Recent publications

Phenotypic variation

• Lucchini, N., Martínez-Freiría, F., Freitas, I., Zuazo, Ó., Dezetter, M., & Lourdais, O. (2025). Hydrothermal physiology and vulnerability to climatic change: insight from European vipers. Journal of Thermal Biology, 104115. DOI: 10.1016/j.jtherbio.2025.104115  

• Lakušić, M., Martínez-Freiría, F., Anđelković, M. & Hempel, B.-F. (2025) Beyond sexual maturity: Importance of dietary changes in venom variation in Vipera ammodytes. Toxicon, 2025: 108291. DOI: 10.1016/j.toxicon.2025.108291

• Buldain, J., Vitorino, R., Lima, T., Avella, I., Zuazo, Ó., & Martínez-Freiría, F. (2025). Intraspecific venom variation in the Iberian asp viper (Vipera aspis zinnikeri) across natural and intensive agricultural habitats. Journal of Proteomics, 310: 105337. DOI 10.1016/j.jprot.2024.105337 

• Lakušić, M., Damm, M., Bjelica, V., Anđelković, M., Tomović, L., Bonnet, X., ... & Martínez-Freiría, F. (2025). Ontogeny, not prey availability, underlies allopatric venom variability in insular and mainland populations of Vipera ammodytes. Journal of Proteomics, 310: 105320. DOI: 10.1016/j.jprot.2024.105320

• Freitas, I., Velo-Antón, G., Kaliontzopoulou, A., Zuazo, Ó., Martínez-Freiría, F. (2024). Association between Genetic Admixture and Morphological Patterns in a Hybrid Zone between the Two Iberian Vipers, Vipera aspis and V. latastei. Journal of Zoological Systematics and Evolutionary Research. DOI: 10.1155/2024/3800363

• Buldain, J., Kaliontzopoulou, A., Zuazo, Ó., Martínez-Freiría, F. (2024) Intensive agriculture increases fluctuating asymmetry of dorsal head shape in Iberian asp viper populations. Journal of Zoology. DOI: 10.1111/jzo.13203

• Damm, M., Avella, I., Merzara, R., Lucchini, N., Buldain, J., Corga, F., ... & Martínez-Freiría, F. (2024). Venom variation among the three subspecies of the North African mountain viper Vipera monticola Saint Girons 1953. Biochimie. DOI: 10.1016/j.biochi.2024.07.008

• Guillon M., Martínez-Freiría, F., Lucchini, N., …. & Lourdais, O. (2024). Inferring current and Last Glacial Maximum distributions are improved by physiology-relevant climatic variables in cold-adapted ectotherms. Journal of Biogeography. DOI: 10.1111/jbi.14828 

• Freitas, I., Tarroso, P., Zuazo, Ó., Zaldívar, R., Álvarez, J., Meijide-Fuentes, M., ... & Martínez-Freiría, F. (2023). Local niches explain coexistence in environmentally-distinct contact zones between Western Mediterranean vipers. Scientific Reports, 13(1), 21113. DOI: 10.1038/s41598-023-48204-3 

• Avella, I., Damm, M., Freitas, I., Wüster, W., Lucchini, N., Zuazo, Ó., Süssmuth, R. & Martínez-Freiría, F. (2023). One Size Fits All - Venomics of the Iberian Adder (Vipera seoanei, Lataste 1878) Reveals Low Levels of Venom Variation across Its Distributional Range. Toxins 15, 371. DOI: 10.3390/toxins15060371

• Scaramuzzi, A., Freitas, I., Sillero, N. & Martínez-Freiría, F. (2023). Meso-habitat distribution patterns and ecological requirements of two Mediterranean vipers depict weak competition in a contact zone. Journal of Zoology DOI: 10.1111/jzo.13087

• Oliveira, D., Halpern, B., Martínez-Freiría, F. & Kaliontzopoulou, A. (2023). Head Shape Heritability in the Hungarian Meadow Viper Vipera ursinii rakosiensis. Animals 13, 322. DOI: 10.3390/ani13020322

• Espasandín, I., Galán, P., & Martínez-Freiría, F. (2022). Sex, size and eco-geographic factors affect the feeding ecology of the Iberian adder, Vipera seoanei. Amphibia-Reptilia. DOI: 10.1163/15685381-bja10093

• Avella, I., Calvete, J. J., Sanz, L., Wüster, W., Licata, F., Quesada-Bernat, S., Rodríguez, Y. & Martínez-Freiría, F. (2022). Interpopulational variation and ontogenetic shift in the venom composition of Lataste's viper (Vipera latastei, Boscá 1878) from northern Portugal. Journal of Proteomics, 104613. DOI: 10.1016/j.jprot.2022.104613

• Martínez-Gil, H., Martínez-Freiría, F., Perera, A., Enriquez-Urzelai, U., Martínez-Solano, I., Velo-Antón, G. & Kaliontzopoulou, A. (2022). Morphological diversification of Mediterranean anurans: the roles of evolutionary history and climate. Biological Journal of the Linnean Society 165: 462-477. DOI: 10.1093/biolinnean/blab156

• Yusefi, H.G., Godinho, R., Khalatbari, L., Broomand, S., Fahimi, H., Martínez-Freiría, F. & Álvares, F. (2021) Habitat use and population genetics of golden jackals in Iran: Insights from a generalist species in a highly heterogeneous landscape. Journal of Zoological Systematics and Evolutionary Research 59: 1503-1515. DOI: 10.1111/jzs.12519

• Martínez-Castro, A., Kaliontzopoulou, A., Freitas, I. & Martínez-Freiría, F. (2021). Macroevolutionary variation and environmental correlates of scalation traits in Eurasian vipers (Serpentes: Viperinae). Biological Journal of the Linnean Society 132: 318-327. DOI: 10.1093/biolonnean/blaa197 

• Lucchini, N., Kaliontzopoulou, A., Val, G. A., & Martinez-Freiría, F. (2021). Sources of Intraspecific Morphological Variation in Vipera seoanei: Allometry, Sex, and Colour Phenotype. Amphibia-Reptilia 42: 1-16. DOI:10.1163/15685381-bja10024

• Martínez-Freiría, F., Toyama, K., Freitas I. & Kaliontzopoulou, A. (2020) Thermal melanism explains macroevolutionary variation of dorsal pigmentation in Eurasian vipers. Scientific Reports 10: 16122. DOI 10.1038/s41598-020-72871-1

• Martínez-Freiría, F., Lorenzo, M. & Lizana, M. (2019). Zamenis scalaris prefers abandoned citrus orchards in Eastern Spain. Ecological insights from a radio-tracking survey. Amphibia-Reptilia 40: 113-119. DOI: 10.1163/15685381-18000003

• Hemami, M.R., Esmaeili, S., Brito, J.C., Ahmadi, M., Omidi, M., & Martínez-Freiría, F. (2018). Using ecological models to explore niche partitioning within a guild of desert felids. Hystrix, the Italian Journal of Mammalogy 29: 216-222. DOI: 10.4404/hystrix-00042-2017

• Kaliontzopoulou, A., Pinho, C. & Martínez-Freiría, F. (2018). Where does diversity come from? Linking geographical patterns of morphological, genetic, and environmental variation in wall lizards. BMC Evolutionary Biology 18: 124. DOI: 10.1186/s12862-018-1237-7 

Evolutionary history

• Ambu, J., Litvinchuk, S. N., Caballero‐Díaz, C., Nicieza, A., Velo‐Antón, G., Gonçalves, H., ... & Dufresnes, C. (2025). Genomic, phenotypic and environmental correlates of speciation in the midwife toads (Alytes). Molecular Ecology, 34(8), e17736. DOI: 10.1111/mec.17736 

• Dufresnes, C., Ghielmi, S., Halpern, B., Martínez-Freiría, F., Mebert, K., Jelić, D., ... & Ursenbacher, S. (2024). Phylogenomic insights into the diversity and evolution of Palearctic vipers. Molecular Phylogenetics and Evolution, 197, 108095. DOI: 10.1016/j.ympev.2024.108095 

• Liz, A. V., Mochales-Riaño, G., Velo-Antón, G., García-Cardenete, L., Brito, J. C., Carranza, S., & Martínez-Freiría, F. (2024). Historical biogeography of the Saharan horned viper enlightens past dynamics of hyperarid desert habitats. Journal of Arid Environments, 222, 105171. DOI: 10.1016/j.jaridenv.2024.105171 

• Freitas, I., Velo-Antón, G., Lopes, S., Muñoz-Merida, A., & Martínez-Freiría, F. (2024). Isolation and characterization of polymorphic microsatellite loci for the three Iberian vipers, Vipera aspis, V. latastei and V. seoanei by Illumina MiSeq sequencing. Molecular Biology Reports, 51(1), 294. DOI: 10.1007/s11033-024-09263-5 

• Mochales-Riaño, G., Burriel-Carranza, B., Barros, M. I., Velo-Antón, G., Talavera, A., Spilani, L., ... & Martínez-Freiría, F. (2024). Hidden in the sand: Phylogenomics unravel an unexpected evolutionary history for the desert-adapted vipers of the genus Cerastes. Molecular Phylogenetics and Evolution, 191, 107979. DOI: 10.1016/j.ympev.2023.107979 

• Lucchini, N., Kaliontzopoulou, A., Lourdais, O., & Martínez‐Freiría, F. (2023). Climatic adaptation explains responses to Pleistocene oscillations and diversification in European vipers. Journal of Biogeography, 50(11), 1838-1851. DOI: 10.1111/jbi.14694 

• Romero-Iraola, I., Freitas, I., Jiménez-Ruíz, Y., Geniez, P., García-París, M., & Martínez-Freiría, F. (2023). Phylogeographic and paleoclimatic modelling tools improve our understanding of the biogeographic history of Hierophis viridiflavus (Colubridae). Animals, 13(13), 2143. DOI: 10.3390/ani13132143 

• Enriquez‐Urzelai, U., Martínez‐Freiría, F., Freitas, I., Perera, A., Martínez‐Solano, Í., Salvi, D., ... & Kaliontzopoulou, A. (2022). Allopatric speciation, niche conservatism and gradual phenotypic change in the evolution of European green lizards. Journal of Biogeography, 49: 2193-2205. DOI: 10.1111/jbi.14497

• Velo-Antón, G., Henrique, M., Liz, A. V., Martínez-Freiría, F., Pleguezuelos, J. M., Geniez, P., ... & Brito, J. C. (2022). DNA barcode reference library for the West Sahara-Sahel reptiles. Scientific Data, 9: 459. DOI: 10.1038/s41597-022-01582-1

• Martínez-Freiría, F., Freitas, I., Velo-Antón, G., Lucchini, N., Fahd, S., Larbes, S., Pleguezuelos, J.M, Santos, X., & Brito, J.C. (2021). Integrative taxonomy reveals two species and intraspecific differentiation in the Vipera latastei–monticola complex. Journal of Zoological Systematics and Evolutionary Research. DOI: 10.1111/ jzs.12534

• Pizzigalli, C., Pierre-André, C., Geniez, P., Martínez-Freiría, F., Velo-Antón, G. & Brito, J.C. (2021) Phylogeographic diversification of the Mesalina olivieri species complex (Squamata: Lacertidae) with the description of a new species and a new subspecies endemic from North West Africa. Journal of Zoological Systematics and Evolutionary Research. DOI: 10.1111/jzs.12516

• Chamorro, D., Martínez-Freiría, F., Real, R. & Muñoz, A.-R. (2021). Understanding parapatry: How do environment and competitive interactions shape Iberian vipers’ distributions? Journal of Biogeography. DOI: 10.1111/jbi.14078

• Torres-Roig, E., Mitchell, K.J., Alcover, J.A., Martínez-Freiría, F., Bailón, S., Heiniger, H., Williams, M., Cooper, A., Pons, J. & Bover, P. (2020). Origin, extinction and ancient DNA of a new fossil insular viper: molecular clues of overseas immigration. Zoological Journal of the Linnean Society. DOI: 10.1093/zoolinnean/zlaa094/5909523

• Martínez-Freiría, F., Freitas, I., Zuffi, M.A.L., Golay, P., Ursenbacher, S. & Velo-Antón, G. (2020) Climatic refugia boosted allopatric diversification in western Mediterranean vipers. Journal of Biogeography 47: 1698-1713. DOI: 10.1111/jbi.13861 

• Freitas, I., Ursenbacher, S., Mebert, K., Zinenko, O., Schweiger, S., Wüster, W., Brito, J.C., Crnobrnja-Isailović, J., Halpern, B., Fahd, S., Santos, X., Pleguezuelos, J.M., Joger, U., Orlov, N., Mizsei, E., Lourdais, O., Zuffi, M.A.L., Strugariu, A., Zamfirescu, Ş.R., Martínez-Solano, Í., Velo-Antón, G., Kaliontzopoulou, A. & Martínez-Freiría, F. (2020) Evaluating taxonomic inflation: towards evidence-based species delimitation in Eurasian vipers (Serpentes: Viperinae). Amphibia-Reptilia 41: 285–311. DOI: 10.1163/15685381-bja10007

• Moutinho, F., Serén, N., Paupério, J., Silva, T.L., Martínez-Freiría, F., Sotelo, G., Faria, R., Mappes, T., Alves, P-C-, Brito, J.C. & Boratynski, Z. (2020). Evolutionary history of two cryptic species of northern African jerboas. BMC Evolutionary Biology 20. DOI: 10.1186/s12862-020-1592-z

• Dinis, M., Merabet, K., Martínez-Freiría, F., Steinfartz, S., Vences, M., Burgon, J. D., Elmer, K.M., Donaire, D., Hinckley, A., Fahd, S., Joger, U., Fawzi, A., Slimani, T. & Velo-Antón, G. (2019). Allopatric diversification and evolutionary melting pot in a North African Palearctic relict: The biogeographic history of Salamandra algira. Molecular Phylogenetics and Evolution 130: 81-91. DOI: 10.1016/j.ympev.2018.10.018

• Velo-Antón, G., Martínez-Freiría, F., Pereira, P., Crochet, P.-A. & Brito J.C. (2018). Living on the edge: ecological and genetic connectivity of the spiny-footed lizard, Acanthodactylus aureus, confirms the Atlantic Sahara desert as a biogeographic corridor and centre of lineage diversification. Journal of Biogeography 45: 1031-1042. DOI: 10.1111/jbi.13176

• Gonçalves D. V., Martínez-Freiría, F., Crochet, P. A., Geniez, P., Carranza, S. & Brito, J.C. (2018). The role of climatic cycles and trans-Saharan migration corridors in species diversification: biogeography of Psammophis schokari group in North Africa. Molecular Phylogenetics and Evolution 118: 64–74. DOI: 10.1016/j.ympev.2017.09.009

Conservation

• Carmona-González, R., Carro, F., González de la Vega, J.P., & Martínez-Freiría, F. (2024). Temporal Range Dynamics of the Lataste’s Viper (Vipera latastei Boscá, 1878) in Doñana (Spain): Insights into Anthropogenically Driven Factors. Animas 14, 3024. DOI: 10.3390/ani14203025

• Martínez-Freiría, F., Fahd, S., Bouazza, A., Crochet, P.-A. & Beddek, M. 2024. Vipera monticola. The IUCN Red List of Threatened Species 2024: e.T221202628A3105206. 

• Mebert, K., Cattaneo, A., Ioannidis, I., Strugariu, A., Stümpel, N. & Martínez-Freiría, F. 2024. Macrovipera schweizeri. The IUCN Red List of Threatened Species 2024: e.T12654A137843008.

• Martínez-Freiría, F., Santos, X. & Pleguezuelos, J.M. 2024. Vipera latastei. The IUCN Red List of Threatened Species 2024: e.T221202039A137859691. 

• Lucchini, N., Freitas, I., Avella, I. & Martínez-Freiría, F. 2024. Vipera seoanei. The IUCN Red List of Threatened Species 2024: e.T61594A137859941. 

• Ursenbacher, S., Bowles, P., Martínez-Freiría, F., Pleguezuelos, J.M., Joger, U., Zuffi, M.A.L. & Crochet, P.-A. 2024. Vipera aspis. The IUCN Red List of Threatened Species 2024: e.T61591A137859549.

• Brito, J. C., Sow, A. S., Vale, C. G., Pizzigalli, C., Hamidou, D., Gonçalves, D.V., Martínez-Freiría, F., ... & Álvares, F. (2022). Diversity, distribution and conservation of land mammals in Mauritania, North-West Africa. Plos one, 17(8): e0269870 DOI: 10.1371/journal.pone.0269870 

• Naia, M., Tarroso, P., Sow, A.S., Liz, A.V., Gonçalves, D.V., Martínez-Freiría, F., Santarém, F., Yusefi, G.H., Velo-Antón, G., Avella, I., Hanson, J.O., Khalatbari, L., Silva, M.J.F., Camacho-Sanchez, M., Boraynski, I., Carvalho, S.B. & Brito, J.C. (2021). Potential negative effects of the Green Wall on Sahel's biodiversity. Conservation Biology. DOI: 10.1111/cobi.13755

• Sampaio, M., Velo-Antón, G., Martínez-Freiría, F., Sánchez-Vialas, A., Pleguezuelos, J.M., Geniez, P., Crochet, P.-A. & Brito, J.C. (2021) Beyond the comfort zone: Amphibian diversity and distribution in the West Sahara-Sahel using mtDNA and nuDNA barcoding and spatial modelling. Conservation Genetics. DOI: 10.1007/s10592-021-01331-8

• Goudarzi, F., Hemami, M.-R., Makelian, M., Fakheran, S. & Martínez-Freiría, F. (2021). Species versus within-species niches: a multi-modelling approach to assess range size of a spring-dwelling amphibian. Scientific Reports, 11: 597. DOI: 10.1038/s41598-020-79783-0

• Campos, J., Bernhardt, J., Aquilué, N., Brotons, L., Domínguez, J., Lomba, Á., Marcos, B., Martínez-Freiría, F., Moreira, F., Pais, S., Honrado, J.P. & Regos, A. (2021) Using fire to enhance rewilding when agricultural policies fail. Science of Total Environment, 755: 142897. DOI: 10.1016/j.scitotenv.2020.142897

• Pais, S., Aquilué, N., Campos, J., Sila, Â., Marcosa, B., Martínez-Freiría, F., Domínguez, J., Brotons, L., Honrado, J.P. & Regos, A. (2020). Mountain farmland protection and fire-smart management jointly reduce fire hazard and enhance biodiversity and carbon sequestration. Ecosystem Services, 44: 101143. DOI: 10.1016/j.ecoser.2020.101143

• Khalatbari, L., Yusefi, G.H., Martínez-Freiría, F., Jowkar, H. & Brito, J.C. (2018). Availability of prey and natural habitats are related with temporal dynamics in range and habitat suitability for Asiatic Cheetah. Hystrix, the Italian Journal of Mammalogy 29: 145–151. DOI: 10.4404/hystrix-00080-2018

• Brito, J.C., Durant, S.M., Pettorelli, N., Newby, J., Canney, S., Algadafi, W., Rabeil, T., Crochet, P.A., Pleguezuelos, J.M., Wacher, T., de Smet, K., Gonçalves, D.V., Ferreira da Silva, M.J., Martínez-Freiría, F., Abáigar, T., Campos, J.C., Comizzoli, P., Fahd, S., Fellous, A., Garba, A.H.M., Hamidou, D., Harouna, A., Hatcha, M.H., Nagy, A., Silva, T.L., Sow, A.S., Vale, C.G., Boratyński, Z., Rebelo, H. & Carvalho, S.B. (2018). Armed conflicts and wildlife decline: Challenges and recommendations for effective conservation policy in the Sahara‐Sahel. Conservation Letters e12446. DOI: 10.1111/conl.12446

• Freitas, I., Fahd, S., Velo-Antón, G. & Martínez-Freiría, F. (2018). Chasing the phantom: biogeography and conservation of Vipera latastei-monticola in the Maghreb (North Africa). Amphibia-Reptilia 39: 145-161. DOI: 10.1163/15685381-17000197

Distribution

• Lobón-Rovira, J., Sierra, P., Portas, R., Vaz Pinto, P., & Martínez-Freiría, F. (2025) Noteworthy herpetological notes from Namibe Province, Angola. Herpetology Notes 18: 99–105.

• Martínez-Freiría, F., Bouazza, A., Buldain, J., Freitas, I., Avella, I., Scaramuzzi, A., Sioumpora, K., Oliveira, D. & Fahd, S. (2023) Fieldwork campaigns and citizen science data increase the distributional range of the elusive Vipera monticola in Morocco. Basic & Applied Herpetology 37

• Scaramuzzi, A., Avella, I., Lansari, A., Bouazza, A. & Martínez-Freiría, F. (2022) New high-elevation records of Malpolon monspessulanus and Hemorrhois hippocrepis from the High Atlas Mountains (Morocco). Boletín de la Asociación Herpetológica Española 33

• Vázquez, R. & Martínez-Freiría, F. (2021) First record of autumnal mating in the iberian adder, Vipera seoanei. Boletín de la Asociación Herpetológica Española 32 (1).

• Argaz, H., Brito, J.C., Fahd, S., Martínez-Freiría, F., Boudajbir, C. & Geniez, P. (2020) Atlas des amphibiens et reptiles endémiques du Maroc: répartition et état de conservation. Bull. Soc. Herp. Fr. 176: 61-94.

• Avella, I., Lucchini, N., Enriquez-Urzelai, U., Corga, F. & Martínez-Freiría, F. (2019). New records for the reptile fauna of the Tichka plateau (western High Atlas, Morocco). Boletín de la Asociación Herpetológica Española, 30(2): 42-47.

• Zuazo, O., Freitas, I., Zaldívar, R. & Martínez-Freiría, F. (2019). Coexistence and intermediate morphological forms between Vipera aspis and V. latastei in the intensive agriculture fields of north-western Iberian System. Boletín de la Asociación Herpetológica Española, 30(1): 35-41.

Other topics

• Sánchez-Vialas, A., Buckley, D., Recuero, E., Martínez-Freiría, F., Velo-Antón, G., Bisbal-Chinesta, J.F., Nartínez-Solano, I. (2024). Species list of the Spanish herpetofauna: an update. Basic and Applied Herpetology 38: 5-23. 

• Blanvillain, G., Lorch, J. M., Joudrier, N., Bury, S., Cuenot, T., Franzen, M., ... & Hoyt, J. R. (2024). Contribution of host species and pathogen clade to snake fungal disease hotspots in Europe. Communications biology, 7(1), 440. DOI: 10.1038/s42003-024-06092-x 

• Blanvillain, G., Martínez-Freiría, F., Hoyt, J. R., Lorch, J. M., & Martinez-Silvestre, A. (2024). Paranannizziopsis spp. Infection in Wild Vipers, Europe. Emerging Infectious Diseases, 30(5), 1000. DOI: 10.3201/eid3005.231317 

• Roca, V., Gómez-Ramírez, F., Espasandín, I., Megía-Palma, R., Perera, A. & Martínez-Freiría (2023). First helminthological data on the Iberian adder, Vipera seoanei. Parasitology Research DOI: 10.1007/s00436-023-07849-9

• Avella, I., Wüster, W., Luiselli, L., & Martínez-Freiría, F. (2022). Toxic Habits: An Analysis of general trends and biases in snake venom research. Toxins 14: 884. DOI: 10.3390/toxins14120884

• Valdoleiros, S.R., Gonçalves, I.C., Silva, C., Guerra, D., Silva, L.A., Martínez-Freiría, F., Rato, F. & Xará, S. (2021). Animais Venenosos em Território Português: Abordagem Clínica de Picadas e Mordeduras. Acta Medica Portuguesa 34(11): 784-795. DOI: 10.20344/amp.15589

General audience

• Martínez-Freiría, F. (2025). Red de Seguimiento de poblaciones de víboras ibéricas. Boletín UICN, primavera 2025.

• Tejado Leseros, C. & Martínez-Freiría, F. (2024) Víboras ibéricas. AHE

• Martínez-Freiría, F., Brito, J.C., Freitas, I., Pleguezuelos, J.M., Santos, X. &  Velo-Antón, G. (2023). Luces y sombras en la biología de la víbora hocicuda. Quercus, 452: 20-27.

• Martínez-Freiría, F. & Velo-Antón, G. (2023). Estructura genética y estado de conservación de la víbora hocicuda (Vipera latastei) y la víbora del Magreb (Daboia mauritanica) en el Norte de África. Instituto de Estudios Ceutíes, Ceuta, España.