📜 Publications

Research articles


Arthofer P, Panhölzl F, Delafont V, Hay A, Reipert S, Cyran N, Wienkoop S, Willemsen A, Sifaoui I, Arberas-Jiménez I, Schulz F, Lorenzo-Morales J, and Horn M. A giant virus infecting the amoeboflagellate Naegleria. Nature Communications, 15(1):3307. doi: 10.1038/s41467-024-47308-2.


Picard MA, Leblay F, Cassan C, Willemsen A, Daron J, Bauffe F, Decourcelle M, Demange A, and Bravo IG. Transcriptomic, proteomic, and functional consequences of codon usage bias in human cells during heterologous gene expression. Protein Science, 32(3):e4576. doi: 10.1002/pro.4576. biorχiv: 10.1101/2022.01.07.475042.


Arthofer P, Delafont V, Willemsen A, Panhölzl F, and Horn M. Defensive symbiosis against giant viruses in amoebae. PNAS, 119(36):e2205856119. doi: 10.1073/pnas.2205856119.


Willemsen A and Bravo IG. Ecological opportunity, radiation events and genomic innovations shaped the episodic evolutionary history of papillomaviruses. biorχiv: 10.1101/2020.03.08.982421.


Willemsen A, van den Boom A, Dietz J, Dagalp SB, Dogan F, Bravo IG, Ehrhardt A, and Ehrke-Schulz E. Genomic and phylogenetic characterization of ChPV2, a novel goat PV closely related to the Xi-PV1 species infecting bovines. Virology Journal, 17(1):167. doi: 10.1186/s12985-020-01440-9.

Fanni Borvető, Bravo IG and Willemsen A. Papillomaviruses infecting cetaceans exhibit signs of genome adaptation following a recombination event. Virus Evolution, veaa038. doi: 10.1093/ve/veaa038.

Flores-Miramontes MG, Olszewski D, Artaza-Irigaray C, Willemsen A, Bravo IG, Vallejo-Ruíz V, Leal- Herrera YA, Piña-Sánchez P, Molina-Pineda A, Cantón-Romero JC, Martínez-Silva MG, Jave-Suárez LF and Aguilar-Lemarroy A. Detection of Alpha, Beta, Gamma and unclassified Human Papillomaviruses in cervical cancer samples from Mexican women. Frontiers in Cellular and Infection Microbiology, 10:234. doi: 10.3389/fcimb.2020.00234.


Willemsen A, Félez-Sánchez M, and Bravo IG. Genome plasticity in Papillomaviruses and de novo emergence of E5 Oncogenes. Genome Biology and Evolution, 11(6):1602-1617. doi: 10.1093/gbe/evz095. biorχiv: 10.1101/337477. Recommended by PCI Evol Biol, doi: 10.24072/pci.evolbiol.100067.

Willemsen A and Bravo IG. Origin and evolution of papillomavirus (onco) genes and genomes. Philosophical Transactions of the Royal Society B, 374(1773):20180303. doi: 10.1098/rstb.2018.0303. biorχiv: 10.1101/428912.


↳ 2018

Willemsen A, Carrasco JL, Elena SF, and Zwart MP. Going, going, gone: predicting the fate of genomic insertions in plant RNA viruses. Heredity, 121(5):499-509. doi: 10.1038/s41437-018-0086-x.

↳ 2017

Willemsen A, Zwart MP, and Elena SF. High virulence does not necessarily impede viral adaptation to a new host: a case study using a plant RNA virus. BMC Evolutionary Biology, 17(1):25. doi: 10.1186/s12862-017-0881-7. bioRχiv: 10.1101/060137.

Willemsen A, Zwart MP, Ambrós S, Carrasco JL, and Elena SF. 2b or Not 2b: Experimental evolution of functional exogenous sequences in a plant RNA virus. Genome Biology and Evolution, 9(2):297-310. doi: 10.1093/gbe/evw300. bioRχiv: 10.1101/079970.

↳ 2016

Willemsen A, Zwart MP, Higueras P, Sardanyés J, and Elena SF. Predicting the stability of homologous gene duplications in a plant RNA virus. Genome Biology and Evolution, 8(9):3065-3082. doi: 10.1093/gbe/evw219. bioRχiv: 10.1101/060517.

Velasco L, Salem N, Willemsen A, Lapidot M, Mansour AN, Rubio L, and Galipienso L. Genetic variation and evolutionary forces shaping Cucumber vein yellowing virus populations: risk of emergence of virulent isolates in Europe. Plant Pathology, 65(5):847-856. doi: 10.1111/ppa.12465.

Willemsen A, Zwart MP, Tromas N, Majer E, Daròs JA, and Elena SF. Multiple barriers to the evolution of alternative gene orders in a positive-strand RNA virus. Genetics, 202(4):1503-1521. doi: 10.1534/genetics.115.185017.

↳ 2015

Galipienso L, Martínez C, Willemsen A, Alfaro-Férnandez A, Font-San Ambrosio I, Davino S, and Rubio L. Genetic variability and evolutionary analysis of parietaria mottle virus: role of selection and genetic exchange. Archives of Virology, 160(10):2611-2616. doi: 10.1007/s00705-015-2550-8.

Janzac B, Willemsen A, Cuevas JM, Glais L, Tribodet M, Verrier J-L, Elena SF, and Jacquot E. Brazilian Potato virus Y isolates identified as members of a new clade facilitate the reconstruction of evolutionary traits within this species Plant Pathology, 64(4):799-807. doi: 10.1111/ppa.12318.

Cuevas J M, Willemsen A, Hillung J, Zwart MP, and Elena SF. Temporal dynamics of intrahost molecular evolution for a plant RNA virus. Molecular Biology and Evolution, 32(5):1132-1147. doi: 10.1093/molbev/msv028.

↳ 2014

Walia J J, Willemsen A, Elci E, Caglayan K, Falk BW, and Rubio L. Genetic variation and possible mechanisms driving the evolution of worldwide Fig mosaic virus isolates. Phytopathology, 104(1):108-114. doi: 10.1094/PHYTO-05-13-0145-R.

Majer E, Salvador Z, Zwart MP, Willemsen A, Elena SF, and Daròs JA. Relocation of the NIb gene in the Tobacco etch potyvirus genome. Journal of Virology, 88(8):4586-4590. doi: 10.1128/JVI.03336-13.

Zwart MP, Willemsen A, Daròs JA, and Elena SF. Experimental evolution of pseudogenization and gene loss in a plant RNA virus. Molecular Biology and Evolution, 31(1):121-134. doi: 10.1093/molbev/mst175.

↳ 2013

Davino S, Willemsen A, Panno S, Davino M, Catara A, Elena SF, and Rubio L. Emergence and phylodynamics of Citrus tristeza virus in Sicily, Italy. PLoS One, 8(6):e66700. doi: 10.1371/journal.pone.0066700.

↳ 2012

Münk C, Willemsen A, and Bravo IG. An ancient history of gene duplications, fusions and losses in the evolution of APOBEC3 mutators in mammals. BMC evolutionary biology, 12:71. doi: 10.1186/1471-2148-12-71.

Bogaert L, Willemsen A, Vanderstraeten E, Bracho MA, de Baere C, Bravo IG, and Martens A. EcPV2 DNA in equine genital squamous cell carcinomas and normal genital mucosa. Veterinary Microbiology, 158(1-2):33-41. doi: 10.1016/j.vetmic.2012.02.005.

Review articles


Willemsen A and Zwart MP. On the stability of sequences inserted into viral genomes. Virus Evolution, 5(2):vez045. doi: 10.1093/ve/vez045.