{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T18:09:16Z","timestamp":1784830156845,"version":"3.55.0"},"reference-count":58,"publisher":"MDPI AG","issue":"8","license":[{"start":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T00:00:00Z","timestamp":1784764800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["J. Fungi","JoF"],"abstract":"<jats:p>Mainland Portugal has a lichenological tradition spanning more than three centuries, yet it still lacks a modern, critically revised checklist of its lichen biota. This gap is particularly relevant given the increasing role of species checklists as authoritative references for DNA metabarcoding applications and the expansion of large biodiversity datasets, including those derived from citizen science initiatives. Here we present the first comprehensive checklist of lichens and lichenicolous fungi recorded from Mainland Portugal, based on the critical assessment of 43,718 bibliographic records drawn from 381 publications issued between 1788 and 2021. A total of 2012 species and infraspecific taxa are documented from a universe of 4440 taxonomic names, including synonyms. Nomenclature was updated in accordance with the ITALIC 8.0 database. Species distributions are provided by province, following the scheme adopted in Portuguese botanical floras, with additional information on substrates and altitudinal ranges where available. With over 2000 taxa recorded, Mainland Portugal can be considered a lichen-rich territory relative to its land area, comparing favourably with other European checklists. We discuss the factors underlying this diversity, with particular emphasis on the biogeographical position of Portugal at the south-western edge of the European continent.<\/jats:p>","DOI":"10.3390\/jof12080549","type":"journal-article","created":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T17:13:17Z","timestamp":1784826797000},"page":"549","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Checklist of Lichens and Lichenicolous Fungi from Mainland Portugal"],"prefix":"10.3390","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-3661-6272","authenticated-orcid":false,"given":"Palmira","family":"Carvalho","sequence":"first","affiliation":[{"name":"Museu Nacional de Hist\u00f3ria Natural e da Ci\u00eancia, Universidade de Lisboa, Rua da Escola Polit\u00e9cnica 58, 1250-102 Lisboa, Portugal"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5265-7459","authenticated-orcid":false,"given":"Mar\u00eda Eugenia","family":"L\u00f3pez de Silanes","sequence":"additional","affiliation":[{"name":"Departamento de Enxe\u00f1ar\u00eda dos Recursos Naturais e Medio Ambiente, Escola de Enxe\u00f1ar\u00eda Forestal, Universidade de Vigo, 36005 Pontevedra, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3724-8483","authenticated-orcid":false,"given":"Graciela","family":"Paz-Berm\u00fadez","sequence":"additional","affiliation":[{"name":"Departamento de Enxe\u00f1ar\u00eda dos Recursos Naturais e Medio Ambiente, Escola de Enxe\u00f1ar\u00eda Forestal, Universidade de Vigo, 36005 Pontevedra, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3702-9782","authenticated-orcid":false,"given":"Joana","family":"Marques","sequence":"additional","affiliation":[{"name":"CIBIO\u2014Centro de Investiga\u00e7\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, InBIO Laborat\u00f3rio Associado, Campus de Vair\u00e3o, Universidade do Porto, 4485-661 Vair\u00e3o, Portugal"},{"name":"BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO\u2014Centro de Investiga\u00e7\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Campus de Vair\u00e3o, 4485-661 Vair\u00e3o, Portugal"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8351-4028","authenticated-orcid":false,"given":"Rui","family":"Figueira","sequence":"additional","affiliation":[{"name":"CIBIO\u2014Centro de Investiga\u00e7\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, InBIO Laborat\u00f3rio Associado, Campus de Vair\u00e3o, Universidade do Porto, 4485-661 Vair\u00e3o, Portugal"},{"name":"CIBIO\u2014Centro de Investiga\u00e7\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, InBIO Laborat\u00f3rio Associado, Instituto Superior de Agronomia, Universidade de Lisboa, Tapada da Ajuda, 1349-017 Lisboa, Portugal"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,7,23]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"5146","DOI":"10.5943\/mycosphere\/15\/1\/25","article-title":"The 2024 Outline of Fungi and fungus-like taxa","volume":"15","author":"Hyde","year":"2024","journal-title":"Mycosphere"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"136002","DOI":"10.65390\/fdiv.2026.136002","article-title":"Global polypore diversity and distribution patterns","volume":"136","author":"Zhao","year":"2026","journal-title":"Fungal Divers."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"R502","DOI":"10.1016\/j.cub.2009.04.034","article-title":"Lichens","volume":"19","author":"Lutzoni","year":"2009","journal-title":"Curr. Biol."},{"key":"ref_4","unstructured":"Coppins, B.J. (2002). Checklist of Lichens of Great Britain and Ireland, British Lichen Society."},{"key":"ref_5","unstructured":"Nimis, P.L. (2025). The Information System on Italian Lichens, University of Trieste, Dept. of Biology. Available online: https:\/\/italic.units.it\/."},{"key":"ref_6","first-page":"1","article-title":"Checklist of the lichen and lichenicolous fungi of Greece","volume":"103","author":"Abbott","year":"2009","journal-title":"Bibl. Lichenol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"400","DOI":"10.13158\/heia.34.2.2021.400","article-title":"Checklist of the lichens and lichenicolous fungi of Tunisia","volume":"34","author":"Seaward","year":"2021","journal-title":"Herzogia"},{"key":"ref_8","unstructured":"Y\u00e1nez-Ayabaca, A., Benitez, A., Mollina, R.B., Naranjo, D., Etayo, J., Prieto, M., Sol\u00f3rzano, G.C., Caicedo, E., Scharnagl, K., and McNerlin, B. (2025, November 20). Lichens, Lichenicolous and Allied Fungi of Ecuador, Version #01. Consortium of Lichen Herbaria. Symbiota Checklist. Available online: https:\/\/lichenportal.org\/portal\/checklists\/checklist.php?clid=1283&pid."},{"key":"ref_9","unstructured":"(2025, November 14). GBIF Occurrence Download. Available online: https:\/\/www.gbif.org."},{"key":"ref_10","unstructured":"(2025, October 12). Catalogue of Life. Catalogue of Life Annual Checklist 2024. Available online: https:\/\/www.catalogueoflife.org."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1423","DOI":"10.1111\/j.1523-1739.2007.00827.x","article-title":"The effectiveness of Iberian protected areas in conserving terrestrial biodiversity","volume":"21","author":"Lobo","year":"2007","journal-title":"Conserv. Biol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1121","DOI":"10.1007\/s10531-016-1289-1","article-title":"Case study of the implications of climate change for lichen diversity and distributions","volume":"26","author":"Psomas","year":"2017","journal-title":"Biodivers. Conserv."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1017\/S0024282916000153","article-title":"Seven species of Graphis from Portugal reported new to Europe","volume":"48","author":"Lepista","year":"2016","journal-title":"Lichenologist"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1556\/034.59.2017.3-4.11","article-title":"Lichens and lichenicolous fungi of Estremadura, Portugal, collected in 2015","volume":"59","year":"2017","journal-title":"Acta Bot. Hung."},{"key":"ref_15","unstructured":"Burgaz, A.R., and Mart\u00ednez, I. (2003). Peltigerales: Lobariaceae, Nephromataceae, Peltigeraceae, Sociedad Espa\u00f1ola de Liquenolog\u00eda."},{"key":"ref_16","first-page":"149","article-title":"Checklist and three new species of lichens and lichenicolous fungi of the Algarve (Portugal)","volume":"64","author":"Giralt","year":"2012","journal-title":"Sydowia"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1017\/S0024282913000261","article-title":"A Peltula Nyl. diversity hotspot in north-east Portugal, with one species new to science and three species new to mainland Europe","volume":"45","author":"Marques","year":"2013","journal-title":"Lichenologist"},{"key":"ref_18","unstructured":"Vandelli, D. (1789). Viridarium Grisley Lusitanicum, Linnaeanis Nominibus Illustratum, Jussu Academiae in Lucem Editum, Ex Typographia Regalis Academiae Scientiarum Olisiponensis."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Brotero, F.A. (1804). Flora Lusitanica, seu, Plantarum, Quae in Lusitania vel Sponte Crescunt, vel Frequentius Coluntur, ex Florum Praesertim Sexubus Systematice Distributarum, Synopsis, Ex Typographia Regia.","DOI":"10.5962\/bhl.title.45110"},{"key":"ref_20","first-page":"241","article-title":"Lichenes Lusitaniae","volume":"16","author":"Arnold","year":"1868","journal-title":"Flora"},{"key":"ref_21","first-page":"1","article-title":"Contribui\u00e7\u00e3o para o estudo das Parmeli\u00e1ceas portuguesas","volume":"1","author":"Tavares","year":"1945","journal-title":"Port. Acta Biol."},{"key":"ref_22","first-page":"123","article-title":"L\u00edquenes da Serra do Ger\u00eas. Cat\u00e1logo","volume":"12","author":"Tavares","year":"1950","journal-title":"Agron. Lusit."},{"key":"ref_23","first-page":"199","article-title":"Novas esp\u00e9cies de l\u00edquenes para a flora portuguesa","volume":"1","author":"Tavares","year":"1951","journal-title":"Rev. Fac. Ci\u00eanc. Lisb."},{"key":"ref_24","first-page":"199","article-title":"Epiphytic lichens of the Algarve, Portugal","volume":"14","author":"Jones","year":"1985","journal-title":"Port. Acta Biol. S\u00e9r. B"},{"key":"ref_25","first-page":"51","article-title":"Notes on the distribution and composition of epiphytic lichen communities with Nephroma laevigatum Ach. Portugal","volume":"18","author":"Jones","year":"1999","journal-title":"Port. Acta Biol. S\u00e9r. B"},{"key":"ref_26","first-page":"71","article-title":"Twelve new lichens for Portugal collected from the Serra da Estrela","volume":"13","author":"Aptroot","year":"1992","journal-title":"Cryptogam. Bryol. Lich."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1127\/nova.hedwigia\/63\/1996\/145","article-title":"Contribution to the flora of Portugal, lichens and lichenicolous fungi I","volume":"63","author":"Giralt","year":"1996","journal-title":"Nova Hedwig."},{"key":"ref_28","first-page":"5","article-title":"Checklist of the lichens and lichenicolous fungi of the Iberian Peninsula and Balearic Islands","volume":"14","author":"Llimona","year":"2001","journal-title":"Bocconea"},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Vieira, G., Z\u00eazere, J., and Mora, C. (2020). The climate of Portugal. Landscapes and Landforms of Portugal, Springer. World Geomorphological Landscapes.","DOI":"10.1007\/978-3-319-03641-0"},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Wieczorek, J., Bloom, D., Guralnick, R., Blum, S., D\u00f6ring, M., Giovanni, R., Robertson, T., and Vieglais, D. (2012). Darwin Core: An Evolving Community-Developed Biodiversity Data Standard. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0029715"},{"key":"ref_31","unstructured":"Delpeuch, A., Morris, T., Huynh, D., Mazzocchi, S., Guidry, T., Stephens, O., Matsunami, I., Sproat, I., Santos, S., and Larsson, A. (2026, June 16). OpenRefine\/OpenRefine: 3.10.1. Zenodo 2026. Available online: https:\/\/zenodo.org\/records\/18868465."},{"key":"ref_32","unstructured":"Chapman, A.D., and Wieczorek, J.R. (2020). Georeferencing Best Practices, GBIF Secretariat."},{"key":"ref_33","unstructured":"Wieczorek, J. (2026, June 16). tdwg\/dwc: 2023-07-10. Zenodo 2023. Available online: https:\/\/zenodo.org\/records\/8132249."},{"key":"ref_34","unstructured":"GBIF (2026, April 12). GBIF Registered Extensions 2026. Available online: https:\/\/rs.gbif.org\/extensions.html."},{"key":"ref_35","unstructured":"Carvalho, P., L\u00f3pez de Silanes, M.E., Paz-Berm\u00fadez, G., Marques, J., and Figueira, R. (2026). Checklist of Lichens and Lichenicolous Fungi from Mainland Portugal, Checklist dataset."},{"key":"ref_36","first-page":"193","article-title":"Lichens, lichenicolous and allied fungi of Germany\u2014A revised checklist","volume":"35","author":"Printzen","year":"2022","journal-title":"Herzogia"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Martellos, S., Conti, M., and Nimis, P.L. (2023). Aggregation of Italian lichen data in ITALIC 7.0. J. Fungi, 9.","DOI":"10.3390\/jof9050556"},{"key":"ref_38","unstructured":"Clerc, P., and Blaise, P. (2026, March 20). Catalogue des Lichens et des Champignons Lich\u00e9nicoles de Suisse, Available online: https:\/\/catlich.ch."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Martellos, S., d\u2019Agostino, M., Chiarucci, A., Nimis, P.L., and Nascimbene, J. (2020). Lichen distribution patterns in the ecoregions of Italy. Diversity, 12.","DOI":"10.3390\/d12080294"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"992","DOI":"10.1111\/ele.12624","article-title":"Multidimensional biases, gaps and uncertainties in global plant occurrence information","volume":"19","author":"Meyer","year":"2016","journal-title":"Ecol. Lett."},{"key":"ref_41","first-page":"57","article-title":"Flora liqu\u00e9nica do Parque Natural da Serra de S. Mamede","volume":"17","author":"Carvalho","year":"1997","journal-title":"Port. Acta Biol. S\u00e9r. B Sist."},{"key":"ref_42","first-page":"213","article-title":"Estudo da polui\u00e7\u00e3o atmosf\u00e9rica no estu\u00e1rio do Tejo. A vegeta\u00e7\u00e3o epif\u00edtica como bioindicadora","volume":"14","year":"1985","journal-title":"Port. Acta Biol. B"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"153","DOI":"10.1016\/S1290-0796(00)00110-3","article-title":"Cryptogamic epiphytes as indicators of air quality around an industrial complex in the Tagus valley, Portugal: Factor analysis and environmental variables","volume":"21","author":"Carvalho","year":"2000","journal-title":"Cryptogam. Bryol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1007\/BF00395121","article-title":"The Sines industrial complex monitoring programme: A preliminary report","volume":"1","author":"Jones","year":"1981","journal-title":"Environ. Monit. Assess."},{"key":"ref_45","unstructured":"Llimona, X., Lumbsch, H.T., and Ott, S. (2002). Dynamics of epiphytic lichen communities in an industrial area of Portugal. Progress and Problems in Lichenology at the Turn of the Millennium, Bibliotheca Lichenologica, J. Cramer."},{"key":"ref_46","unstructured":"Pereira, M., Branquinho, C., and Soares, A. (1995). Modelling air pollution in a copper mine by using physical and biological monitoring. Proceedings of the 3rd International Conference on Air Pollution, WIT Press."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1016\/S0098-8472(97)00015-4","article-title":"The cellular location of Cu in lichens and its effects on membrane integrity and chlorophyll fluorescence","volume":"38","author":"Branquinho","year":"1997","journal-title":"Environ. Exp. Bot."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1093\/sysbio\/syr030","article-title":"Determining species boundaries in a world full of rarity: Singletons, species delimitation methods","volume":"61","author":"Lim","year":"2012","journal-title":"Syst. Biol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1","DOI":"10.3897\/mycokeys.31.23568","article-title":"The lichens of the Alps\u2014An annotated checklist","volume":"31","author":"Nimis","year":"2018","journal-title":"MycoKeys"},{"key":"ref_50","first-page":"57","article-title":"Annotated checklist of Hong Kong lichens","volume":"17","author":"Aptroot","year":"1999","journal-title":"Trop. Bryol."},{"key":"ref_51","first-page":"279","article-title":"Lichens and lichenicolous fungi in the Montesinho Natural Park, the Serra da Nogueira and the Rio Sabor Valley (Portugal)","volume":"30","author":"Arroyo","year":"2009","journal-title":"Cryptogam. Mycol."},{"key":"ref_52","first-page":"37","article-title":"New and interesting lichen records for the Portuguese funga from the Upper Douro region (north-east Portugal)","volume":"23","author":"Marques","year":"2014","journal-title":"Austrian J. Mycol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"47","DOI":"10.3114\/persoonia.2025.54.02","article-title":"A novel, exclusively lichen-inhabiting lineage of Hypocrealean fungi revealed in the Sordariomycetes","volume":"54","author":"Darmostuk","year":"2025","journal-title":"Persoonia"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1016\/j.scitotenv.2019.04.247","article-title":"DNA barcode reference libraries for the monitoring of aquatic biota in Europe: Gap-analysis and recommendations for future work","volume":"678","author":"Weigand","year":"2019","journal-title":"Sci. Total Environ."},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Tzafesta, E., Saccomanno, B., Zangaro, F., Vadrucci, M.R., Specchia, V., and Pinna, M. (2022). DNA Barcode Gap Analysis for Multiple Marker Genes for Phytoplankton Species Biodiversity in Mediterranean Aquatic Ecosystems. Biology, 11.","DOI":"10.3390\/biology11091277"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"119374","DOI":"10.1016\/j.envres.2024.119374","article-title":"Barcoding Brazilian mammals to monitor biological diversity and threats: Trends, perspectives, and knowledge gaps","volume":"258","author":"Oliveira","year":"2024","journal-title":"Environ. Res."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Hollingsworth, P.M., Fantoni, K., Koureas, D., Arakelyan, M., Aravanopoulos, F.A., Bacela-Spychalska, K., Ballesteros-Mejia, L., Beja, P., Bonchev, G.N., and Bou Dagher Kharrat, M. (2026). Species identification, discovery, and biomonitoring: Strategic priorities for DNA barcoding in Europe, set in a global context. BioScience, biag052.","DOI":"10.1093\/biosci\/biag052"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"939","DOI":"10.1111\/nph.14855","article-title":"Widespread sampling biases in herbaria revealed from large-scale digitization","volume":"217","author":"Daru","year":"2018","journal-title":"New Phytol."}],"container-title":["Journal of Fungi"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2309-608X\/12\/8\/549\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,23]],"date-time":"2026-07-23T17:36:43Z","timestamp":1784828203000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2309-608X\/12\/8\/549"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,23]]},"references-count":58,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["jof12080549"],"URL":"https:\/\/doi.org\/10.3390\/jof12080549","relation":{},"ISSN":["2309-608X"],"issn-type":[{"value":"2309-608X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,23]]}}}