Prof. Fernando González Candelas, PhD

Instituto Cavanilles de Biodiversidad y Biología Evolutiva

Universidad de Valencia


phone: +34 96 354 3653

fax: +34 96 354 3670

Last update: January 19, 2015


Full Professor of Genetics, Department of Genetics
University of Valencia (Spain)


I am also Research Associate in the following institutions:

Institut Cavanilles de Biodiversitat i Biologia Evolutiva (ICBiBE), University of Valencia.

CIBER en Epidemiología y Salud Pública (CIBERESP), Instituto de Salud Carlos III

FISABIO-Salud Pública (Formerly CSISP), Generalitat Valenciana

Some of my activities are reflected in these blogs:

Evolutionaria, my “academic” blog, in Spanish

A-probando la evolución, ESTIC Project with Biology majors, cotutored with Juli Peretó. In Spanish, English and Catalán. This project has now expanded to include a wiki site with learning resources for evolutionary biology: wikievo

Our research on molecular phylogenetics applied to forensic analysis of a large hepatitis C outbreak (BMC Biol 11: 76) has been highlighted in Nature , BMC Biology and The Scientist


You can also follow my work in ORCID Scopus GoogleScholar 

My book in Spanish ("La evolución, de Darwin al genoma") and Catalan ("L'evolució, de Darwin al genoma") versions has just been published.

I have coauthored a book on GMOs ("Transgénicos") with Carmen Fenoll, from UCLM.



1982      B.Sc. (Biology), University of Valencia

1987      Ph.D. (Genetics), University of Valencia

Major research interests

My main research interests are in population and evolutionary genetics, molecular and evolutionary epidemiology, molecular systematics and genomics, bioinformatics and conservation biology. I am currently working on the molecular evolutionary epidemiology of different pathogens, mainly RNA viruses, such as hepatitis C virus (HCV) and human immunodeficiency virus (HIV), and bacteria, such as Legionella pneumophila. Our basic approach is the analysis of nucleotide sequence variability at different levels, from intrapatient to world-wide samples, depending on the speficic goals of the different projects.

Previously I have studied the population and evolutionary biology of Mediterranean endemic Limonium (Plumbaginaceae) species, by using an array of genetic markers (RAPDs, AFLPs, microsatellites, isozymes) and including the analysis of quantitative traits. Besides, I have been very involved in the establishment of the Bioinformatics Service at my university, of which I was in charge from 1994 to 2000. I have also helped in the development of the Automated Sequencing Service. I have served as head of the Bioinformatics Service at the University of Valencia from 1994 until 2001 and as Scientific Expert on the Spanish National Biosafety Council from 1998 until 2010.

Current research projects

  • Molecular and evolutionary epidemiology of RNA virus
  • Molecular populations genetics and evolutionary epidemiology of Legionella pneumophila
  • Genomic epidemiology
  • Comparative and evolutionary genomics, phylogenomics
  • Population and evolutionary genetics of pathogens, parasites and their vectors
  • Conservation genetics of endangered plant species


My research is developed in the Evolution and Health group at the University of Valencia and in the Joint Research Unit "Infection and Health" FISABIO-Salud Pública/IUniversity of Valencia.

Research Associates

Dr. María Alma Bracho Lapiedra, CIBERESP, CSISP

Dr. Iñaki Comas, CIBERESP, CSISP

Dr. Xavier López-Labrador, CIBERESP, CSISP

Dr. Rafael Sanjuán, ICBiBE


Graduate students

Leonor Sánchez Busó (PhD Thesis: Genomic epidemiology of Legionella pneumophila)

Juan Angel Patiño Galindo (PhD Thesis: Molecular epidemiology of HIV in the Comunidad Valenciana). Co-advisors: María Alma Bracho and Fco. Xavier López-Labrador.

Paula Ruiz Hueso (Master Thesis: Genetic and genomic characterization of antibiotic resistance genes and strains in hospitals from the Comunidad Valenciana)

Vicente Sentandreu, (PhD thesis: Evolutionary dynamics of HCV and HIV in coinfected patients)

Bárbara Gomila (PhD Thesis: Molecular epidemiology of Mycobacterium tuberculosis), co-advisor: Iñaki Comas

Alvaro Chiner (PhD Thesis: Evolutionary systems biology of Mycobacterium tuberculosis), co-advisor: Iñaki Comas

Claudia García (MSc Thesis: Defining species in the Mycobacteria), co-advisor: Iñaki Comas

Francisco J. Roig (MSc Thesis: Phylogenomics of Vibrio vulnificus)

Visiting scholars

Undergraduate students

Beatriz Beamud Aranguren

Manuel Ferrando Bernal

Luis Moll Martínez

Marta Pla Díaz

Neris García González

Carlos Valiente Mullor

Ana Martínez

Yaiza Arenas Ortiz

Paula Izquierdo


Dr. Natahsa Arora, Univ. Zurich

Dr. José Manuel Cuevas, ICBiBE, CIBERESP

Dr. Juan Carlos Galán, CIBERESP, Hosp. Ramón y Cajal

Dr. África Holguín, CIBERESP, Hosp. Ramón y Cajal

Dr. Vicente Ausina, CIBERES, Hosp. Germans Trias i Pujol

Dr. Elisa Martró, CIBERESP, Hosp. Germans Trias i Pujol

Dr. Miguel Thomson, Instituto de Salud Carlos III

Dr. Manuel Zúñiga, IATA

Dr. Carmen Amaro, Dpto. Microbiología, Univ. Valencia

Dr. Pilar Catalán, Univ. Zaragoza

Dr. José Gabriel Segarra, Univ. Zaragoza

Dr. María Dolores Bargues, Dpt. Parasitología, Univ. Valencia

 Former PhD students

Dr. Alicia Amadoz,  CIMA (PhD thesis: Bioinformatic methods for molecular epidemiology analysis, 2011)

Dr. Mireia Coscollá, Swiss TPH (PhD thesis: Population genetics and molecular epidemiology of Legionella pneumophila, 2009)

Dr. Iñaki Comas, FISABIO-Salud Pública (PhD thesis: Phylogenomics of Proteobacteria, 2008)

Dr. Marisa Palop Esteban, (PhD thesis: Population genetics with SSR markers of endangered Limonium sp, 2003.)

Dr. Manuela Torres-Puente, (PhD thesis: Molecular epidemiology of HCV in the Comunidad Valenciana, 2004)

Dr. Carmen Palacios, (PhD thesis: Conservation genetics and evolution of Limonium spp, 1997)

Dr. Adoración Hernández, (PhD thesis: Effects of recurrent bottlenecks in experimental populations of Drosophila psuedobbscura, 1993)

Former Postdocs

Dr. Borys Wróbel, Institut of Oceanography - Polish Academy of Sciences, Methods for dating viral infections.

Dr. Ferrán Palero

Former Visiting Scientists

Maria Nowakowaka

Henry Arenas Castro

José María González Alba

Maria Keramarou

Jovani Barajas



Selected publications PubMed

Pérez-Losada, Marcos; Arenas, Miguel; Galán, Juan Carlos; Palero, Ferran; González-Candelas, Fernando. 2015. Recombination in viruses: mechanisms, methods of study, and evolutionary consequences. Infection, Genetics and Evolution 30: 296-307.
[Abstract] [PDF] [Full Text]

Sánchez-Busó, Leonor; Olmos, María Piedad; Camaró, María Luisa; Adrián, Francisco; Calafat, Juan Miguel; González-Candelas, Fernando. 2015. Phylogenetic analysis of environmental Legionella pneumophila isolates from an endemic area (Alcoy, Spain). Infection, Genetics and Evolution 30:45-54.
[Abstract] [PDF] [Full Text]

Sánchez-Busó, Leonor; Comas, Iñaki; Jorques, Guillermo; González-Candelas, Fernando. 2014. Recombination drives genome evolution in outbreak-related Legionella pneumophila isolates. Nature Genetics 46(11): 1205-1211.
[Abstract] [PDF] [Full Text]

Lara, James; López-Labrador, F. Xavier; González-Candelas, Fernando; Berenguer, Marina; Khudyakov, Yury E. 2014. Computational models of liver fibrosis progression for hepatitis C virus chronic infection. BMC Bioinformatics 15(Suppl 8): S5.
[Abstract] [PDF] [Full Text]

Coscollá, Mireia; Fernández, Carmen; Colomina, Javier; Sánchez-Busó, Leonor; González-Candelas, Fernando. 2014. Mixed infection by Legionella pneumophila in outbreak patients. International Journal of Medical Microbiology 304 (3-4): 307-313.
[Abstract] [PDF] [Full Text]

Domínguez, Angela; Godoy, Pere; Castilla, Jesús; Mayoral, José María; Soldevila, Nuria; Torner, Nuria; Toledo, Diana; Astray,  Jenaro; Tamames, Sonia; García-Gutiérrez, Susana; González-Candelas, Fernando; Martín, Vicente; Díaz, José; and the CIBERESP Working Group for the Survey on Influenza Vaccination in Primary Health Care Workers. 2013. Knowledges of and attitudes to influenza vaccination in primary healthcare workers in Spain, 2011-2012.
PLoS  ONE  8(11): e81200.
[Abstract] [PDF] [Full Text]

Pérez-Lago, Laura; Comas, Iñaki; Navarro, Yurena; González-Candelas, Fernando; Herranz, Marta; Bouza, Emilio; García-de-Viedma, Darío.
2014. Whole Genome Sequencing analysis of intrapatient microevolution in Mycobacterium tuberculosis: Potential impact on the inference of tuberculosis transmission Journal of Infectious Diseases 209(1): 98-108.
[Abstract] [PDF] [Full Text]

Bracho, M.A.; Sentandreu, V.; Alastrué, I.; Belda, J.; Juan, A.; Fernández-García, E.; Santos, C.; Zafra, T.; Tasa, T.; González-Candelas, F. 2014.
Emerging trends in CRF02_AG transmission among men who have sex with men in Spain. Journal of Acquired Immune Deficiency Syndromes 65(3): e130-e133.  

Castilla, Jesús; Martínez-Baz, Iván; Godoy, Pere; Toledo, Diana; Astray, Jenaro; García, Susana; Mayoral, José María; Martín, Vicente; González-Candelas, Fernando; Guevara, Marcela; Diaz-Borrego, José; Torner, Núria; Baricot, Maretva; Tamames, Sonia; Domínguez, Angela; Working Group for the Survey on Influenza Vaccination in Primary Health Care Professionals.
2013. Trends in influenza vaccine coverage among primary healthcare workers in Spain, 2008-2011. Preventive Medicine 57: 206-211.
[Abstract] [PDF] [Full Text]

Delgado-Rodríguez, Miguel; Castilla, Jesús; Godoy, Pere; Martín, Vicente; Soldevila, Nuria; Alonso, Jordi; Astray, Jenaro; Baricot, Maretva; Galán, Juan Carlos; Castro, Ady; González-Candelas, Fernando; Mayoral, José María; Quintana, José María; Pumarola, Tomás; Tamames, Sonia; Sáez, Marc; Domínguez, Àngela; and the CIBERESP Cases and Controls in Pandemic Influenza Working Group, Spain.
2013. Different prognosis in hospitalized patients with influenza one season after the pandemic H1N1 flu of 2009-2010 in Spain. Influenza and Other Respiratory Viruses 7(6): 1336-1342.
[Abstract] [PDF] [Full Text]

González-Candelas, F.; Bracho, M.A.; Wróbel, B.; Moya, A. 2013. Molecular evolution in court: Analysis of a large hepatitis C virus outbreak from an evolving source. BMC Biology 11: 76.
[Abstract] [PDF] [Additional files]

Castilla, Jesús; Godoy, Pere; Domínguez, Ángela; Martínez-Baz, Iván; Astray, Jenaro; Martín, Vicente; Delgado-Rodríguez, Miguel; Baricot, Maretva; Soldevila, Nuria; Mayoral, José María; Quintana, José María; Galán, Juan Carlos; Castro, Ady; González-Candelas, Fernando; Garín, Olatz; Saez, Marc; Tamames, Sonia; Pumarola, Tomás and the CIBERESP Cases and Controls in Influenza Working Group. 2013. Influenza vaccine effectiveness in preventing outpatient, hospitalized and severe cases of laboratory-confirmed influenza: multicase-control study. Clinical Infectious Diseases 57(2): 167-175.
[Abstract] [PDF]

Sánchez-Busó, Leonor; Coscollá, Mireia; Pinto-Carbó, Marta; Catalán, Vicente; González-Candelas, Fernando. 2013. Genetic characterization of Legionella pneumophila isolated from a common watershed in Comunidad Valenciana, Spain. PLoS ONE 8(4): e61564.
[Abstract] [PDF]

Galán, Juan-Carlos; González-Candelas, Fernando; Rolain, Jean-Marc; Cantón, Rafael. 2013. Antibiotics as selectors and accelerators 1 of diversity in the mechanisms of resistance: From the resistome to genetic plasticity in the β -lactamases world. Frontiers in  Microbiology (Antimicrobials, Resistance and Chemotherapy) 4:9.
[Abstract] [PDF]

Matas, Marina; Picornell, Antònia; Cifuentes, Carmen; Payeras, Antoni; Bassa, Antoni; Homar, Francesc; González-Candelas, Fernando; López-Labrador, F. Xavier; Moya, Andrés; Ramon, Maria M.; Castro, José A. 2013. Generalized Linear Model (GLM) framework for the association of host variables and viral strains with liver fibrosis in HCV/HIV coinfected patients. Infection, Genetics and Evolution 13(1): 284-291.
[Abstract] [PDF]

Saludes, Verónica; González-Candelas, Fernando; Planas, Ramon; Solà, Ricard; Ausina, Vicente; Martró, Elisa. 2013. Evolutionary dynamics of the E1-E2 viral populations during combination therapy in non-responder patients chronically infected by hepatitis C virus subtype 1b. Infection, Genetics and Evolution 13(1): 1-10.
[Abstract] [PDF]

Roy, K.M.; Galmés Truyols, A.; Giménez Duran, J.; Anderson, E.; Prempeh, H.; González Candelas, F.; Bracho, M.A.; Aitkin, C.; McTier, R.; Tejera, E. for and on behalf of the Incident Groups in Scotland and Spain. 2012. Epidemiology and molecular investigation of hepatitis C infection following holiday haemodialysis. Journal of Hospital Infection 82(3): 158-163.
[Abstract] [PDF]

Martínez, Fernando; Lafforgue, Guillaume; Morelli, Marco J.; González-Candelas, Fernando; Chua, Nam-Hai; Daròs, José-Antonio; Elena, Santiago F. 2012. Ultra-deep Sequencing Analysis of Population Dynamics of Virus Escape Mutants in RNAi-mediated Resistant Plants. Molecular Biology and Evolution 29(11): 3297-3307.
[Abstract] [PDF]

Garcia-Garcerà, Marc; Coscollà, Mireia; Garcia-Etxebarria, Koldo; Martín-Caballero, Juan; González-Candelas, Fernando; Latorre, Amparo; Calafell, Francesc. 2012. Staphylococcus prevails in the skin microbiota of long-term immunodeficient mice. Environmental Microbiology 14(8): 2087-2098.
[Abstract] [PDF]

Fernando González-Candelas, Jenaro Astray, Jordi Alonso, Ady Castro, Rafael Cantón, Juan Carlos Galán, Olatz Garin,Marc Saez, Nuria Soldevila, Maretva Baricot, Jesús Castilla, Pere Godoy, Miguel Delgado-Rodríguez, Vicente Martín, José María Mayoral, Tomás Pumarola, José María Quintana, Sonia Tamames, Angela Domínguez and the CIBERESP Cases and Controls in Pandemic Influenza Working Group, Spain. 2012. Sociodemographic Factors and Clinical Conditions Associated with Hospitalization in Influenza A (H1N1) 2009 Virus Infected Patients in Spain, 2009-2010. PLoS ONE 7(3): e33139.
[Abstract] [PDF]

Zúñiga, Manuel; Gómez-Escoín, Ciara Luna; and González-Candelas, Fernando. 2011. Evolutionary history of the OmpR/IIIA family of signal transduction two component systems in Lactobacillaceae and Leuconostocaceae. BMC Evolutionary Biology 11: 34.
[Abstract] [PDF]

López Davia, Javier; Hernández Bel, Pablo; Zaragoza Ninet, Violeta; Bracho, María Alma; González-Candelas, Fernando; Salazar, Antonio; Gobernado, Miguel; Febrer Bosch, Isabel. 2011. Onychomadesis outbreak in Valencia, Spain associated with Hand, Foot, and Mouth disease caused by enteroviruses. Pediatric Dermatology 28(1): 1-5.
[Abstract] [PDF]

Belén Rodríguez Sánchez; María Alonso; Pilar Catalán; Matilde Sánchez Conde; Fernando González-Candelas; Maddalena Giannella; Emilio Bouza; Darío García de Viedma. 2011. Genotyping of a nosocomial outbreak of pandemic influenza A/H1N1 2009. Journal of Clinical Virology 52(2): 129-132.
[Abstract] [PDF]

J.M. González-Alba, A. Holguín, R. García, S. García-Bujalance, R. Alonso , A. Suárez, R. Delgado, L. Cardeñoso, R. González, I. García-Bermejo, F. Portero ,C de Mendoza, F. González-Candelas, J.C. Galán. 2011. Molecular surveillance of HIV-1 in Madrid, Spain: a phylogeographic analysis. Journal of Virology 85(20): 10755-10763.
[Abstract] [PDF]

Bracho, Maria A.; González-Candelas, Fernando; Valero, Ana; Córdoba, Juan; Salazar, Antonio. 2011. Enterovirus Co-infections and Onychomadesis after Hand, Foot, and Mouth Disease, Spain, 2008. Emerging Infectious Diseases 17(12): 2223-2231.
[Abstract] [PDF]

González-Candelas, Fernando; López-Labrador, Fco. Xavier; Bracho, María Alma. 2011. Recombination in HCV. Viruses 3(10): 2006-2024.
[Abstract] [PDF]

Roig, F.; González-Candelas, F.; Amaro, C. 2011. Domain organization and evolution of Multifunctional Autoprocessing Repeats-in-Toxin (MARTX) toxin in Vibrio vulnificus. Applied and Environmental Microbiology 77(2): 657-668.

Sanjuán, E.; González-Candelas, F.; Amaro, C. 2011. Polyphyletic origin of Vibrio vulnificus biotype 2 as revealed by sequence-based analysis. Applied and Environmental Microbiology 77(2): 688-695.
[Abstract] [PDF]

Coscollá, M.; Comas, I.; González-Candelas, F. 2011. Quantifying Nonvertical Inheritance in the Evolution of Legionella pneumophila. Molecular Biology and Evolution 28(2): 985-1001.
[Abstract] [PDF]

Coscollá, M.; Fenollar, J.; Escribano, I.; González-Candelas, F. 2010. Legionellosis outbreak associated with asphalt paving machine, Spain, 2009. Emerging Infectious Diseases 16(9): 1381-1387.

González-Candelas, F.; Guiral, S.; Carbó, R.; Valero, A.; Vanaclocha, H.; González, F.; Bracho, M.A. 2010. Patient-to-patient transmission of hepatitis C virus (HCV) during colonoscopy diagnosis. Virology Journal 7: 217.

Saludes, V.; Bracho, M.A.; Valero, O.; Ardèvol, M.; Planas, R.; González-Candelas, F.; Ausina, V.; Martró, E. 2010. Baseline prediction of combination therapy outcome in hepatitis C virus genotype 1b infected patients by discriminant analysis using viral and host factors. PLoS ONE 5(11): e14132.
[Abstract] [PDF]

González-Candelas, F. 2010. Molecular Phylogenetic Analyses in Court Trials. In: Encyclopedia of Life Sciences (ELS). John Wiley & Sons, Ltd: Chichester. DOI: 10.1002/9780470015902.a0021575

González-Candelas, F.; López-Labrador, F.X. 2010. Clinical relevance of genetic heterogeneity in HCV. Future Virology 5(1): 33-49

Novais, Â.; Comas, I.; Baquero, F.; Cantón, R.; Coque, T.M.; Moya, A.; González-Candelas, F.; Galán, J.C. 2010. Evolutionary Trajectories of Beta-Lactamase CTX-M-1 Cluster Enzymes: Predicting Antibiotic Resistance. PLoS Pathogens 6(1): e1000735. doi:10.1371/journal.ppat.1000735. [Full text] [PDF] [Supplementary Material]

Sánchez, B.; Zúñiga, M.; González-Candelas, F.; González de los Reyes-Gavilán,C.; Margolles,A.. 2010. Bacterial and eukaryotic phosphoketolases: phylogeny, distribution and evolution. Journal of Molecular Microbiology and Biotechnology 18: 37-51.
[Abstract] [PDF]

Coscollá, M.; González-Candelas, F. 2009. Comparison of clinical and environmental samples of Legionella pneumophila at the nucleotide sequence level. Infection, Genetics and Evolution 9: 882-888.
[Abstract] [PDF] [Supplementary Data]

Coscollá, M.; González-Candelas, F. 2009. Direct sequencing of Legionella pneumophila from respiratory samples for Sequence-Based Typing analysis. Journal of Clinical Microbiology 47(9): 2901-2905.
[Abstract] [PDF] [Supplemental Data]

Cuevas, J.M.; González, M.;Torres-Puente, M.; Jiménez-Hernández, N.; Bracho, M.A.; García-Robles, I.; González-Candelas, F.; Moya, A. 2009.The role of positive selection in hepatitis C virus. Infection, Genetics and Evolution 9: 882-888.
[Abstract] [PDF]

Cuevas,J.M.; Gonzalez-Candelas,F.; Moya,A.; Sanjuan,R. 2009. The effect of ribavirin on the mutation rate and spectrum of Hepatitis C virus in vivo. J. Virol. 83: 5760-5764.
[Abstract] [PDF] [Supplemental material]

Cuevas, J.M.; Torres-Puente, M.; Jiménez-Hernández, N.; Bracho, M.A.; García-Robles, I.; Carnicer, F.; Olmo, J.D.; Ortega, E.; González-Candelas, F.; Moya, A. 2009. Combined therapy of interferon plus ribavirin promotes multiple adaptive solutions in hepatitis C virus. Journal of Medical Virology 81(4): 650-656.
[Abstract] [PDF]

Domingo-Calap, P.; Sentandreu,V.; Bracho, M.A.; González-Candelas, F.; Moya,A.; Sanjuán, R. 2009. Unequal distribution of RT-PCR artifacts along the E1-E2 region of Hepatitis C virus. Journal of Virological Methods 161: 136-140.

Lourenço Jr., V.; Moya, A.; González-Candelas, F.; Carbone, I.; Maffia, L.A.; Mizubuti, E. 2009. Molecular diversity and evolutionary processes of Alternaria solani in Brazil inferred using genealogical and coalescent approaches. Phytopathology 99: 765-774.

Juárez, P.; Comas, I.; González-Candelas, F.; Calvete, J.J. 2008. Evolution of snake venom disintegrins by positive Darwinian selection. Molecular Biology and Evolution 25: 2391-2407.
[Abstract] [PDF] [Full Text]

Bargues, M.D.; Klisiowicz, D.R.; González-Candelas, F.; Ramsey, J.M.; Monroy, C.; Ponce, C-; Salazar-Schettino, P.M.; Panzera, F.; Abad-Franch, F.; Sousa, O.E.; Schofield, C.J.; Dujardin, J.P.; Guhl, F.; Mas-Coma, S. 2008. Phylogeography and genetic variation of Triatoma dimidiata, the main Chagas disease vector in Central America, and its position within the genus Triatoma. PLoS Neglected Tropical Diseases 2(5): e233.
[Full Text] [PDF] [Suppl. Material]

Bracho, M.A.; Saludes, V.; Martro, E.; Bargallo, A.; González-Candelas, F.; Ausina, V. 2008. Complete genome of a European hepatitis C virus subtype 1g isolate: phylogenetic and genetic analyses. Virology Journal 5: 72.
[Abstract] [Provisional PDF]

Comas, I.; González-Candelas, F.; Zúñiga, M. 2008. Unravelling the evolutionary history of the phosphoryl-transfer chain of the phosphoenolpyruvate:phosphotransferase system by phylogenetic analyses and genome context. BMC Evolutionary Biology 8:147.
[Abstract] [PDF]

Cuevas, J. M.; Torres-Puente, M.; Jiménez-Hernández, N.; Bracho, M. A.; García-Robles, I.; Carnicer, F.; Olmo, J. D.; Ortega, E.; Moya, A.; González-Candelas, F. 2008. Refined analysis of genetic variability parameters in hepatitis C virus and the ability to predict antiviral treatment response. Journal of Viral Hepatitis. Online Early, May 12, 2008. doi:10.1111/j.1365-2893.2008.00991.x.
[Abstract] [Full Text] [PDF] [Suppl.Material]

Cuevas, J.M.; Torres-Puente, M.; Jiménez-Hernández, N.; Bracho, M.A.; García-Robles, I.; Wrobel, B.; Carnicer,F.; del Olmo, F.; Ortega,E.; Moya,A.; González-Candelas,F. (2008) Genetic variability of Hepatitis C Virus before and after combined therapy of interferon plus ri bavirin. PLoSONE 3(8): e3058. doi:10.1371/journal.pone.0003058. [Full Text] [PDF]

Sentandreu, V.; Jiménez-Hernández, N; Torres-Puente, M.; Bracho, M.A.; Valero, A.; Gosalbes, M.J.; Ortega, E.; Moya, A.; González-Candelas, F. (2008) Evidence of Recombination in Intrapatient Populations of Hepatitis C Virus. PLoS ONE 3(9): e3239. doi:10.1371/journal.pone.0003239. [Full Text] [PDF] [Supporting Information]

Torres-Puente, M.; Cuevas, J.N.; Jiménez-Hernández, N.; Bracho, M.A.; García-Robles, I.; Wróbel, B.;Carnicer, F.; del Olmo, F.; Ortega, E.; Moya, A.; González-Candelas, F. 2008. Using evolutionary tools to refine the new hypervariable region 3 within the envelope 2 protein of hepatitis C virus. Infection, Genetics and Evolution 8(1): 74-82. [PDF]

Torres-Puente, M.; Cuevas, J.N.; Jiménez-Hernández, N.; Bracho, M.A.; García-Robles, I.; Wrobel, B.; Carnicer, F.; del Olmo, F.; Ortega, E.; Moya, A.; González-Candelas, F. 2008. Genetic variability in hepatitis C virus and its role in antiviral treatment response. The Journal of Viral Hepatitis 15: 188-199. [Abstract] [PDF] [Suppl.Material]

Torres-Puente, M.; Cuevas, J.N.; Jiménez-Hernández, N.; Bracho, M.A.; García-Robles, I.; Carnicer, F.; del Olmo, F.; Ortega, E.; Moya, A.; González-Candelas, F. 2007. Hepatitis C virus and the controversial role of the interferon sensitivity determining region in the response to interferon treatment. The Journal of Medical Virology 80(2): 247-253. [Abstract] [PDF]

Jimenez-Hernandez N, Sentandreu V, Castro JA, Torres-Puente M, Bracho A, Garcia-Robles I, Ortega E, Del Olmo J, Carnicer F, Gonzalez-Candelas F, Moya A. 2008. Effect of antiviral treatment and host susceptibility on positive selection in hepatitis C virus (HCV). Virus Res. 2008 131(2): 224-232. [Abstract] [PDF]

Palop-Esteban, M.; Segarra-Moragues, J.G.; González-Candelas, F. 2007. Historical and biological determinants of genetic diversity in the highly endemic triploid sea lavender Limonium dufourii (Plumbaginaceae). Molecular Ecology 16(18):3814-27. [PDF]

Torres-Puente, M.; Cuevas, J.M.; Jiménez-Hernéndez, N.; Bracho, M.A.; García-Robles, I.; Carnicer, F.; del Olmo, J.;Ortega, E.; Moya, A.; González-Candelas, F. 2007. Contribution of insertions and deletions to the variability of hepatitis C virus populations. Journal of General Virology 88(8):2198-2203. [PDF]

Amadoz, A.; González-Candelas, F. 2007. epiPATH: an information system for the storage and management of molecular epidemiology data from infectious pathogens. BMC Infectious Diseases 7: 32.

Comas, I.; Moya, A.; González-Candelas, F. 2007. From phylogenetics to phylogenomics: the evolutionary relationships of insect endosymbiotic gamma-Proteobacteria as a test case. Systematic Biology 56(1): 1-16.

Comas, I.; Moya, A.; González-Candelas, F. 2007. Phylogenetic signal and functional categories in Proteobacteria genomes. BMC Evolutionary Biology 7 (Suppl. 1): S7 (8 February 2007)

Garcia-Martinez, J.; González-Candelas, F.; Perez-Ortin, J. 2007. Common gene expression strategies revealed by genome-wide analysis in yeast. Genome Biology 8:R222.
[Abstract] [PDF] [PubMed]
Highly accessedOpen Access

Coscollá, M.; González-Candelas, F. 2007. Population structure and recombination in environmental isolates of Legionella pneumophila. Environmental Microbiology 9(3): 643-656.

Segarra-Moragues J.G.; Palop-Esteban M.; González-Candelas F.; Catalán P. 2007. Nunatak survival vs. tabula rasa in the Central Pyrenees: a study on the endemic plant species Borderea pyrenaica (Dioscoreaceae). Journal of Biogeography 34: 1893-1906.
[Abstract] [PDF]

Jiménez-Hernández, N.; Torres-Puente, M.; Bracho, M. A.; García-Robles, I.; Ortega, E.; del Olmo, J.;Carnicer, F. González-Candelas, F.; Moya, A. 2007. Epidemic dynamics of two coexisting hepatitis C virus subtypes. Journal of General Virology 88(1): 123-133.
[Abstract] [PDF]

Comas, I.; Moya, A.; Azad, R.K.; Lawrence, J.G.; González-Candelas, F. 2006. The Evolutionary Origin of Xanthomonadales Genomes and the Nature of the Horizontal Gene Transfer Process. Molecular Biology and Evolution 23(11): 2049-2057.

Coscollá, M.; Gosalbes, M.J.; Catalán, V.; González-Candelas, F. 2006. Genetic variability in environmental isolates of Legionella pneumophila from Comunidad Valenciana (Spain). Environmental Microbiology 8 (6): 1056-1063.

Wróbel, B.; Torres-Puente, M.; Jiménez, N.; Bracho, M.A.; García-Robles, I.; Moya, A.; González-Candelas, F. 2006. Analysis of the overdispersed clock in the short term evolution of hepatitis C virus: using the E1/E2 gene sequences to infer infection dates in a single source outbreak. Molecular Biology and Evolution 23(6):1242-1253.

Czarna, A.; Sanjuán, R.; González-Candelas, F.; Wróbel, B. 2006.Topology testing of phylogenies using least squares methods. BMC Evolutionary Biology 6:105.

Bargues, M.D.; Klisiowicz, D.R.; Panzera, F.; Noireau, F.; Marcilla, A.; Perez, R.; Rojas, M.G.; O’Connor, J.E.; González-Candelas, F.; Galvão, C.; Jurberg, J.; Carcavallo, R.U.; Dujardin, J.P.; Mas-Coma, S. 2006. Origin and phylogeography of the Chagas disease main vector Triatoma infestans based on nuclear rDNA sequences and genome size. Infection, Genetics and Evolution 6(1): 46-62.

López-Labrador, F.-X.; Bracho, M.A.; Berenguer, M.; Coscolla, M.; Rayón, J.M.; Prieto, M.; Carrasco, D.; Gómez, M. D.; Moya, A.; González-Candelas, F. 2006. Genetic similarity of hepatitis C virus and fibrosis progression in chronic and recurrent infection after liver transplantation. Journal of Viral Hepatitis 13: 104-115.

Bracho, M.A.; Carrillo-Cruz, F.Y.; Ortega, E.; Moya, A.; González-Candelas, F. 2006. A new subtype of hepatitis C virus genotype 1: complete genome and phylogenetic relationships of an Equatorial Guinea isolate. Journal of General Virology 87 (6): 1697-1702.

Maria Alma Bracho, María José Gosalbes, Francisco González, Andrés Moya and Fernando González-Candelas. 2006. Molecular epidemiology and evolution in a fulminant hepatitis B virus outbreak. Journal of Clinical Microbiology 44(4): 1288-1294.

Pilar Catalán, José Gabriel Segarra-Moragues, Marisa Palop-Esteban, Carlos Moreno and Fernando González-Candelas. 2006. A Bayesian approach for discriminating among alternative inheritance hypotheses in plant polyploids: the allotetraploid origin of genus Borderea (Dioscoreaceae). Genetics 172(3): 1939-1953.

González-Candelas, F.; Moya, A. 2005. Time and rate of evolution are the key to establish transmission cases. AIDS 19(14): 1552-1553.

Zúñiga, M.; Comas,I.; Linaje, R.; Monedero, V.; Yebra, M. J.; Esteban, C.D.; Deutscher, J.;Pérez-Martínez, G.; González-Candelas, F. 2005. Horizontal Gene Transfer in the Molecular Evolution of Mannose PTS Transporters. Molecular Biology and Evolution 22(8):1673-1685. doi:10.1093/molbev/msi156
[Abstract] [Full Text] [PDF]

Bracho,M. A.; Gosalbes, M. J.; Blasco, D.; Moya, A.; González-Candelas, F. 2005. Molecular Epidemiology of a Hepatitis C Virus Outbreak in a Hemodialysis Unit. J. Clin. Microbiol. 43: 2750-2755.
[Abstract] [Full Text] [PDF]

Comas, I.; Moya, A.; González-Candelas, F. 2005. Validating viral quasispecies with digital organisms: a re-examination of the critical mutation rate. BMC Evolutionary Biology 5: 5.
[Abstract] [Full text] [PDF] [PubMed] [Related articles]

Segarra-Moragues, J.G.; Palop-Esteban, M.; González-Candelas; F. and Catalán, P.; 2005. On the verge of extinction: Genetics of the critically endangered Iberian plant species Borderea chouardii (Dioscoreaceae) and implications for conservation management. Molecular Ecology 14: 969-982.
[Abstract] [PDF] [Supplementary Material]

Bracho, M. A.; García-Robles, I.; Jiménez, N.; Torres-Puente, M.; Moya, A., and González-Candelas. F. 2004. Effect of oligonucleotide primers in determining viral variability within hosts. Virology Journal, 1:13.
[Abstract] [Full Text] [PDF] [PubMed] [Related articles]

Segarra-Moragues, J.G., M. Palop-Esteban, F. González-Candelas & P. Catalán. 2004. Characterization of seven (CTT)n microsatellite loci in the Pyrenean endemic Borderea pyrenaica (Dioscoreaceae). Remarks on ploidy level and hybrid origin assessed through allozymes and microsatellite analyses. Journal of Heredity 95(2):177–183.

Moya, A., E.C. Holmes & F.González-Candelas. 2004. Population genetics and the evolutionary epidemiology of RNA viruses. Nature Reviews in Microbiology 2: 279-288.

López-Labrador,F.X., X.-S. He, M. Berenguer, R.C. Cheung, F. González-Candelas, T.L. Wright y H.B. Greenberg. 2004. Genetic variability of hepatitis C virus non-structural protein 3 and virus-specific CD8+ response in patients with chronic hepatitis C. Journal of Medical Virology 72(4): 575-585.

Gil, R. , F. J. Silva, E. Zientz, F. Delmotte, F. González-Candelas, A. Latorre, C. Rausell, J. Kamerbeek, J. Gadau, B. Hölldobler, R. C. H. J. van Ham, R. Gross & A. Moya. 2003. The genome sequence of Blochmannia floridanus: comparative analysis of reduced genomes. Proceedings of the National Academy of Sciences USA 100(16): 9388-9393.

Segarra-Moragues, J.G., M. Palop-Esteban, F. González-Candelas & P. Catalán. 2003. Characterization of ten microsatellite loci in the critically endangered Pyrenean yam Borderea chouardii (Dioscoraceae). Molecular Ecology Notes 3(2): 265-267.

Torres-Puente, M., M.A. Bracho , N. Jimenez , I. Garcia-Robles , A. Moya & F. Gonzalez-Candelas. 2003. Sampling and repeatability in the evaluation of hepatitis C virus genetic variability. Journal of General Virology 84(9):2343-50.
[PubMed] [PDF]

González-Candelas, F., M.A. Bracho & A. Moya. 2003. Molecular epidemiology and forensic genetics: application to a hepatitis C virus transmission event in a hemodialysis unit. Journal of Infectious Diseases 187(3): 352-358.

Rodríguez, S., M. Palop, C. Palacios & F. González-Candelas. 2003. Molecular and morphological differentiation in Limonium dufourii (Plumbaginaceae), an endangered Mediterranean plant. Conservation Genetics 4(3): 383-391.

Zúñiga, M., G. Pérez & F. González-Candelas. 2002. Evolution of arginine deiminase (ADI) pathway genes. Molecular Phylogenetics and Evolution 25(3): 429-444.

Marín, I., M.A. Farés, F. González Candelas, E. Barrio & A. Moya. 2001. Detecting changes in the functional constraints of paralogous genes. Journal of Molecular Evolution 52(1): 17-28.

Palop, M., Palacios, C. & F. González Candelas. 2000. Development and across species transferability of microsatellite markers in the genus Limonium (Plumbaginaceae). Conservation Genetics 1(2): 177-179.

VanHam, R.C.H.J., F. González Candelas, F. Silva, A. Moya & A. Latorre. 2000. Post-symbiotic plasmid acquisition and evolution of the repA1-replicon in Buchnera aphidicola. Proceedings National Academy Sciences USA. 97(20): 10855-10860.

Sanchis, A., A. Latorre, F. González Candelas & J.M. Michelena. 2000. An 18S rDNA based molecular phylogeny of Aphidiinae (Himenoptera: Braconidae). Molecular Phylogenetics and Evolution 14 (2): 180-194.

Palacios, C., J.A. Rosselló & F. González Candelas. 2000. Study of the evolutionary relationships among Limonium species (Plumbaginaceae) using nuclear and cytoplasmic molecular markers. Molecular Phylogenetics and Evolution 14(2): 232-249.

Palacios, C., S. Kresovich & F. González Candelas. 1999. A population genetic study of the endandered plant species Limonium dufourii based on AFLPs. Molecular Ecology 8: 645-657.

Palacios, C. & F. González Candelas. 1997. Analysis of population genetic structure and variability using RAPD markers in the endemic and endangered Limonium dufourii (Plumbaginaceae). Molecular Ecology 6 (12): 1107-1121.

Phippen, W.B., S. Kresovich, F. González Candelas & J.R. McFerson. 1997. Molecular characterization can discriminate and partition variation among genebank holdings: a case study with phenotypically similar accessions of Brassica oleracea L. var. capitata (cabbage) 'Golden Acre'. Theoretical and Applied Genetics 94: 227-234.

Brown, S.M.; M.S. Hopkins, S.E. Mitchel, T.Y. Wang, S. Kresovich, R.R. Duncan, M.L. Senior & F. González Candelas. 1996. Multiple methods for the identification of polymorphic simple sequence repeats (SSRs) in sorghum [Sorghum bicolor (L.) Moench.?. Theor. Appl. Genet. 93: 190-198.

Elena, S.F., F. González-Candelas, I.S. Novella, E.A. Duarte, D.K. Clarke, E. Domingo, J.J. Holland & A. Moya. 1996. Evolution of fitness in experimental populations of Vesicular Stomatitis Virus. Genetics 142: 673-679.

Galiana, A., F. González Candelas y A. Moya. 1996. Postmating isolation analysis in founder-flush experimental populations of Drosophila pseudoobscura. Evolution 50 (2): 941-944.

González Candelas, F., S.F. Elena y A. Moya. 1995. Approximate variance of nucleotide divergence between two sequences when restriction fragment data are available. Genetics 140: 1443-1446.

Elena, S.F., F. González Candelas y A. Moya. 1992. Does the VP1 gene of foot-and-mouth disease virus behave as a molecular clock? Journal of Molecular Evolution 35: 223-229.

Martínez, M.A., Carrillo, C., González Candelas, F., Moya, A., Domingo, E. y Sobrino, F. 1991. Fitness alteration of foot-and-mouth disease virus mutants: measurement of the adaptability of viral quasispecies. Journal of Virology 65(7): 3954-3957.

Mueller, L.D., González Candelas, F. y Sweet, V.F. 1991. Components of density-dependent population dynamics: models and tests with Drosophila. The American Naturalist 137(4): 457-475.


Courses currently taught

The Tree of Life  (undergrad)

Evolutionary Processes and Mechanisms (undergrad)

Major Transitions in Evolution (undergrad)

Evolutionary Bioinformatics (Master in Bioinformatics, Univ. Valencia)

Evolutionary Genetics (Master in Integrative Evolutionary Biology, Univ. Valencia)

Molecular Epidemiology (Summer School, CSISP)


Previous teaching

Introduction to Evolutionary Theory (undergrad)

Molecular Population Genetics (graduate)

Systematics, Classification and Phylogeny (graduate)

Biological Basis for Conservation (graduate)

Phylogenography and Population Structure (graduate)

Molecular Phylogenetics (graduate level)

Bioinformatics (graduate level)




Mailing address

            Instituto Cavanilles de Biodiversidad y Biología Evolutiva
            Universidad de Valencia
            c/ Catedrático José Beltrán, 2
46180 – Paterna
            Valencia - Spain