156. Bakonyi T, Lussy H, Weissenbock H, Hornyak A, Nowotny N (2005) *In vitro* host-cell susceptibility to Usutu virus. *Emerg Infect Dis* 11:298–301. https://doi.org/10.3201/eid1102.041016

157. Hamel R, Phanitchat T, Wichit S, Morales Vargas RE, Jaroenpool J, Diagne CT, Pompon J, Misse D (2021) New insights into the biology of the emerging Tembusu virus. *Pathogens* 10:1010. https://doi.org/10.3390/pathogens10011010

158. Wang CW, Chen YL, Yao SJT, Lin TC, Wu CW, Thongchan D, Wang CY, Wu HY (2022) Pathogenicity of avian polyomaviruses and prospect of vaccine development. *Viruses* 14:2079. https://doi.org/10.3390/v14092079

159. Guerin JL, Gelfi J, Dubois L, Vuillaume A, Boucraut-Baralon C, Pingret JL (2000) A novel polyomavirus (goose hemorrhagic polyomavirus) is the agent of hemorrhagic nephritis enteritis of geese. *J Virol* 74:4523–4529. https://doi.org/10.1128/jvi.74.10.4523-4529.2000

160. Garmyn A, Verlinden M, Bosseler L, Adriaensen C, Martel A (2017) Persistent goose hemorrhagic polyomavirus infection on a Belgian goose farm. *Avian Dis* 61:536–538. https://doi.org/10.1637/11604-020317-Case.1

161. Krautwald ME, Muller H, Kaleta EF (1989) Polyomavirus infection in budgerigers (*Melopsittacus undulatus*): clinical and aetiological studies. *Zentralbl Veterinarmed B* 36:459–467. https://doi.org/10.1111/j.1439-0450.1989.tb00628.x

162. Stoll R, Luo D, Kouwenhoven B, Hobom G, Muller H (1993) Molecular and biological characteristics of avian polyomaviruses: isolates from different species of birds indicate that avian polyomaviruses form a distinct subgenus within the polyomavirus genus. *J Gen Virol* 74:229–237. https://doi.org/10.1099/0022-1317-74-2-229

163. Varsani A, Porzig EL, Jennings S, Krabberger S, Farkas K, Julian L, Massaro M, Ballard G, Ainley DG (2015) Identification of an avian polyomavirus associated with Adelie penguins (*Pygoscelis adeliae*). *J Gen Virol* 96:851–857. https://doi.org/10.1099/vir.0.000038

164. Fawaz M, Vijayakumar P, Mishra A, Gandhale PN, Dutta R, Kamble NM, Sudhakar SB, Roychoudhary P, Kumar H, Kulkarni DD, Raut AA (2016) Duck gut viral metagenome analysis captures snapshot of viral diversity. *Gut Pathog* 8:30. https://doi.org/10.1186/s13099-016-0113-5

165. Williams RAJ, Tolf C, Waldenstrom J (2018) Molecular identification of papillomavirus in ducks. *Sci Rep* 8:9096. https://doi.org/10.1038/s41598-018-27373-6

166. Truchado DA, Williams RAJ, Benitez L (2018) Natural history of avian papillomaviruses. *Virus Res* 252:58–67. https://doi.org/10.1016/j.viruses.2018.05.014

167. Canuti M, Munro HJ, Robertson GJ, Kroyer ANK, Roul S, Ojkic D, Whitney HG, Lang AS (2019) New insight into avian papillomavirus ecology and evolution from characterization of novel wild bird papillomaviruses. *Front Microbiol* 10:701. https://doi.org/10.3389/fmicb.2019.000701

168. Rosenbaum CS, Wunschmann A, Armien D, Burguiera A, Manuguerra JC, Caro V, Eloit M (2012) Human skin microbiota: high diversity of DNA viruses identified on the human skin by high throughput sequencing. *PLoS One* 7:e38499. https://doi.org/10.1371/journal.pone.0038499

169. Van Doorslaer K, Sidi AO, Zanier K, Rybin V, Deryckere F, Rector A, Burk RD, Lienau EK, van Ranst M, Trave G (2009) Identification of unusual E6 and E7 proteins within avian papillomaviruses: cellular localization, biophysical characterization, and phylogenetic analysis. *J Virol* 83:8759–8770. https://doi.org/10.1128/JVI.01777-08

170. Varsani A, Krabberger S, Jennings S, Porzig EL, Julian L, Massaro M, Pollard A, Ballard G, Ainley DG (2014) A novel papillomavirus in Adelie penguin (*Pygoscelis adeliae*) faeces sampled at the Cape Crozier colony, Antarc- tica. *J Gen Virol* 95:1352–1365. https://doi.org/10.1099/vir.0.064436-0

171. Lina PH, van Noord MJ, de Groot FG (1973) Detection of virus in squa- mous papillomas of the wild bird species *Fringilla coelebs*. *J Natl Cancer Inst* 50:567–571. https://doi.org/10.1093/jnci/50.2.567

172. Jacobson ER, Mladinich CR, Clubb S, Sundberg JP, Lancaster WD (1983) Papilloma-like virus infection in an African gray parrot. *J Am Vet Med Assoc* 183:1307–1308

173. Wieland U, Mauch C, Kreuter A, Krieg T, Pfister H (2009) Merkel cell poly- omavirus DNA in persons without merkel cell carcinoma. *Emerg Infect Dis* 15:1496–1498. https://doi.org/10.3201/eid1509.081575

174. Schowalter RM, Pastrana DV, Pumphrey KA, Moyer AL, Buck CB (2010) Merkel cell polyomavirus and two previously unknown polyomaviruses

are chronically shed from human skin. *Cell Host Microbe* 7:509–515. https://doi.org/10.1016/j.chom.2010.05.006

175. Foulongne V, Sauvage V, Hebert C, Dereure O, Cheval J, Gouilh MA, Pari- ente K, Segondy M, Burguiera A, Manuguerra JC, Caro V, Eloit M (2012) Human skin microbiota: high diversity of DNA viruses identified on the human skin by high throughput sequencing. *PLoS One* 7:e38499. https://doi.org/10.1371/journal.pone.0038499

176. Palya V, Zolnai A, Benyeda Z, Kovács E, Kardi V, Mató T (2009) Short beak and dwarfism syndrome of mule duck is caused by a distinct lineage of goose parvovirus. *Avian Pathol* 38:175–180. https://doi.org/10.1080/03079450902737839

177. Ning K, Wang M, Qu S, Lv J, Yang L, Zhang D (2017) Pathogenicity of Pekin duck- and goose-origin parvoviruses in Pekin ducklings. *Vet Microbiol* 210:17–23. https://doi.org/10.1016/j.vetmic.2017.08.020

178. Yu K, Ma X, Sheng Z, Qi L, Liu C, Wang D, Huang B, Li F, Song M (2016) Identification of goose-origin parvovirus as a cause of newly emerging beak atrophy and dwarfism syndrome in ducklings. *J Clin Microbiol* 54:1999–2007. https://doi.org/10.1128/jcm.03244-15

179. Ning K, Liang T, Wang M, Dong Y, Qu S, Zhang D (2018) Pathogenicity of a variant goose parvovirus, from short beak and dwarfism syndrome of Pekin ducks, in goose embryos and goslings. *Avian Pathol* 47:391–399. https://doi.org/10.1080/03079457.2018.1459040

180. Rheinwald JG, Green H (1975) Serial cultivation of strains of human epi- dermal keratinocytes: the formation of keratinizing colonies from single cells. *Cell* 6:331–343. https://doi.org/10.1016/s0092-8674(75)80001-8

181. Couteaudier M, Trapp-Fragnet L, Auger N, Courvoisier K, Pain B, Denes- vre C, Vautherot JF (2015) Derivation of keratinocytes from chicken embryonic stem cells: establishment and characterization of differenti- ated proliferative cell populations. *Stem Cell Res* 14:224–237. https:// doi.org/10.1016/j.scr.2015.01.002

182. Olivier S, Jacoby M, Brillon C, Bouletreau S, Mollet T, Nerriere O, Angel A, Danet S, Souttou B, Guehenneux F, Gauthier L, Berthome M, Vie H, Beltraminelli N, Mehtali M (2010) EB66 cell line, a duck embryonic stem cell-derived substrate for the industrial production of therapeutic monoclonal antibodies with enhanced ADCC activity. *MAbs* 2:405–415. https://doi.org/10.4161/mabs.12350

183. Wolff ET, Laforest P (1960) On the specific histopathological lesions induced by the fowl-pox virus in explants of chick embryo skin in organ culture in vitro. *Ann Inst Pasteur* 98:325–334

184. Dhouailly D (1973) Dermo-epidermal interactions between birds and mammals: differentiation of cutaneous appendages. *J Embryol Exp Morphol* 30:587–603. https://doi.org/10.1242/dev.30.3.587

185. Souci L, Denesvre C (2021) 3D skin models in domestic animals. *Vet Res* 52:21. https://doi.org/10.1186/s13567-020-00888-5

186. Chuong CM (2000) Skin morphogenesis. *Embryonic chicken skin* explant cultures. *Methods Mol Biol* 136:101–106. https://doi.org/10.1385/1-59259-065-9-101

187. Chuong C-M, Ting SA, Widelitz RB, Lee Y-S (1992) Mechanism of skin morphogenesis. *II. Retinoic acid modulates axis orientation and pheno- types of skin appendages*. *Development* 115:839–852. https://doi.org/10.1242/dev.115.3.839

188. Philpott MP (2013) Hair follicle culture—a historical perspective. *Int Soc Hair Restor Surg* 23:130–133. https://doi.org/10.33589/23.2.0130

189. Mallet C, Souci L, Ledevin M, Georgeault S, Larcher T, Denesvre C (2022) Establishment of a culture model for the prolonged maintenance of chicken feather follicles structure in vitro. *PLoS One* 17:e0271448. https://doi.org/10.1371/journal.pone.0271448

190. Andrei G, Duraffour S, Van den Oord J, Snoeck R (2010) Epithelial raft cultures for investigations of virus growth, pathogenesis and efficacy of antiviral agents. *Antiviral Res* 85:431–449. https://doi.org/10.1016/j.antiviral.2009.10.019

191. De La Cruz NC, Mockel M, Niehues H, Rubsam M, Malter W, Zinser M, Krummenacher C, Knebel-Morsdorf D (2023) Ex vivo infection of human skin models with herpes simplex virus 1: accessibility of the receptor nectin-1 during formation or impairment of epidermal barriers is restricted by tight junctions. *J Virol* 97:e002623. https://doi.org/10.1128/jvi.00262-23

192. Muhsen M, Protschka M, Schneider LE, Muller U, Kohler G, Magin TM, Buttner M, Alber G, Siegemund S (2019) Orf virus (ORFV) infection in a three-dimensional human skin model: characteristic cellular
156. Bakonyi T, Lussy H, Weissenbock H, Hornyak A, Nowotny N (2005) *In vitro* host-cell susceptibility to Usutu virus. *Emerg Infect Dis* 11:298–301. https://doi.org/10.3201/eid1102.041016

157. Hamel R, Phanitchat T, Wichit S, Morales Vargas RE, Jaroenpool J, Diagne CT, Pompon J, Misse D (2021) New insights into the biology of the emerging Tembusu virus. *Pathogens* 10:1010. https://doi.org/10.3390/pathogens10011010

158. Wang CW, Chen YL, Yao SJT, Lin TC, Wu CW, Thongchan D, Wang CY, Wu HY (2022) Pathogenicity of avian polyomaviruses and prospect of vaccine development. *Viruses* 14:2079. https://doi.org/10.3390/v14092079

159. Guerin JL, Gelfi J, Dubois L, Vuillaume A, Boucraut-Baralon C, Pingret JL (2000) A novel polyomavirus (goose hemorrhagic polyomavirus) is the agent of hemorrhagic nephritis enteritis of geese. *J Virol* 74:4523–4529. https://doi.org/10.1128/jvi.74.10.4523-4529.2000

160. Garmyn A, Verlinden M, Bosseler L, Adriaensen C, Martel A (2017) Persistent goose hemorrhagic polyomavirus infection on a Belgian goose farm. *Avian Dis* 61:536–538. https://doi.org/10.1637/11604-020317-Case.1

161. Krautwald ME, Muller H, Kaleta EF (1989) Polyomavirus infection in budgerigers (*Melopsittacus undulatus*): clinical and aetiological studies. *Zentralbl Veterinarmed B* 36:459–467. https://doi.org/10.1111/j.1439-0450.1989.tb00628.x

162. Stoll R, Luo D, Kouwenhoven B, Hobom G, Muller H (1993) Molecular and biological characteristics of avian polyomaviruses: isolates from different species of birds indicate that avian polyomaviruses form a distinct subgenus within the polyomavirus genus. *J Gen Virol* 74:229–237. https://doi.org/10.1099/0022-1317-74-2-229

163. Varsani A, Porzig EL, Jennings S, Krabberger S, Farkas K, Julian L, Massaro M, Ballard G, Ainley DG (2015) Identification of an avian polyomavirus associated with Adelie penguins (*Pygoscelis adeliae*). *J Gen Virol* 96:851–857. https://doi.org/10.1099/vir.0.000038

164. Fawaz M, Vijayakumar P, Mishra A, Gandhale PN, Dutta R, Kamble NM, Sudhakar SB, Roychoudhary P, Kumar H, Kulkarni DD, Raut AA (2016) Duck gut viral metagenome analysis captures snapshot of viral diversity. *Gut Pathog* 8:30. https://doi.org/10.1186/s13099-016-0113-5

165. Williams RAJ, Tolf C, Waldenstrom J (2018) Molecular identification of papillomavirus in ducks. *Sci Rep* 8:9096. https://doi.org/10.1038/s41598-018-27373-6

166. Truchado DA, Williams RAJ, Benitez L (2018) Natural history of avian papillomaviruses. *Virus Res* 252:58–67. https://doi.org/10.1016/j.viruses.2018.05.014

167. Canuti M, Munro HJ, Robertson GJ, Kroyer ANK, Roul S, Ojkic D, Whitney HG, Lang AS (2019) New insight into avian papillomavirus ecology and evolution from characterization of novel wild bird papillomaviruses. *Front Microbiol* 10:701. https://doi.org/10.3389/fmicb.2019.000701

168. Rosenbaum CS, Wunschmann A, Armien D, Burguiere A, Manuguerra JC, Caro V, Eloit M (2012) Novel papillomavirus in a mallard duck with mesenchymal chondroid dermal tumors. *J Vet Diagn Invest* 34:231–236. https://doi.org/10.1177/10406387221075607

169. Van Doorslaer K, Sidi AO, Zanier K, Rybin V, Deryckere F, Rector A, Burk RD, Lienau EK, van Ranst M, Trave G (2009) Identification of unusual E6 and E7 proteins within avian papillomaviruses: cellular localization, biophysical characterization, and phylogenetic analysis. *J Virol* 83:8759–8770. https://doi.org/10.1128/JVI.01777-08

170. Varsani A, Krabberger S, Jennings S, Porzig EL, Julian L, Massaro M, Pollard A, Ballard G, Ainley DG (2014) A novel papillomavirus in Adelie penguin (*Pygoscelis adeliae*) faeces sampled at the Cape Crozier colony, Antarc- tica. *J Gen Virol* 95:1352–1365. https://doi.org/10.1099/vir.0.064436-0

171. Lina PH, van Noord MJ, de Groot FG (1973) Detection of virus in squamous papillomas of the wild bird species *Fringilla coelebs*. *J Natl Cancer Inst* 50:567–571. https://doi.org/10.1093/jnci/50.2.567

172. Jacobson ER, Mladinich CR, Clubb S, Sundberg JP, Lancaster WD (1983) Papilloma-like virus infection in an African gray parrot. *J Am Vet Med Assoc* 183:1307–1308

173. Wieland U, Mauch C, Kreuter A, Krieg T, Pfister H (2009) Merkel cell polyomavirus DNA in persons without merkel cell carcinoma. *Emerg Infect Dis* 15:1496–1498. https://doi.org/10.3201/eid1509.081575

174. Schowalter RM, Pastrana DV, Pumphrey KA, Moyer AL, Buck CB (2010) Merkel cell polyomavirus and two previously unknown polyomaviruses

are chronically shed from human skin. *Cell Host Microbe* 7:509–515. https://doi.org/10.1016/j.chom.2010.05.006

175. Foulongne V, Sauvage V, Hebert C, Dereure O, Cheval J, Gouilh MA, Pariente K, Segondy M, Burguiere A, Manuguerra JC, Caro V, Eloit M (2012) Human skin microbiota: high diversity of DNA viruses identified on the human skin by high throughput sequencing. *PLoS One* 7:e38499. https://doi.org/10.1371/journal.pone.0038499

176. Palya V, Zolnai A, Benyeda Z, Kovács E, Kardi V, Mató T (2009) Short beak and dwarfism syndrome of mule duck is caused by a distinct lineage of goose parvovirus. *Avian Pathol* 38:175–180. https://doi.org/10.1080/03079450902737839

177. Ning K, Wang M, Qu S, Lv J, Yang L, Zhang D (2017) Pathogenicity of Pekin duck- and goose-origin parvoviruses in Pekin ducklings. *Vet Microbiol* 210:17–23. https://doi.org/10.1016/j.vetmic.2017.08.020

178. Yu K, Ma X, Sheng Z, Qi L, Liu C, Wang D, Huang B, Li F, Song M (2016) Identification of goose-origin parvovirus as a cause of newly emerging beak atrophy and dwarfism syndrome in ducklings. *J Clin Microbiol* 54:1999–2007. https://doi.org/10.1128/jcm.03244-15

179. Ning K, Liang T, Wang M, Dong Y, Qu S, Zhang D (2018) Pathogenicity of a variant goose parvovirus, from short beak and dwarfism syndrome of Pekin ducks, in goose embryos and goslings. *Avian Pathol* 47:391–399. https://doi.org/10.1080/03079457.2018.1459040

180. Rheinwald JG, Green H (1975) Serial cultivation of strains of human epidermal keratinocytes: the formation of keratinizing colonies from single cells. *Cell* 6:331–343. https://doi.org/10.1016/s0092-8674(75)80001-8

181. Couteaudier M, Trapp-Fragnet L, Auger N, Courvoisier K, Pain B, Denesvre C, Vautherot JF (2015) Derivation of keratinocytes from chicken embryonic stem cells: establishment and characterization of differentiated proliferative cell populations. *Stem Cell Res* 14:224–237. https://doi.org/10.1016/j.scr.2015.01.002

182. Olivier S, Jacoby M, Brillon C, Bouletreau S, Mollet T, Nerriere O, Angel A, Danet S, Souttou B, Guehenneux F, Gauthier L, Berthome M, Vie H, Beltraminelli N, Mehtali M (2010) EB66 cell line, a duck embryonic stem cell-derived substrate for the industrial production of therapeutic monoclonal antibodies with enhanced ADCC activity. *MAbs* 2:405–415. https://doi.org/10.4161/mabs.12350

183. Wolff ET, Laforest P (1960) On the specific histopathological lesions induced by the fowl-pox virus in explants of chick embryo skin in organ culture in vitro. *Ann Inst Pasteur* 98:325–334

184. Dhouailly D (1973) Dermo-epidermal interactions between birds and mammals: differentiation of cutaneous appendages. *J Embryol Exp Morphol* 30:587–603. https://doi.org/10.1242/dev.30.3.587

185. Souci L, Denesvre C (2021) 3D skin models in domestic animals. *Vet Res* 52:21. https://doi.org/10.1186/s13567-020-00888-5

186. Chuong CM (2000) Skin morphogenesis. *Embryonic chicken skin* explant cultures. *Methods Mol Biol* 136:101–106. https://doi.org/10.1385/1-59259-065-9-101

187. Chuong C-M, Ting SA, Widelitz RB, Lee Y-S (1992) Mechanism of skin morphogenesis. II. Retinoic acid modulates axis orientation and phenotypes of skin appendages. *Development* 115:839–852. https://doi.org/10.1242/dev.115.3.839

188. Philpott MP (2013) Hair follicle culture—a historical perspective. *Int Soc Hair Restor Surg* 23:130–133. https://doi.org/10.33589/23.2.0130

189. Mallet C, Souci L, Ledevin M, Georgeault S, Larcher T, Denesvre C (2022) Establishment of a culture model for the prolonged maintenance of chicken feather follicles structure in vitro. *PLoS One* 17:e0271448. https://doi.org/10.1371/journal.pone.0271448

190. Andrei G, Duraffour S, Van den Oord J, Snoeck R (2010) Epithelial raft cultures for investigations of virus growth, pathogenesis and efficacy of antiviral agents. *Antiviral Res* 85:431–449. https://doi.org/10.1016/j.antiviral.2009.10.019

191. De La Cruz NC, Mockel M, Niehues H, Rubsam M, Malter W, Zinser M, Krummenacher C, Knebel-Morsdorf D (2023) Ex vivo infection of human skin models with herpes simplex virus 1: accessibility of the receptor nectin-1 during formation or impairment of epidermal barriers is restricted by tight junctions. *J Virol* 97:e002623. https://doi.org/10.1128/jvi.00262-23

192. Muhsen M, Protschka M, Schneider LE, Muller U, Kohler G, Magin TM, Buttner M, Alber G, Siegemund S (2019) Orf virus (ORFV) infection in a three-dimensional human skin model: characteristic cellular
alterations and interference with keratinocyte differentiation. PLoS One
14:e0210504. https://doi.org/10.1371/journal.pone.0210504

193. Loke ASW, Longley BJ, Lambert PF, Spurgeon ME (2021) A novel in vitro culture model system to study merkel cell polyomavirus-associated MCC using three-dimensional organotypic raft equivalents of human skin. Viruses 13:138. https://doi.org/10.3390/v13010138

194. Jiang Y, Tsoi LC, Billi AC, Ward NL, Harms PW, Zeng C, Maverakis E, Kahlenberg JM, Gudjonsson JE (2020) Cytokinocytes: the diverse contribution of keratinocytes to immune responses in skin. JCI Insight 5:e142067. https://doi.org/10.1172/jci.insight.142067

195. Monson EA, Trenerry AM, Laws JL, Mackenzie JM, Helbig KJ (2021) Lipid droplets and lipid mediators in viral infection and immunity. FEMS Microbiol Rev 45:fuaa066. https://doi.org/10.1093/femsre/fuaa066

**Publisher's Note**

Springer Nature remains neutral with regard to jurisdictional claims in pub-
lished maps and institutional affiliations.
alterations and interference with keratinocyte differentiation. PLoS One
14:e0210504. https://doi.org/10.1371/journal.pone.0210504

193. Loke ASW, Longley BJ, Lambert PF, Spurgeon ME (2021) A novel in vitro culture model system to study merkel cell polyomavirus-associated MCC using three-dimensional organotypic raft equivalents of human skin. Viruses 13:138. https://doi.org/10.3390/v13010138

194. Jiang Y, Tsoi LC, Billi AC, Ward NL, Harms PW, Zeng C, Maverakis E, Kahlenberg JM, Gudjonsson JE (2020) Cytokinocytes: the diverse contribution of keratinocytes to immune responses in skin. JCI Insight 5:e142067. https://doi.org/10.1172/jci.insight.142067

195. Monson EA, Trenerry AM, Laws JL, Mackenzie JM, Helbig KJ (2021) Lipid droplets and lipid mediators in viral infection and immunity. FEMS Microbiol Rev 45:fuaa066. https://doi.org/10.1093/femsre/fuaa066

**Publisher's Note**

Springer Nature remains neutral with regard to jurisdictional claims in pub-
lished maps and institutional affiliations.