 (0%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e1/1(100%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003ePlatyrrhinus vittatus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e0/1(0%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e1/1 (100%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eSturnira luisi\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e1/3(33.33%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e1/3(33.33%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eVampyrodes major\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e1/1(100%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e1/1(100%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cem\u003eVampyriscus nymphaea\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e0/2(0%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e1/2(50%)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd width=\"24.242424242424242%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003eTotal\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"39.39393939393939%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e18/123 (14.63%)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd width=\"36.36363636363637%\" valign=\"top\"\u003e\n \u003cp\u003e\u003cstrong\u003e28/123 (22.76%)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"
[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"","lastPublishedDoi":"10.21203/rs.3.rs-4030611/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4030611/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"Bats are important reservoirs for pathogenic viruses. Amidst rising human-wildlife interactions, the risk of potential spillover of zoonotic diseases escalates. We aimed to identify Coronavirus and Paramyxovirus in bat saliva samples for species inhabiting Ecuador’s northwestern Choco region. Sampling was undertaken at FCAT reserve in the Esmeraldas province in November 2022. A total of 123 samples from individuals appertaining 20 species, were analyzed using RT-PCR. We recorded 28 positive cases for paramyxovirus, 18 for coronavirus and four co-infections. These findings underscore the necessity of enhancing zoonotic surveillance and establishing viral diversity monitoring strategies.","manuscriptTitle":"Molecular Detection of Coronavirus and Paramyxovirus in Phyllostimid Bat Communities From Ecuador’s Northwestern Chocoan Forests","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-06-07 19:05:38","doi":"10.21203/rs.3.rs-4030611/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"
[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"85666d47-ce26-45e1-b9ee-fb54613db1b2","owner":[],"postedDate":"June 7th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2024-06-07T19:05:47+00:00","versionOfRecord":[],"versionCreatedAt":"2024-06-07 19:05:38","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-4030611","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4030611","identity":"rs-4030611","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}
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