

1Korea Zoonosis Research Institute, Jeonbuk National University, Iksan 54596, Republic of Korea
2Department of Veterinary and Animal Science, College of Veterinary Medicine, Jeonbuk National University, Iksan 54596, Republic of Korea
© The Author(s), under exclusive licence to Microbiological Society of Korea 2026
This is an Open Access article distributed under the terms of the Creative Commons Attribution 4.0 International License (CC BY 4.0) (https://creativecommons.org/licenses/by/4.0/) which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Acknowledgments
This work was supported by the National Research Foundation of Korea (NRF) grant funded by the Korea government (MSIT) (RS-2026-25489765).
Author Contributions
Jun-Gu Kang conceived the study and designed the experiments. Yun-Seon Shin and Jun-Gu Kang prepared the manuscript Jeong-Yoon Lee and Jun-Gu Kang provided significant intellectual input and revised the manuscript. All the authors have read and approved the final version of the manuscript.
Conflict of Interest
The authors declare no conflict of interest.
| Country | Host species | Specimen type | Detection method | Serology (+/tested) | RNA detected (+/tested) | Reference | ||
|---|---|---|---|---|---|---|---|---|
| China | Bird (unspecified) | Serum | ELISA, qRT-PCR | 7 | 155 | – | – | Li et al. (2016) |
| Cattle | Serum, Blood, Viscera | ELISA, qRT-PCR | 828 | 1501 | 48 | 909 | Cheng et al. (2025); Huang et al. (2019); Ishijima et al. (2023); Jiao et al. (2012); Li et al. (2014); Niu et al. (2013); Xing et al. (2016) | |
| Chicken | Serum, Viscera | ELISA, qRT-PCR | 344 | 1398 | 25 | 609 | Cheng et al. (2025); Huang et al. (2019); Ishijima et al. (2023); Jiao et al. (2012); Li et al. (2014); Niu et al. (2013) | |
| Dog | Serum, Plasma, Viscera | ELISA, qRT-PCR, IFA | 230 | 804 | 26 | 405 | Cheng et al. (2025); Cui et al. (2013); Ding et al. (2014); Huang et al. (2019); Li et al. (2014); Niu et al. (2013); Wanshuang et al. (2024) | |
| Duck | Serum, Viscera | ELISA, qRT-PCR | 12 | 61 | 9 | 61 | Huang et al. (2019) | |
| Goat | Serum, Blood | ELISA, qRT-PCR | 616 | 991 | 32 | 194 | Cheng et al. (2025); Cui et al. (2013); Ding et al. (2014); Hu et al. (2019); Jiao et al. (2012); Li et al. (2014); Lu et al. (2025); Xing et al. (2016); Zhao et al. (2012) | |
| Geese | Serum | ELISA | 2 | 120 | – | – | Li et al. (2014) | |
| Hares | Serum, Viscera | ELISA, qRT-PCR | 46 | 73 | 16 | 73 | Huang et al. (2019) | |
| Hedgehog | Serum, Viscera | ELISA, RT-PCR, qRT-PCR | 56 | 332 | 15 | 232 | Hu et al. (2019); Huang et al. (2019); Jiao et al. (2012); Li et al. (2014, 2016); Sun et al. (2017); Zhao et al. (2022) | |
| Mink | Serum | ELISA | 15 | 178 | – | – | Wang et al. (2017) | |
| Pheasant | Serum, Viscera | ELISA, qRT-PCR | 6 | 14 | 4 | 14 | Huang et al. (2019) | |
| Pig | Serum, Viscera | ELISA, qRT-PCR | 36 | 1369 | 29 | 902 | Hu et al. (2019); Huang et al. (2019); Jiao et al. (2012); Li et al. (2014); Niu et al. (2013) | |
| Rock pigeon | Serum, Viscera | ELISA, qRT-PCR | 8 | 23 | 5 | 23 | Huang et al. (2019) | |
| Rodent | Serum, Viscera | ELISA, qRT-PCR, nested RT-PCR | 67 | 2718 | 23 | 1577 | Cheng et al. (2025); Gu et al. (2022); Hu et al. (2019); Huang et al. (2019); Li et al. (2014, 2016); Liu et al. (2014); Ni et al. (2015) | |
| Sheep | Serum, Viscera | ELISA, qRT-PCR | 381 | 548 | 30 | 1232 | Huang et al. (2019); Niu et al. (2013) | |
| Turtledoves | Serum, Viscera | ELISA, qRT-PCR | 1 | 7 | 1 | 7 | Huang et al. (2019) | |
| Wild boar | Serum | qRT-PCR | – | – | 2 | 6 | Kuan et al. (2023a) | |
| Yellow weasel | Serum, Viscera | ELISA, qRT-PCR | 41 | 45 | 17 | 45 | Huang et al. (2019) | |
| Japan | Cat | Serum, Plasma, Oral/Rectal/Ocular swab, Blood | RT-PCR, qRT-PCR, ELISA, VNT, GLIPS | 109 | 639 | 80 | 201 | Ando et al. (2021); Kida et al. (2019); Matsuu et al. (2019, 2021, 2022); Osako et al. (2024); Sasaki et al. (2026); Yamanaka et al. (2020) |
| Tsushima leopard cats | Serum | ELISA, VNT | 1 | 6 | – | – | Matsuu et al. (2022) | |
| Cattle | Serum | Indirect immunoperoxidase technique | 11 | 510 | – | – | Tabara et al. (2016) | |
| Cheetah | Plasma, Serum, Lymph node, Viscera | qRT-PCR | – | – | 2 | 2 | Matsuno et al. (2018) | |
| Deer | Serum | ELISA, VNT, GLIPS | 552 | 916 | 1 | 229 | Kimura et al. (2018); Matsuu et al. (2021); Tatemoto et al. (2022) | |
| Dog | Serum | ELISA, GLIPS, RT-PCR | 18 | 245 | 1 | 114 | Kimura et al. (2018); Matsuu et al. (2021, 2022) | |
| Japanese badgers | Blood, Viscera, Serum | qRT-PCR, ELISA | 43 | 63 | 2 | 63 | Kaneko et al. (2023) | |
| Raccoon dogs | Blood, Viscera, Serum | qRT-PCR, ELISA, VNT | 12 | 53 | – | – | Kaneko et al. (2023) | |
| Wild boar | Serum | ELISA, GLIPS, VNT, RT-PCR, FRNT, SRIP-neutralization | 421 | 4063 | 9 | 207 | Hayasaka et al. (2016); Kimura et al. (2018); Kirino et al. (2022); Matsuu et al. (2021, 2022); Tatemoto et al. (2022); Yazawa et al. (2025) | |
| Korea | Cat | Serum | ELISA, RT-PCR, nested RT-PCR, IFA | 11 | 203 | 35 | 564 | Han et al. (2022); Hwang et al. (2017); Kang et al. (2020); Lee et al. (2017) |
| Leopard cat | Spleen | RT-PCR | – | – | 1 | 1 | Byun et al. (2026a) | |
| Cattle | Serum | ELISA, nested RT-PCR | 161 | 763 | 34 | 854 | Chae et al. (2024) | |
| Chicken | Serum | ELISA, nested RT-PCR | 83 | 296 | 7 | 312 | Rim et al. (2023) | |
| Deer | Serum, Viscera | ELISA, IFA, RT-PCR, nested RT-PCR, VNT | 45 | 236 | 12 | 450 | Byun et al. (2024); Lee et al. (2020); Oh et al. (2016); Yu et al. (2019) | |
| Dog | Serum, Blood | ELISA, IFA, VNT, RT-PCR, qPCR, nested RT-PCR | 86 | 542 | 21 | 1020 | Byun et al. (2026b); Chung et al. (2020); Kang et al. (2019); Lee et al. (2017, 2018, 2020) | |
| Duck | Serum | ELISA, nested RT-PCR | 81 | 246 | 3 | 249 | Rim et al. (2023) | |
| Goat | Serum, Blood, Whole blood | ELISA, RT-PCR, nested RT-PCR | 73 | 831 | 32 | 1694 | Kang et al. (2018a); Ouh (2026); Yu et al. (2018) | |
| Horse | Serum | ELISA, VNT, RT-PCR | 452 | 887 | 12 | 889 | Han et al. (2023) | |
| Pig | Serum | nested RT-PCR | – | – | 4 | 240 | Kang et al. (2018b) | |
| Wild boar | Serum | ELISA, IFA, nested RT-PCR | 119 | 848 | 50 | 867 | Oh et al. (2016); Rim et al. (2021) | |
| Wild geese | Serum | ELISA, nested RT-PCR | 8 | 26 | 5 | 45 | Rim et al. (2023) | |
| Thailand | Dog | Serum, Blood | VNT, RT-PCR | 76 | 458 | 2 | 508 | Ishijima et al. (2023) |
| Rodent | Viscera | qRT-PCR | – | – | 11 | 1019 | Linsuwanon et al. (2024) | |
| Taiwan | Sheep | Serum | qRT-PCR, nested RT-PCR | 12 | 31 | 39 | 81 | Kuan et al. (2023a), Lin et al. (2020) |
| Goat | Serum | qRT-PCR | – | – | 47 | 166 | Kuan et al. (2023a), Lin et al. (2020) | |
| Cattle | Serum | qRT-PCR, nested RT-PCR | – | – | 166 | 650 | Kuan et al. (2023a) | |
| Wild boar | Serum, Viscera | qRT-PCR | – | – | 2 | 6 | Kuan et al. (2023a) | |
| Mammals (unspecified) | Serum, Viscera | qRT-PCR | – | – | 4 | 257 | Kuan et al. (2023a) | |
| Bird (unspecified) | Serum, Viscera | qRT-PCR | – | – | 1 | 258 | Kuan et al. (2023a) | |
| Dog | Serum, Plasma | RT-PCR | – | – | 111 | 429 | Kuan et al. (2023b) | |
| Cat | Serum, Plasma | RT-PCR | – | – | 59 | 306 | Kuan et al. (2023b) | |
ELISA, enzyme-linked immunosorbent assay; IFA, immunofluorescence assay; VNT, virus neutralization test; GLIPS, gaussia luciferase immunoprecipitation system; FRNT, focus reduction neutralization test; IPT, immunoperoxidase test; SRIP neutralization assay, single-round infectious particles neutralization assay; RT-PCR, reverse transcription polymerase chain reaction; qRT-PCR, quantitative reverse transcription polymerase chain reaction.
| Host species | Specimen | Cell line | Virus isolate names | GenBank accession | Reference |
|---|---|---|---|---|---|
| Apodemus agrarius | Lung | Vero E6 | rodent01 / Zhejiang / 2014, rodent02 / Zhejiang / 2014 | KR698330–KR698331, KR698343–KR698344, KR698356–KR698357 | Hu et al. (2019) |
| Domestic cat | Serum | Vero E6 | Cat/Miyazaki/2018 | LC462229–LC462237 | Kida et al. (2019) |
| Erinaceus europaeus | Spleen | Vero E6, DH82 cells | JS2014-hedgehog-01 / 02 | KR230768–KR230769, KR230788–KR230789, KR230808–KR230809 | Li et al. (2016) |
| Cheetah | Plasma, spleen, lymph node, brain | Huh-7 cells, Vero E6 cells | SkrP/2017, ArtSp/2017 | LC325234–LC325239 | Matsuno et al. (2018) |
| Yellow weasel | Heart | Vero E6 | - | MF574211–MF574213 | Huang et al. (2019) |
| Domestic cat | Serum, mouse spleen homogenate | A129 mouse, Vero E6 | NFe1, NFe3, NFe4, NFe8, NFe10, NFe11, NFe13, NFe37, NFe43, NFe45, NFe57, NFe66, NFe71, NFe76, NFe77, NFe79, NFe82, NFe88, NFe92, NFe93, NFe94, NFe97, NFe100, NFe101, NFe108, NFe111, NFe112, NFe114, NFe115 | MN995224–MN995310 | Matsuu et al. (2019) |
| Sheep, Cattle, Dog | Serum | Vero E6 | SDLZSheep01/2011, SDLZCattle05/2011, SDPLDog01/2011 | JQ693001–JQ693003 | Niu et al. (2013) |
| Dog | Serum | Vero E6 | Dog 22 | KY968712, MH464251, MH464252 | Kang et al. (2019) |
| Cat | Serum | Vero E6 | P2021-03, P2021-09, P2021-31, P2022-03, P2022-06, P2022-08, P2022-10, P2022-13, P2022-17, P2022-29, P2022-31, P2023-06, P2023-12, P2023-13, P2023-18, P2023-24 | PP273701–PP273716 | Sasaki et al. (2026) |
| Cattle | Serum | Vero E6 | - | OR987936–OR987938 | Chae et al. (2024) |
| Korean water deer | Spleen | Vero E6 | - | PP790964, PP790965, PP790966 | Yu et al. (2019) |
| Dog | Serum | Vero E6 | Dog 1, 12 | OL773687, OK423754, OM179920–OM179923 | Han et al. (2022) |
| Wild leopard cat | Spleen | Vero E6 | KLC2025 | PV816800, PV816801, PV816802 | Byun et al. (2026a) |
| Cat | Serum | Vero E6 | Cat 3, 4 | OK423755, OL773688, OL773689, MZ342903, MZ352108, MZ363633 | Lee et al. (2017) |
| Country | Host species | Specimen type | Detection method | Serology (+/tested) | RNA detected (+/tested) | Reference | ||
|---|---|---|---|---|---|---|---|---|
| China | Bird (unspecified) | Serum | ELISA, qRT-PCR | 7 | 155 | – | – | |
| Cattle | Serum, Blood, Viscera | ELISA, qRT-PCR | 828 | 1501 | 48 | 909 | ||
| Chicken | Serum, Viscera | ELISA, qRT-PCR | 344 | 1398 | 25 | 609 | ||
| Dog | Serum, Plasma, Viscera | ELISA, qRT-PCR, IFA | 230 | 804 | 26 | 405 | ||
| Duck | Serum, Viscera | ELISA, qRT-PCR | 12 | 61 | 9 | 61 | ||
| Goat | Serum, Blood | ELISA, qRT-PCR | 616 | 991 | 32 | 194 | ||
| Geese | Serum | ELISA | 2 | 120 | – | – | ||
| Hares | Serum, Viscera | ELISA, qRT-PCR | 46 | 73 | 16 | 73 | ||
| Hedgehog | Serum, Viscera | ELISA, RT-PCR, qRT-PCR | 56 | 332 | 15 | 232 | ||
| Mink | Serum | ELISA | 15 | 178 | – | – | ||
| Pheasant | Serum, Viscera | ELISA, qRT-PCR | 6 | 14 | 4 | 14 | ||
| Pig | Serum, Viscera | ELISA, qRT-PCR | 36 | 1369 | 29 | 902 | ||
| Rock pigeon | Serum, Viscera | ELISA, qRT-PCR | 8 | 23 | 5 | 23 | ||
| Rodent | Serum, Viscera | ELISA, qRT-PCR, nested RT-PCR | 67 | 2718 | 23 | 1577 | ||
| Sheep | Serum, Viscera | ELISA, qRT-PCR | 381 | 548 | 30 | 1232 | ||
| Turtledoves | Serum, Viscera | ELISA, qRT-PCR | 1 | 7 | 1 | 7 | ||
| Wild boar | Serum | qRT-PCR | – | – | 2 | 6 | ||
| Yellow weasel | Serum, Viscera | ELISA, qRT-PCR | 41 | 45 | 17 | 45 | ||
| Japan | Cat | Serum, Plasma, Oral/Rectal/Ocular swab, Blood | RT-PCR, qRT-PCR, ELISA, VNT, GLIPS | 109 | 639 | 80 | 201 | |
| Tsushima leopard cats | Serum | ELISA, VNT | 1 | 6 | – | – | ||
| Cattle | Serum | Indirect immunoperoxidase technique | 11 | 510 | – | – | ||
| Cheetah | Plasma, Serum, Lymph node, Viscera | qRT-PCR | – | – | 2 | 2 | ||
| Deer | Serum | ELISA, VNT, GLIPS | 552 | 916 | 1 | 229 | ||
| Dog | Serum | ELISA, GLIPS, RT-PCR | 18 | 245 | 1 | 114 | ||
| Japanese badgers | Blood, Viscera, Serum | qRT-PCR, ELISA | 43 | 63 | 2 | 63 | ||
| Raccoon dogs | Blood, Viscera, Serum | qRT-PCR, ELISA, VNT | 12 | 53 | – | – | ||
| Wild boar | Serum | ELISA, GLIPS, VNT, RT-PCR, FRNT, SRIP-neutralization | 421 | 4063 | 9 | 207 | ||
| Korea | Cat | Serum | ELISA, RT-PCR, nested RT-PCR, IFA | 11 | 203 | 35 | 564 | |
| Leopard cat | Spleen | RT-PCR | – | – | 1 | 1 | ||
| Cattle | Serum | ELISA, nested RT-PCR | 161 | 763 | 34 | 854 | ||
| Chicken | Serum | ELISA, nested RT-PCR | 83 | 296 | 7 | 312 | ||
| Deer | Serum, Viscera | ELISA, IFA, RT-PCR, nested RT-PCR, VNT | 45 | 236 | 12 | 450 | ||
| Dog | Serum, Blood | ELISA, IFA, VNT, RT-PCR, qPCR, nested RT-PCR | 86 | 542 | 21 | 1020 | ||
| Duck | Serum | ELISA, nested RT-PCR | 81 | 246 | 3 | 249 | ||
| Goat | Serum, Blood, Whole blood | ELISA, RT-PCR, nested RT-PCR | 73 | 831 | 32 | 1694 | ||
| Horse | Serum | ELISA, VNT, RT-PCR | 452 | 887 | 12 | 889 | ||
| Pig | Serum | nested RT-PCR | – | – | 4 | 240 | ||
| Wild boar | Serum | ELISA, IFA, nested RT-PCR | 119 | 848 | 50 | 867 | ||
| Wild geese | Serum | ELISA, nested RT-PCR | 8 | 26 | 5 | 45 | ||
| Thailand | Dog | Serum, Blood | VNT, RT-PCR | 76 | 458 | 2 | 508 | |
| Rodent | Viscera | qRT-PCR | – | – | 11 | 1019 | ||
| Taiwan | Sheep | Serum | qRT-PCR, nested RT-PCR | 12 | 31 | 39 | 81 | |
| Goat | Serum | qRT-PCR | – | – | 47 | 166 | ||
| Cattle | Serum | qRT-PCR, nested RT-PCR | – | – | 166 | 650 | ||
| Wild boar | Serum, Viscera | qRT-PCR | – | – | 2 | 6 | ||
| Mammals (unspecified) | Serum, Viscera | qRT-PCR | – | – | 4 | 257 | ||
| Bird (unspecified) | Serum, Viscera | qRT-PCR | – | – | 1 | 258 | ||
| Dog | Serum, Plasma | RT-PCR | – | – | 111 | 429 | ||
| Cat | Serum, Plasma | RT-PCR | – | – | 59 | 306 | ||
| Host species | Specimen | Cell line | Virus isolate names | GenBank accession | Reference |
|---|---|---|---|---|---|
| Apodemus agrarius | Lung | Vero E6 | rodent01 / Zhejiang / 2014, rodent02 / Zhejiang / 2014 | KR698330–KR698331, KR698343–KR698344, KR698356–KR698357 | |
| Domestic cat | Serum | Vero E6 | Cat/Miyazaki/2018 | LC462229–LC462237 | |
| Erinaceus europaeus | Spleen | Vero E6, DH82 cells | JS2014-hedgehog-01 / 02 | KR230768–KR230769, KR230788–KR230789, KR230808–KR230809 | |
| Cheetah | Plasma, spleen, lymph node, brain | Huh-7 cells, Vero E6 cells | SkrP/2017, ArtSp/2017 | LC325234–LC325239 | |
| Yellow weasel | Heart | Vero E6 | - | MF574211–MF574213 | |
| Domestic cat | Serum, mouse spleen homogenate | A129 mouse, Vero E6 | NFe1, NFe3, NFe4, NFe8, NFe10, NFe11, NFe13, NFe37, NFe43, NFe45, NFe57, NFe66, NFe71, NFe76, NFe77, NFe79, NFe82, NFe88, NFe92, NFe93, NFe94, NFe97, NFe100, NFe101, NFe108, NFe111, NFe112, NFe114, NFe115 | MN995224–MN995310 | |
| Sheep, Cattle, Dog | Serum | Vero E6 | SDLZSheep01/2011, SDLZCattle05/2011, SDPLDog01/2011 | JQ693001–JQ693003 | |
| Dog | Serum | Vero E6 | Dog 22 | KY968712, MH464251, MH464252 | |
| Cat | Serum | Vero E6 | P2021-03, P2021-09, P2021-31, P2022-03, P2022-06, P2022-08, P2022-10, P2022-13, P2022-17, P2022-29, P2022-31, P2023-06, P2023-12, P2023-13, P2023-18, P2023-24 | PP273701–PP273716 | |
| Cattle | Serum | Vero E6 | - | OR987936–OR987938 | |
| Korean water deer | Spleen | Vero E6 | - | PP790964, PP790965, PP790966 | |
| Dog | Serum | Vero E6 | Dog 1, 12 | OL773687, OK423754, OM179920–OM179923 | |
| Wild leopard cat | Spleen | Vero E6 | KLC2025 | PV816800, PV816801, PV816802 | |
| Cat | Serum | Vero E6 | Cat 3, 4 | OK423755, OL773688, OL773689, MZ342903, MZ352108, MZ363633 |
ELISA, enzyme-linked immunosorbent assay; IFA, immunofluorescence assay; VNT, virus neutralization test; GLIPS, gaussia luciferase immunoprecipitation system; FRNT, focus reduction neutralization test; IPT, immunoperoxidase test; SRIP neutralization assay, single-round infectious particles neutralization assay; RT-PCR, reverse transcription polymerase chain reaction; qRT-PCR, quantitative reverse transcription polymerase chain reaction.