Vol. 21 No. 1 (2026)
Articles

Detection of CMTV-like ranavirus following a Rana temporaria mass mortality event in a northern Italian alpine lake

Matteo Riccardo Di Nicola
Istituto Zooprofilattico Sperimentale del Piemonte, Liguria e Valle d’Aosta, Via Bologna 148, 10154 Torino, Italy
Chiara Beltramo
Istituto Zooprofilattico Sperimentale del Piemonte, Liguria e Valle d’Aosta, Via Bologna 148, 10154 Torino, Italy
Paolo Pastorino
Istituto Zooprofilattico Sperimentale del Piemonte, Liguria e Valle d’Aosta, Via Bologna 148, 10154 Torino, Italy
Giuseppe Esposito
Istituto Zooprofilattico Sperimentale del Piemonte, Liguria e Valle d’Aosta, Via Bologna 148, 10154 Torino, Italy
Anna Cerullo
Istituto Zooprofilattico Sperimentale del Piemonte, Liguria e Valle d’Aosta, Via Bologna 148, 10154 Torino, Italy
Arianna Meletiadis
Istituto Zooprofilattico Sperimentale del Piemonte, Liguria e Valle d’Aosta, Via Bologna 148, 10154 Torino, Italy
Marino Prearo
Istituto Zooprofilattico Sperimentale del Piemonte, Liguria e Valle d’Aosta, Via Bologna 148, 10154 Torino, Italy
Simone Peletto
Istituto Zooprofilattico Sperimentale del Piemonte, Liguria e Valle d’Aosta, Via Bologna 148, 10154 Torino, Italy
Pier Luigi Acutis
Istituto Zooprofilattico Sperimentale del Piemonte, Liguria e Valle d’Aosta, Via Bologna 148, 10154 Torino, Italy

Published 2026-02-09

Keywords

  • Acinetobacter,
  • CMTV,
  • Cottian Alps,
  • Disease ecology,
  • Emerging infectious diseases,
  • Germanasca Valley,
  • Outbreak,
  • Ranavirus
  • ...More
    Less

How to Cite

Di Nicola, M. R., Beltramo, C., Pastorino, P., Esposito, G., Cerullo, A., Meletiadis, A., Prearo, M., Peletto, S., & Acutis, P. L. (2026). Detection of CMTV-like ranavirus following a Rana temporaria mass mortality event in a northern Italian alpine lake. Acta Herpetologica, 21(1), 49–62. https://doi.org/10.36253/a_h-18382

Abstract

High-mountain lakes are vulnerable to climatic and anthropogenic stressors, and infectious disease may further exacerbate impacts on alpine communities, particularly during seasonal temperature peaks. In August 2024, a mass mortality event of common frogs (Rana temporaria) occurred in a high-altitude lake in the Cottian Alps (Piedmont, Italy). During a one-hour survey, 78 dead frogs were recorded; nine carcasses (all adult males) were sampled for diagnostic tests. Six showed ventral red discolouration and three of them had ulcerative lesions of the digits. Eight out of nine vitreous humour samples were culture-positive, with isolates including Hafnia alvei, Acinetobacter guillouiae, Acinetobacter proteolyticus, and Serratia proteamaculans. PCR screening of skin and pooled organs detected ranavirus in four out of nine frogs, while Batrachochytrium dendrobatidis and herpesvirus tested negative. Phylogenetic analysis of sequenced major capsid protein and DNA polymerase fragments grouped the virus within the CMTV-like clade, with high similarity to reference sequences. This represents the first geolocated detection of a CMTV-like ranavirus in free-ranging amphibians in Italy. Although the advanced state of decomposition precluded histopathological evaluation and causality cannot be conclusively established, the concordance between molecular detection and gross lesions consistent with ranaviral infection supports a plausible role of ranavirus in the observed die-off. Our findings highlight the need for targeted surveillance in Italy’s alpine amphibians, including environmental DNA sampling and screening of non-native fish. Given ecological simplification and short reproductive seasons at high altitude, longitudinal monitoring is advisable to assess persistence, seasonality and potential spillover across life stages and sympatric species.

References

  1. Akçakaya, H.R., Neam, K., Hobin, L., Lötters, S., Martel, A., Pasmans, F. (2023): Assessing the extinction risks of amphibians impacted by infectious diseases. Biol. Conserv. 284: 110205.
  2. Allain, S.J., Duffus, A.L. (2019): Emerging infectious disease threats to European herpetofauna. Herpetol. J. 29: 189-206.
  3. Allasia, P., Audisio, C., Baldo, M., Cirio, C.G., Lollino, G., Giordan, D., Godone, F., Nigrelli, G., Alpe, F., Ambrogio, S., Giardino, M., Perotti, L., Sambuelli, L., De Renzo, G., Fontan, D., Barbero, T. (2004): Instrumented experimental sites for the control of landslide hazards in mountain environments: the Germanasca and Susa Valleys (Northwestern Italy). In: Field trip guidebooks, 32nd International Geological Congress, Florence, Italy, 20-28 August 2004, pp. 37-54. APAT, Roma.
  4. Aree Protette Alpi Marittime (2019): Ranavirus a rischio la rana temporaria. Aree Protette Alpi Marittime.
  5. Ariel, E., Holopainen, R., Olesen, N.J., Tapiovaara, H. (2010): Comparative study of ranavirus isolates from cod (Gadus morhua) and turbot (Psetta maxima) with reference to other ranaviruses. Arch. Virol. 155: 1261-1271.
  6. Ariel, E., Subramaniam, K., Imnoi, K., Sriwanayos, P., Ahasan, M.S., Olesen, N.J., Amedeo, M., Toffan, A., Waltzek, T.B. (2017): Genomic sequencing of Ranaviruses isolated from edible frogs (Pelophylax esculentus). Genome Announc. 5: 10-1128.
  7. Balseiro, A., Dalton, K., Del Cerro, A., Marquez, I., Cunningham, A., Parra, F., Prieto, J., Casais, R. (2009): Pathology, isolation and molecular characterisation of a Ranavirus from the common midwife toad Alytes obstetricans on the Iberian Peninsula. Dis. Aquat. Org. 84: 95-104.
  8. Bates, K.A., Rosa, G.M., Gamer, T.W.J. (2025): Ranavirus. Trends Microbiol. 33: 701-702.
  9. Bergò, P.E., Andreone, F. (2002): A terrestrial viviparous salamander into water: Notes on the unusual larval aquatic development in Salamandra lanzai. Herpetol. Bull. 81: 29-31.
  10. Bianchessi, L., Flach, E., Monacchia, G., Dagleish, M., Maley, M., Turin, L., Rocchi, M.S. (2024): Identification and characterisation of gamma-herpesviruses in zoo artiodactyla. Virol. J. 21: 1-15.
  11. Bison, M., Yoccoz, N.G., Carlson, B.Z., Klein, G., Laigle, I., Van Reeth, C., Delestrade, A. (2021): Earlier snowmelt advances breeding phenology of the common frog (Rana temporaria) but increases the risk of frost exposure and wetland drying. Front. Ecol. Evol. 9: 1-12.
  12. Bosch, J., Carrascal, L.M., Durán, L., Walker, S., Fisher, M.C. (2007): Climate change and outbreaks of amphibian chytridiomycosis in a montane area of Central Spain: Is there a link? Proc. R. Soc. B. 274: 253-260.
  13. Bosch, J., Monsalve-Carcaño, C., Price, S.J., Bielby, J. (2020): Single infection with Batrachochytrium dendrobatidis or Ranavirus does not increase probability of co-infection in a montane community of amphibians. Sci. Rep. 10: 21115.
  14. Bosch, J., Mora-Cabello De Alba, A., Marquínez, S., Price, S.J., Thumsová, B., Bielby, J. (2021): Long-term monitoring of amphibian populations of a national park in northern Spain reveals negative persisting effects of Ranavirus, but not Batrachochytrium dendrobatidis. Front. Vet. Sci. 8: 645491.
  15. Box, E.K., Cleveland, C.A., Subramaniam, K., Waltzek, T.B., Yabsley, M.J. (2021): Molecular confirmation of Ranavirus infection in amphibians from Chad, Africa. Front. Vet. Sci. 8: 733939.
  16. Boyle, D., Boyle, D., Olsen, V., Morgan, J., Hyatt, A. (2004): Rapid quantitative detection of chytridiomycosis (Batrachochytrium dendrobatidis) in amphibian samples using real-time Taqman PCR assay. Dis. Aquat. Org. 60: 141-148.
  17. Brunner, J.L., Olson, D.H., Gray, M.J., Miller, D.L., Duffus, A.L.J. (2021): Global patterns of Ranavirus detections. FACETS 6: 912-924.
  18. Catalan, J., Ninot, J.M., Aniz, M.M. (2017): High mountain conservation in a changing world. Cham, Springer International Publishing.
  19. Claytor, S.C., Subramaniam, K., Landrau-Giovannetti, N., Chinchar, V.G., Gray, M.J., Miller, D.L., Mavian, C., Salemi, M., Wisely, S., Waltzek, T.B. (2017): Ranavirus phylogenomics: Signatures of recombination and inversions among bullfrog ranaculture isolates. Virology 511: 330-343.
  20. Di Nicola, M.R., Cavigioli, L., Luiselli, L., Andreone, F. (2021): Anfibi & Rettili d’Italia. Edizione aggiornata. Latina; Edizioni Belvedere.
  21. Di Nicola, M.R., Rubiola, S., Cerullo, A., Basciu, A., Massone, C., Zabbia, T., Dorne, J.L.M.C., Acutis, P.L., Marini, D. (2025): Microorganisms in wild European reptiles: Bridging gaps in neglected conditions to inform disease ecology research. Int. J. Parasitol. Parasites. Wildl. 27: 101113.
  22. Duffus, A.L., Nichols, R.A., Garner, T.W. (2014): Detection of a frog virus 3-like Ranavirus in native and introduced amphibians in the United Kingdom in 2007 and 2008. Herpetol. Rev. 45: 608-610.
  23. Ehlers, B., Borchers, K., Grund, C., Frölich, K., Ludwig, H., Buhk, H.-J. (1999): Detection of new DNA polymerase genes of known and potentially novel herpesviruses by PCR with degenerate and deoxyinosine-substituted primers. Virus Genes 18: 211-220.
  24. Farina, T., Ed. (2008): Alpi Cozie. Supplemento a Piemonte Parchi 5(XXIII). Torino, Regione Piemonte.
  25. Fisher, M.C., Garner, T.W.J. (2020): Chytrid fungi and global amphibian declines. Nat. Rev. Microbiol. 18: 332-343.
  26. Fisher, M.C., Henk, D.A., Briggs, C.J., Brownstein, J.S., Madoff, L.C., McCraw, S.L., Gurr, S.J. (2012): Emerging fungal threats to animal, plant and ecosystem health. Nature 484: 186-194.
  27. Flechas, S., Urbina, J., Crawford, A., Gutiérrez, K., Corrales, K., Castellanos, L., González, M., Cuervo, A., Catenazzi, A. (2023): First evidence of ranavirus in native and invasive amphibians in Colombia. Dis. Aquat. Org. 153: 51-58.
  28. Forno, M.G., Lingua, A., Lo Russo, S., Taddia, G. (2011): Improving digital tools for Quaternary field survey: A case study of the Rodoretto Valley (NW Italy). Environ. Earth. Sci. 64: 1487-1495.
  29. Forzán, M.J., Jones, K.M., Ariel, E., Whittington, R.J., Wood, J., Markham, R.J.F., Daoust, P.-Y. (2017): Pathogenesis of frog virus 3 (Ranavirus, Iridoviridae) infection in wood frogs (Rana sylvatica). Vet. Pathol. 54: 531-548.
  30. Franklinos, L., Fernandez, J., Hydeskov, H., Hopkins, K., Everest, D., Cunningham, A., Lawson, B. (2018): Herpesvirus skin disease in free-living common frogs Rana temporaria in Great Britain. Dis. Aquat. Org. 129: 239-244.
  31. Guo, L., Jin, X., Yang, D., Wei, L., Chen, J., Lin, Z., Ma, L. (2025): Identification and characterization of Serratia nematophila and Acinetobacter guillouiae from putrid-skin disease lesions in farmed Chinese spiny frog (Quasipaa spinosa). Microbiol. Spectr. 13: e02096-24.
  32. Hanna, P.E., Bellamy, J.E., Donald, A. (1990): Postmortem eyefluid analysis in dogs, cats and cattle as an estimate of antemortem serum chemistry profiles. Can. J. Vet. Res. 54: 487-494.
  33. Hartmann, A.M., Maddox, M.L., Ossiboff, R.J., Longo, A.V. (2022): Sustained Ranavirus outbreak causes mass mortality and morbidity of imperiled amphibians in Florida. EcoHealth 19: 8-14.
  34. Hick, P.M., Subramaniam, K., Thompson, P., Whittington, R.J., Waltzek, T.B. (2016): Complete genome sequence of a Bohle iridovirus isolate from ornate burrowing frogs (Limnodynastes ornatus) in Australia. Genome Announc. 4: e00632-16.
  35. Herath, J., Sun, D., Ellepola, G., Subramaniam, K., Meegaskumbura, M. (2023): Emerging threat of Ranavirus: Prevalence, genetic diversity, and climatic drivers of Ranavirus (Iridoviridae) in ectothermic vertebrates of Asia. Front. Vet. Sci. 10: 1291872.
  36. Hoberg, E.P., Brooks, D.R. (2015): Evolution in action: Climate change, biodiversity dynamics and emerging infectious disease. Philos. Trans. R. Soc. B 370: 20130553.
  37. Holopainen, R., Ohlemeyer, S., Schütze, H., Bergmann, S., Tapiovaara, H. (2009): Ranavirus phylogeny and differentiation based on major capsid protein, DNA polymerase and neurofilament triplet H1-like protein genes. Dis. Aquat. Org. 85: 81-91.
  38. Hyatt, A.D., Gould, A.R., Zupanovic, Z., Cunningham, A.A., Hengstberger, S., Whittington, R.J., Kattenbelt, J., Coupar, B.E.H. (2000): Comparative studies of piscine and amphibian iridoviruses. Arch. Virol. 145: 301-331.
  39. Ilić, M., Bugarski-Stanojević, V., Jovanović, B., Stamenković, G., Zorić, K., Paunović, M., Crnobrnja-Isailović, J. (2024): Phylogeographic substructuring in the southernmost refugium of the European common frog Rana temporaria. Animals 14: 1430.
  40. Ionescu, M.I., Neagoe, D. Ștefan, Crăciun, A.M., Moldovan, O.T. (2022): The gram-negative bacilli isolated from caves – Sphingomonas paucimobilis and Hafnia alvei and a review of their involvement in human infections. Int. J. Environ. Res. Public Health 19: 2324.
  41. Jancovich, J.K., Steckler, N.K., Waltzek, T.B. (2015): Ranavirus taxonomy and phylogeny. In: Ranaviruses; pp. 59-70. Gray, M.J., Chinchar, V.G., Eds, Cham, Springer International Publishing.
  42. Knapp, R.A., Briggs, C.J., Smith, T.C., Maurer, J.R. (2011): Nowhere to hide: Impact of a temperature-sensitive amphibian pathogen along an elevation gradient in the temperate zone. Ecosphere 2: art93.
  43. Kumar, S., Stecher, G., Tamura, K. (2016): MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for Bigger Datasets. Mol. Biol. Evol. 33: 1870-1874.
  44. Lisachova, L.S., Lisachov, A.P., Ermakov, O.A., Svinin, A.O., Chernigova, P.I., Lyapkov, S.M., Zamaletdinov, R.I., Pavlov, A.V., Zaks, S.S., Fayzulin, A.I., Korzikov, V.A., Simonov, E. (2025): Continent-wide distribution of CMTV-like Ranavirus, from the Urals to the Atlantic Ocean. EcoHealth 22: 173-184.
  45. Luedtke, J.A., Chanson, J., Neam, K., Hobin, L., Maciel, A.O., Catenazzi, A., Borzée, A., Hamidy, A., Aowphol, A., Jean, A., Sosa-Bartuano, Á., Fong G., A., De Silva, A., Fouquet, A., Angulo, A., Kidov, A.A., Muñoz Saravia, A., Diesmos, A.C., Tominaga, A., Shrestha, B., Gratwicke, B., Tjaturadi, B., Martínez Rivera, C.C., Vásquez Almazán, C.R., Señaris, C., Chandramouli, S.R., Strüssmann, C., Cortez Fernández, C.F., Azat, C., Hoskin, C.J., Hilton-Taylor, C., Whyte, D.L., Gower, D.J., Olson, D.H., Cisneros-Heredia, D.F., Santana, D.J., Nagombi, E., Najafi-Majd, E., Quah, E.S.H., Bolaños, F., Xie, F., Brusquetti, F., Álvarez, F.S., Andreone, F., Glaw, F., Castañeda, F.E., Kraus, F., Parra-Olea, G., Chaves, G., Medina-Rangel, G.F., González-Durán, G., Ortega-Andrade, H.M., Machado, I.F., Das, I., Dias, I.R., Urbina-Cardona, J.N., Crnobrnja-Isailović, J., Yang, J.-H., Jianping, J., Wangyal, J.T., Rowley, J.J.L., Measey, J., Vasudevan, K., Chan, K.O., Gururaja, K.V., Ovaska, K., Warr, L.C., Canseco-Márquez, L., Toledo, L.F., Díaz, L.M., Khan, M.M.H., Meegaskumbura, M., Acevedo, M.E., Napoli, M.F., Ponce, M.A., Vaira, M., Lampo, M., Yánez-Muñoz, M.H., Scherz, M.D., Rödel, M.-O., Matsui, M., Fildor, M., Kusrini, M.D., Ahmed, M.F., Rais, M., Kouamé, N.G., García, N., Gonwouo, N.L., Burrowes, P.A., Imbun, P.Y., Wagner, P., Kok, P.J.R., Joglar, R.L., Auguste, R.J., Brandão, R.A., Ibáñez, R., Von May, R., Hedges, S.B., Biju, S.D., Ganesh, S.R., Wren, S., Das, S., Flechas, S.V., Ashpole, S.L., Robleto-Hernández, S.J., Loader, S.P., Incháustegui, S.J., Garg, S., Phimmachak, S., Richards, S.J., Slimani, T., Osborne-Naikatini, T., Abreu-Jardim, T.P.F., Condez, T.H., De Carvalho, T.R., Cutajar, T.P., Pierson, T.W., Nguyen, T.Q., Kaya, U., Yuan, Z., Long, B., Langhammer, P., Stuart, S.N. (2023): Ongoing declines for the world’s amphibians in the face of emerging threats. Nature 622: 308-314.
  46. Machate, O., Schmeller, D.S., Schulze, T., Brack, W. (2023): Review: Mountain lakes as freshwater resources at risk from chemical pollution. Environ. Sci. Eur. 35: 3.
  47. Mahlen, S.D. (2011): Serratia infections: From military experiments to current practice. Clin Microbiol. Rev. 24: 755-791.
  48. Mao, J., Hedrick, R.P., Chinchar, V.G. (1997): Molecular characterization, sequence analysis, and taxonomic position of newly isolated fish iridoviruses. Virology 229: 212-220.
  49. Marschang, R.E., Meddings, J.I., Waltzek, T.B., Hick, P., Allender, M.C., Wirth, W., Duffus, A.L.J. (2025): Ranavirus distribution and host range. In: Ranaviruses; pp. 155-230. Gray, M.J., Chinchar, V.G., Eds, Cham, Springer.
  50. Masli, S., Vega, J.L. (2011): Ocular immune privilege sites. Methods Mol. Biol. 677: 449-458.
  51. Mavian, C., López-Bueno, A., Balseiro, A., Casais, R., Alcamí, A., Alejo, A. (2012): The genome sequence of the emerging common midwife toad virus identifies an evolutionary intermediate within ranaviruses. J. Virol. 86: 3617-3625.
  52. Meletiadis, A., Di Nicola, M.R., Bovero, S., Favelli, M., Pezzolato, M., Grella, S., Rezza, G., Acutis, P.L. (2025): Prevalence of Batrachochytrium dendrobatidis in amphibians in northwestern Italy’s protected areas. Animals 15: 157.
  53. Miaud, C., Arnal, V., Poulain, M., Valentini, A., Dejean, T. (2019): eDNA increases the detectability of Ranavirus infection in an alpine amphibian population. Viruses 11: 526.
  54. Miaud, C., Pozet, F., Gaudin, N.C.G., Martel, A., Pasmans, F., Labrut, S. (2016): Ranavirus causes mass die-offs of alpine amphibians in the southwestern alps, France. J. Wildl. Dis. 52: 242-252.
  55. Miller, D.L., Pessier, A.P., Hick, P., Whittington, R.J., Forzán, M.J. (2025): Pathology and diagnostics. In: Ranaviruses; pp. 271-312. Gray, M.J., Chinchar, V.G., Eds, Cham, Springer.
  56. Nigrelli, G. (2005): Analysis and characteristics of pluviometric events in the Germanasca Valley (Italian Western Alps). Geogr. Fis. Din. Quat. 28: 147-158.
  57. North, A.C., Hodgson, D.J., Price, S.J., Griffiths, A.G.F. (2015): Anthropogenic and ecological drivers of amphibian disease (Ranavirosis). PLoS ONE 10: e0127037.
  58. Origgi, F.C., Schmidt, B.R., Lohmann, P., Otten, P., Meier, R.K., Pisano, S.R.R., Moore-Jones, G., Tecilla, M., Sattler, U., Wahli, T., Gaschen, V., Stoffel, M.H. (2018): Bufonid herpesvirus 1 (BfHV1) associated dermatitis and mortality in free ranging common toads (Bufo bufo) in Switzerland. Sci. Rep. 8: 14737.
  59. Padilla, D., Acosta, F., Ramos-Vivas, J., Grasso, V., Bravo, J., El Aamri, F., Real, F. (2015): The pathogen Hafnia alveiin veterinary medicine: A review. J. Appl. Anim. Res. 43: 231-235.
  60. Pastorino, P., Barceló, D., Prearo, M. (2024): Alps at risk: High-mountain lakes as reservoirs of persistent and emerging contaminants. J. Contam. Hydrol. 264: 104361.
  61. Pastorino, P., Colussi, S., Pizzul, E., Varello, K., Menconi, V., Mugetti, D., Tomasoni, M., Esposito, G., Bertoli, M., Bozzetta, E., Dondo, A., Acutis, P.L., Prearo, M. (2021): The unusual isolation of carnobacteria in eyes of healthy salmonids in high-mountain lakes. Sci. Rep. 11: 2314.
  62. Pastorino, P., Prearo, M. (2020): High-mountain lakes, indicators of global change: Ecological characterization and environmental pressures. Diversity 12: 260.
  63. Peñafiel-Ricaurte, A., Price, S.J., Fisher, M.C., Bosch, J., Cunningham, A.A., Azat, C. (2025): Ranavirus in native amphibians from the Pyrenees National Park, France: A case study. Eur. J. Wildl. Res. 71: 39.
  64. Pigaiani, N., Bertaso, A., De Palo, E.F., Bortolotti, F., Tagliaro, F. (2020): Vitreous humor endogenous compounds analysis for post-mortem forensic investigation. Forensic Sci. Int. 310: 110235.
  65. Piemonte Parchi (2019): Ranavirus minaccia la rana temporaria. Piemonte Parchi.
  66. Price, S.J., Ariel, E., Maclaine, A., Rosa, G.M., Gray, M.J., Brunner, J.L., Garner, T.W.J. (2017): From fish to frogs and beyond: Impact and host range of emergent ranaviruses. Virology 511: 272-279.
  67. Price, S.J., Garner, T.W.J., Nichols, R.A., Balloux, F., Ayres, C., Mora-Cabello de Alba, A., Bosch, J. (2014): Collapse of amphibian communities due to an introduced Ranavirus. Curr. Biol. 24: 2586-2591.
  68. Price, S.J., Leung, W.T.M., Owen, C.J., Puschendorf, R., Sergeant, C., Cunningham, A.A., Balloux, F., Garner, T.W.J., Nichols, R.A. (2019): Effects of historic and projected climate change on the range and impacts of an emerging wildlife disease. Glob. Change Biol. 25: 2648-2660.
  69. Råman Vinnå, L., Medhaug, I., Schmid, M., Bouffard, D. (2021): The vulnerability of lakes to climate change along an altitudinal gradient. Commun. Earth. Environ. 2: 35.
  70. Rondinini, C., Battistoni, A., Teofili, C. (2022): Lista Rossa IUCN dei vertebrati italiani 2022. Roma, Comitato Italiano IUCN e Ministero dell’Ambiente e della Sicurezza Energetica.
  71. Rosa, G.M., Sabino-Pinto, J., Laurentino, T.G., Martel, A., Pasmans, F., Rebelo, R., Griffiths, R.A., Stöhr, A.C., Marschang, R.E., Price, S.J., Garner, T.W.J., Bosch, J. (2017): Impact of asynchronous emergence of two lethal pathogens on amphibian assemblages. Sci. Rep. 7: 43260.
  72. Rosa, G.M., Ayala Botto, G., Mitra, A.T., Almeida, J.S.D., Hofmann, M., Leung, W.T.M., Alves De Matos, A.P., Caeiro, M.F., Froufe, E., Loureiro, A., Price, S.J., Owen, C., Rebelo, R., Soares, C. (2022): Invasive fish disrupt host-pathogen dynamics leading to amphibian declines. Biol. Conserv. 276: 109785.
  73. Ruggeri, J., Ribeiro, L.P., Pontes, M.R., Toffolo, C., Candido, M., Carriero, M.M., Zanella, N., Sousa, R.L.M., Toledo, L.F. (2019): Discovery of wild amphibians infected with Ranavirus in Brazil. J. Wildl. Dis. 55: 897-902.
  74. Schilliger, L., Paillusseau, C., François, C., Bonwitt, J. (2023): Major emerging fungal diseases of reptiles and amphibians. Pathogens 12: 429.
  75. Seglie, D. (2020): L’erpetofauna del Parco Naturale di Conca Cialancia, con una nota sulla colorazione atipica di due individui di Salamandra lanzai. Riv. Piem. Stor. Nat. 41: 157-166.
  76. Smith, K.F., Acevedo-Whitehouse, K., Pedersen, A.B. (2009): The role of infectious diseases in biological conservation. Anim. Conserv. 12: 1-12.
  77. Smith, K.F., Sax, D.F., Lafferty, K.D. (2006): Evidence for the role of infectious disease in species extinction and endangerment. Conserv. Biol. 20: 1349-1357.
  78. Stöhr, A.C., Hoffmann, A., Papp, T., Robert, N., Pruvost, N.B.M., Reyer, H.-U., Marschang, R.E. (2013): Long-term study of an infection with ranaviruses in a group of edible frogs (Pelophylax kl. esculentus) and partial characterization of two viruses based on four genomic regions. Vet. J. 197: 238-244.
  79. Stöhr, A.C., López-Bueno, A., Blahak, S., Caeiro, M.F., Rosa, G.M., Alves De Matos, A.P., Martel, A., Alejo, A., Marschang, R.E. (2015): Phylogeny and differentiation of reptilian and amphibian ranaviruses detected in Europe. PLoS ONE 10: e0118633.
  80. Teacher, A.G.F., Cunningham, A.A., Garner, T.W.J. (2010): Assessing the long‐term impact of Ranavirus infection in wild common frog populations. Anim. Conserv. 13: 514-522.
  81. Thumsová, B., Alarcos, G., Ayres, C., Rosa, G.M., Bosch, J. (2024): Relationship between two pathogens in an amphibian community that experienced mass mortalities. Conserv. Biol. 38: e14196.
  82. Thumsová, B., Martínez-Silvestre, A., Pérez-Vallejo, M., Palomar, G., Rohr, J.R., Bosch, J. (2025): Temperature and precipitation mismatches increase infection risk in amphibians. Curr. Biol. 35: 4285-4292.
  83. Thumsová, B., Price, S.J., González-Cascón, V., Vörös, J., Martínez-Silvestre, A., Rosa, G.M., Machordom, A., Bosch, J. (2022): Climate warming triggers the emergence of native viruses in Iberian amphibians. iScience 25: 105541.
  84. Tiberti, R., Rogora, M., Tartari, G., Callieri, C. (2014b): Ecological impact of transhumance on the trophic state of alpine lakes in Gran Paradiso National Park. Knowl. Managt. Aquat. Ecosyst. 415: 05.
  85. Tiberti, R., Von Hardenberg, A. (2012): Impact of introduced fish on Common frog (Rana temporaria) close to its altitudinal limit in alpine lakes. Amphib.-Reptil. 33: 303-307.
  86. Tiberti, R., Von Hardenberg, A., Bogliani, G. (2014a): Ecological impact of introduced fish in high altitude lakes: A case of study from the European Alps. Hydrobiologia 724: 1-19.
  87. Trinh, L.L., Nguyen, H.H. (2024): Role of plant-associated microbes in plant health and development: The case of the Serratia genus. Technol. Agron. 4: e028.
  88. Vörös, J., Herczeg, D., Papp, T., Monsalve-Carcaño, C., Bosch, J. (2020): First detection of Ranavirus infection in amphibians in Hungary. Herpetol. Notes 13: 213-217.
  89. White, C.L., Dusek, R.J. (2015): Wildlife specimen collection, preservation, and shipment. U.S. Geological survey techniques and methods, book 15, chap. C4, 24 p.
  90. Yu, Z., Zhang, W., Gu, C., Chen, J., Zhao, M., Fu, L., Han, J., He, M., Xiao, Q., Xiao, W., He, L., Zhang, Z. (2021): Genomic analysis of Ranavirus and exploring alternative genes for phylogenetics. Transbound. Emerg. Dis. 68: 2161-2170.