{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T23:44:40Z","timestamp":1792107880796,"version":"4.4.0"},"reference-count":111,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2026,2,25]],"date-time":"2026-02-25T00:00:00Z","timestamp":1771977600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Microorganisms"],"abstract":"<jats:p>Filamentous fungi are among the most significant biological agents responsible for the biodeterioration of organic cultural heritage materials preserved in archives, libraries, and museums. Cellulose-based substrates\u2014such as paper, papyri, and plant-derived textiles\u2014as well as protein-based materials, including parchment and leather, provide favourable conditions for fungal colonization due to their chemical composition and hygroscopic behaviour. Once activated, fungi contribute to deterioration through a combination of mechanical penetration and biochemical processes, including the secretion of hydrolytic enzymes, organic acids, and pigmented metabolites, which progressively compromise the structural integrity and visual appearance of heritage objects. This review aims to critically synthesize current knowledge on the mechanisms of fungal biodeterioration affecting organic heritage materials, with particular attention to material-specific vulnerabilities, indoor environmental drivers, and implications for preventive conservation. Recent advances in fungal ecology have highlighted the presence of xerophilic and extremotolerant taxa capable of persisting under conditions traditionally considered unfavourable for microbial growth, posing new challenges for conservation management. Rather than attributing biodeterioration directly to global climate change, this review explicitly emphasizes the role of indirect and building-mediated climate-related stressors\u2014such as increased frequency of moisture intrusion events, infrastructure vulnerability, and microclimatic instability within buildings\u2014in shaping fungal risk in indoor heritage environments. The integration of environmental monitoring, microbiological diagnostics, and predictive risk-assessment tools emerges as a key strategy for early detection and mitigation. By consolidating interdisciplinary evidence from microbiology, materials science, and heritage conservation, this work underscores the need to shift from reactive restoration toward anticipatory, risk-based preventive approaches to ensure the long-term preservation of organic cultural heritage materials.<\/jats:p>","DOI":"10.3390\/microorganisms14030526","type":"journal-article","created":{"date-parts":[[2026,2,25]],"date-time":"2026-02-25T08:50:27Z","timestamp":1772009427000},"page":"526","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":4,"title":["Filamentous Fungi and the Biodeterioration of Organic Cultural Heritage Materials: A Systematic Review of Mechanisms, Risks, and Preventive Conservation Strategies"],"prefix":"10.3390","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-6651-9794","authenticated-orcid":false,"given":"Giancarlo","family":"Angeles Flores","sequence":"first","affiliation":[{"name":"Department of Chemistry, Biology and Biotechnology, University of Perugia, Via del Giochetto, 06122 Perugia, Italy"},{"name":"Centro di Ricerca per l\u2019Innovazione, Digitalizzazione, Valorizzazione e Fruizione del Patrimonio Culturale e Ambientale (CE.D.I.PA.), Piazza San Gabriele dell\u2019Addolorata, 4, 06049 Spoleto, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7768-0468","authenticated-orcid":false,"given":"Roberto","family":"Venanzoni","sequence":"additional","affiliation":[{"name":"Department of Chemistry, Biology and Biotechnology, University of Perugia, Via del Giochetto, 06122 Perugia, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3942-237X","authenticated-orcid":false,"given":"Sabata","family":"Martino","sequence":"additional","affiliation":[{"name":"Department of Chemistry, Biology and Biotechnology, University of Perugia, Via del Giochetto, 06122 Perugia, Italy"},{"name":"Centro di Ricerca per l\u2019Innovazione, Digitalizzazione, Valorizzazione e Fruizione del Patrimonio Culturale e Ambientale (CE.D.I.PA.), Piazza San Gabriele dell\u2019Addolorata, 4, 06049 Spoleto, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6862-1079","authenticated-orcid":false,"given":"Paola","family":"Angelini","sequence":"additional","affiliation":[{"name":"Department of Chemistry, Biology and Biotechnology, University of Perugia, Via del Giochetto, 06122 Perugia, Italy"},{"name":"Centro di Ricerca per l\u2019Innovazione, Digitalizzazione, Valorizzazione e Fruizione del Patrimonio Culturale e Ambientale (CE.D.I.PA.), Piazza San Gabriele dell\u2019Addolorata, 4, 06049 Spoleto, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,2,25]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"7003","DOI":"10.1038\/s41598-024-57228-2","article-title":"Fungal and bacterial species richness in biodeteriorated seventeenth-century Venetian manuscripts","volume":"14","author":"Stratigaki","year":"2024","journal-title":"Sci. Rep."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"686","DOI":"10.1080\/1040841X.2019.1690420","article-title":"Fungi in archives, libraries, and museums: A review on paper conservation and human health","volume":"45","author":"Pinheiro","year":"2019","journal-title":"Crit. Rev. Microbiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1111\/arcm.12831","article-title":"Mold-deteriorated archaeological Egyptian papyri: Biodeteriogens, monitoring the deterioration, and treatment approach","volume":"65","author":"Saada","year":"2023","journal-title":"Archaeometry"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1038\/s40494-025-01601-5","article-title":"Identification, deterioration, and protection of organic cultural heritages from a modern perspective","volume":"13","author":"Wang","year":"2025","journal-title":"NPJ Herit. Sci."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Kujovi\u0107, A., Gostin\u010dar, C., Kavkler, K., Govedi\u0107, N., Gunde-Cimerman, N., and Zalar, P. (2024). Degradation potential of xerophilic and xerotolerant fungi contaminating historic canvas paintings. J. Fungi, 10.","DOI":"10.3390\/jof10010076"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.ibiod.2014.12.012","article-title":"The Extreme Environment of a Library: Xerophilic Fungi Inhabiting Indoor Niches","volume":"99","author":"Micheluz","year":"2015","journal-title":"Int. Biodeterior. Biodegrad."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1186\/s40494-024-01369-0","article-title":"An overview of gel-based cleaning approaches for art conservation","volume":"12","author":"Koochakzaei","year":"2024","journal-title":"npj Herit. Sci."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"113025","DOI":"10.1016\/j.buildenv.2025.113025","article-title":"Multifaceted assessment of fungal diversity and biodegradation activity in historical library","volume":"278","author":"Branysova","year":"2025","journal-title":"Build. Environ."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"e00200-22","DOI":"10.1128\/mmbr.00200-22","article-title":"Fungal biodeterioration and preservation of cultural heritage, artwork and historical artifacts: Extremophily and adaptation","volume":"88","author":"Gadd","year":"2024","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"e594","DOI":"10.1016\/S2666-5247(24)00039-9","article-title":"Impact of Climate Change and Natural Disasters on Fungal Infections","volume":"5","author":"Seidel","year":"2024","journal-title":"Lancet Microbe"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"n71","DOI":"10.1136\/bmj.n71","article-title":"The PRISMA 2020 statement: An updated guideline for reporting systematic reviews","volume":"372","author":"Page","year":"2021","journal-title":"BMJ"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1136\/emermed-2020-209567","article-title":"Population, Intervention, Comparison, Outcomes and Study (PICOS) design as a framework to formulate eligibility criteria in systematic reviews","volume":"37","author":"Janati","year":"2020","journal-title":"Emerg. Med. J."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Li, M., Gao, Q., and Yu, T. (2023). Kappa statistic considerations in evaluating inter-rater reliability between two raters: Which, when and context matters. BMC Cancer, 23.","DOI":"10.1186\/s12885-023-11325-z"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Pyzik, A., Ciuchci\u0144ski, K., Dziurzy\u0144ski, M., and Dziewit, L. (2021). The Bad and the Good\u2014Microorganisms in Cultural Heritage Environments\u2014An Update on Biodeterioration and Biotreatment Approaches. Materials, 14.","DOI":"10.3390\/ma14010177"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Petraretti, M., Duffy, K.J., Del Mondo, A., Pollio, A., and De Natale, A. (2021). Community composition and ex-situ cultivation of fungi associated with UNESCO heritage monuments in the Bay of Naples. Appl. Sci., 11.","DOI":"10.3390\/app11104327"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1038\/s40494-025-01622-0","article-title":"Application of spectroscopy technique in cultural heritage: Systematic review and bibliometric analysis","volume":"13","author":"Ye","year":"2025","journal-title":"npj Herit. Sci."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Camargo-Caicedo, Y., Borja P\u00e9rez, H., Mu\u00f1oz Fuentes, M., Vergara-V\u00e1squez, E., and V\u00e9lez-Pereira, A.M. (2024). Biodeterioration risk assessment in libraries by airborne fungal spores. J. Fungi, 10.","DOI":"10.3390\/jof10100680"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1623","DOI":"10.1038\/s41598-018-37651-y","article-title":"Three ancient documents solve the jigsaw of the parchment purple spot deterioration and validate the microbial succession model","volume":"9","author":"Migliore","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_19","unstructured":"UNESCO (2021). Heritage and Resilience: Safeguarding Cultural Heritage in the Face of Climate Change, United Nations Educational, Scientific and Cultural Organization. Available online: https:\/\/unesdoc.unesco.org\/ark:\/48223\/pf0000378684."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Barker, T.H., Hasanoff, S., Aromataris, E., Stone, J.C., Leonardi-Bee, J., Sears, K., Klugar, M., Tufanaru, C., Moola, S., and Liu, X.L. (2025). The Revised JBI Critical Appraisal Tool for the Assessment of Risk of Bias for Analytical Cross-Sectional Studies. JBI Evid. Synth., in press.","DOI":"10.11124\/JBIES-24-00523"},{"key":"ref_21","first-page":"2127","article-title":"Methodological Quality of Case Series Studies: An Introduction to the JBI Critical Appraisal Tool","volume":"18","author":"Munn","year":"2020","journal-title":"JBI Evid. Synth."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"j4008","DOI":"10.1136\/bmj.j4008","article-title":"AMSTAR 2: A Critical Appraisal Tool for Systematic Reviews That Include Randomised or Non-Randomised Studies of Healthcare Interventions, or Both","volume":"358","author":"Shea","year":"2017","journal-title":"BMJ"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"105784","DOI":"10.1016\/j.ibiod.2024.105784","article-title":"Unearthing the Global Patterns of Cultural Heritage Microbiome for Conservation","volume":"190","author":"Yu","year":"2024","journal-title":"Int. Biodeterior. Biodegrad."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"106001","DOI":"10.1016\/j.ibiod.2025.106001","article-title":"Bridging the Link Between Microbial Biofilm and Biodeterioration in Cultural Heritage Research","volume":"198","author":"Luo","year":"2025","journal-title":"Int. Biodeterior. Biodegrad."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"5159","DOI":"10.1007\/s00253-021-11343-6","article-title":"Hyphal Growth of Penicillium rubens in Changing Relative Humidity","volume":"105","author":"Ruijten","year":"2021","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Jia, S., Li, C., Wu, K., Qi, D., and Wang, S. (2022). Effect of Water Activity on Conidia Germination in Aspergillus flavus. Microorganisms, 10.","DOI":"10.3390\/microorganisms10091744"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1151","DOI":"10.1007\/s10482-022-01762-4","article-title":"Inter- and Intra-Species Heterogeneity in Germination of Aspergillus Conidia","volume":"115","author":"Ijadpanahsaravi","year":"2022","journal-title":"Antonie Van Leeuwenhoek"},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Hsin, K.-T., Lee, H., Huang, Y.-C., Lin, G.-J., Lin, P.-Y., Lin, Y.-C.J., and Chen, P.-Y. (2025). Lignocellulose Degradation in Bacteria and Fungi: Cellulosomes and Industrial Relevance. Front. Microbiol., 16.","DOI":"10.3389\/fmicb.2025.1583746"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"e01184-25","DOI":"10.1128\/mbio.01184-25","article-title":"Cell Wall Remodeling in a Fungal Pathogen Is Required for Hyphal Growth into Microspaces","volume":"16","author":"Miki","year":"2025","journal-title":"mBio"},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Loukou, E., Jensen, N.F., Rohde, L., and Andersen, B. (2024). Damp buildings: Associated fungi and how to find them. J. Fungi, 10.","DOI":"10.3390\/jof10020108"},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Isola, D., Bartoli, F., Meloni, P., Caneva, G., and Zucconi, L. (2022). Black Fungi and Stone Heritage Conservation: Ecological and Metabolic Assays for Evaluating Colonization Potential and Responses to Traditional Biocides. Appl. Sci., 12.","DOI":"10.3390\/app12042038"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"e24022","DOI":"10.1016\/j.heliyon.2024.e24022","article-title":"Enzymatic Degradation of Cellulose in Soil: A Review","volume":"10","author":"Datta","year":"2024","journal-title":"Heliyon"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"804","DOI":"10.1039\/D3CC05298B","article-title":"The Chemical Logic of Enzymatic Lignin Degradation","volume":"60","author":"Bugg","year":"2024","journal-title":"Chem. Commun."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1308","DOI":"10.3390\/heritage6020072","article-title":"Evaluation of the Irradiation Treatment Effects on Ancient Parchment Samples","volume":"6","author":"Vadrucci","year":"2023","journal-title":"Heritage"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"993","DOI":"10.1140\/epjp\/s13360-023-04635-5","article-title":"Deterioration Effects of X-ray Irradiation in Artificially Aged Parchment","volume":"138","author":"Mercuri","year":"2023","journal-title":"Eur. Phys. J. Plus"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1038\/s41529-022-00296-4","article-title":"Investigate the Role of Fungal Communities Associated with a Historical Manuscript from the 17th Century in Biodegradation","volume":"6","author":"Fouda","year":"2022","journal-title":"NPJ Mater. Degrad."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Gomoiu, I., Cojoc, R., Ruginescu, R., Neagu, S., Enache, M., Dumbravician, M., Olteanu, I., Radvan, R., and Ghervase, L. (2022). The Susceptibility to Biodegradation of Some Consolidants Used in the Restoration of Mural Paintings. Appl. Sci., 12.","DOI":"10.3390\/app12147229"},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Sala-Luis, A., Oliveira-Urquiri, H., Bosch-Roig, P., and Mart\u00edn-Rey, S. (2024). Eco-Sustainable Approaches to Prevent and\/or Eradicate Fungal Biodeterioration on Easel Painting. Coatings, 14.","DOI":"10.3390\/coatings14010124"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"15854","DOI":"10.1038\/s41598-023-42427-0","article-title":"Built Environment Microbiomes Transition from Outdoor to Human-Associated Communities after Construction and Commissioning","volume":"13","author":"Young","year":"2023","journal-title":"Sci. Rep."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Chawla, H., Anand, P., Garg, K., Bhagat, N., Varmani, S.G., Bansal, T., McBain, A.J., and Marwah, R.G. (2023). A Comprehensive Review of Microbial Contamination in the Indoor Environment: Sources, Sampling, Health Risks, and Mitigation Strategies. Front. Public Health, 11.","DOI":"10.3389\/fpubh.2023.1285393"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"3056","DOI":"10.1038\/s41467-024-46920-6","article-title":"Survival and Rapid Resuscitation Permit Limited Productivity in Desert Microbial Communities","volume":"15","author":"Imminger","year":"2024","journal-title":"Nat. Commun."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"302","DOI":"10.1016\/j.culher.2024.12.018","article-title":"Microbial Biodeterioration of Historic Wood Based on Classical and Omics Methods with Model Studies","volume":"71","author":"Gutarowska","year":"2025","journal-title":"J. Cult. Herit."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Derksen, K., Brimblecombe, P., Pi\u00f1ar, G., Waldherr, M., Graf, A., Haltrich, M., Querner, P., and Sterflinger, K. (2024). Fungal Biodeterioration Risk in Monastic Libraries without Climate Control. Microorganisms, 12.","DOI":"10.3390\/microorganisms12071450"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"14","DOI":"10.21014\/actaimeko.v12i4.1508","article-title":"Risk Assessment of Fungal Formations in Historic Buildings Based on Dynamic Thermo-Hygrometric Analysis","volume":"12","author":"Carpino","year":"2023","journal-title":"Acta IMEKO"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"2986","DOI":"10.1038\/s41396-021-00988-w","article-title":"Metabolic Flexibility Allows Bacterial Habitat Generalists to Become Dominant in a Frequently Disturbed Ecosystem","volume":"15","author":"Chen","year":"2021","journal-title":"ISME J."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1016\/j.culher.2023.08.012","article-title":"Exploration of Effects by the \u201cAlum-Glue Solution\u201d on Xuan Paper Degradation Using a 23 Factorial Design Experiment","volume":"64","author":"Zhang","year":"2023","journal-title":"J. Cult. Herit."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"172","DOI":"10.1186\/s40494-024-01261-x","article-title":"Deterioration-Associated Microbiome of a Modern Photographic Artwork: The Case of Skull and Crossbones by Robert Mapplethorpe","volume":"12","author":"Petraretti","year":"2024","journal-title":"Herit. Sci."},{"key":"ref_48","doi-asserted-by":"crossref","unstructured":"Yan, Y., Tang, Y., and Yang, Y. (2025). Chemical Conservation of Paper-Based Cultural Heritage. Molecules, 30.","DOI":"10.3390\/books978-3-7258-3625-3"},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Corbu, V.M., Gheorghe-Barbu, I., Dumbrava, A.S., Vr\u00e2ncianu, C.O., and Sesan, T.E. (2023). Current Insights in Fungal Importance\u2014A Comprehensive Review. Microorganisms, 11.","DOI":"10.3390\/microorganisms11061384"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1186\/s40494-024-01511-y","article-title":"Evaluation of Some Fungicides for Inhibiting Proteolytic Fungi Isolated from Leather Binding of a Historical Manuscript Dated Back to the Mamluk Period","volume":"12","author":"Abdelhafez","year":"2024","journal-title":"Herit. Sci."},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"De Leo, F., Marchetta, A., and Urz\u00ec, C. (2022). Black Fungi on Stone-Built Heritage: Current Knowledge and Future Outlook. Appl. Sci., 12.","DOI":"10.3390\/app12083969"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1186\/s40494-024-01298-y","article-title":"Rainbow Code of Biodeterioration to Cultural Heritage Objects","volume":"12","author":"Avdanina","year":"2024","journal-title":"Herit. Sci."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"3853","DOI":"10.1007\/s13399-022-02815-4","article-title":"Isolated Essential Oils as Antifungal Compounds for Organic Materials","volume":"14","author":"Taha","year":"2024","journal-title":"Biomass Convers. Biorefin."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1007\/s00253-025-13553-8","article-title":"Biodeterioration of Canvas Paintings: Microbial Role and Development of Sustainable Treatments for Biocontrol","volume":"109","author":"Climaco","year":"2025","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"186","DOI":"10.1016\/j.ibiod.2008.01.005","article-title":"Microfungal Biodeterioration of Historic Paper: Preliminary FTIR and Microbiological Anal-yses","volume":"62","author":"Zotti","year":"2008","journal-title":"Int. Biodeterior. Biodegrad."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Yovchevska, L., Gocheva, Y., Stoyancheva, G., Miteva-Staleva, J., Dishliyska, V., Abrashev, R., Stamenova, T., Angelova, M., and Krumova, E. (2025). Halophilic Fungi\u2014Features and Potential Applications. Microorganisms, 13.","DOI":"10.3390\/microorganisms13010175"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"173880","DOI":"10.1016\/j.scitotenv.2024.173880","article-title":"A Novel Contaminant in Museums? A Cross-Sectional Study on Xerophilic Aspergillus Growth in Climate-Controlled Repositories","volume":"944","author":"Bastholm","year":"2024","journal-title":"Sci. Total Environ."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/j.culher.2024.02.016","article-title":"Analysis of Cracking Behavior of Murals in Mogao Grottoes under Environmental Humidity Change","volume":"67","author":"Zhang","year":"2024","journal-title":"J. Cult. Herit."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"354","DOI":"10.1111\/ina.12313","article-title":"Fungal and Bacterial Growth in Floor Dust at Elevated Relative Humidity Levels","volume":"27","author":"Dannemiller","year":"2017","journal-title":"Indoor Air"},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Verticchio, E., Frasca, F., Mat\u00e8, D., Giammusso, F.M., Sani, M., Sebastiani, M.L., Sclocchi, M.C., and Siani, A.M. (2023). Assessing the Impact of Climate Change on the Biodeterioration Risk in Historical Buildings of the Mediterranean Area: The State Archives of Palermo. Atmosphere, 14.","DOI":"10.3390\/atmos14071169"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1007\/s10453-021-09730-7","article-title":"Spatiotemporal Distribution and the Passive Dispersal of Fungal Spores through HVAC Systems","volume":"38","author":"Lin","year":"2022","journal-title":"Aerobiologia"},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"103","DOI":"10.15698\/mic2022.05.776","article-title":"Air- and Dustborne Fungi in Repositories of the National Archive of the Republic of Cuba","volume":"9","author":"Borrego","year":"2022","journal-title":"Microb. Cell"},{"key":"ref_63","doi-asserted-by":"crossref","unstructured":"Riedling, O.L., David, K.T., and Rokas, A. (2024). Global Patterns of Species Diversity and Distribution in the Biomedically and Biotechnologically Important Fungal Genus Aspergillus. bioRxiv.","DOI":"10.1101\/2024.11.29.626055"},{"key":"ref_64","unstructured":"Hocking, A.D., and Pitt, J.I. (2020). Water Activity Limits for Fungal Growth and Spoilage. Fungi and Food Spoilage, Springer. [4th ed.]."},{"key":"ref_65","first-page":"111951","article-title":"Microclimate Modelling for Preventive Conservation: Multi-Sensor Strategies and Risk Forecasting","volume":"261","author":"Bertolin","year":"2022","journal-title":"Energy Build."},{"key":"ref_66","first-page":"211","article-title":"Sorption Isotherms of Heritage Materials and Their Use in Climate Response Modelling","volume":"69","author":"Carter","year":"2024","journal-title":"Stud. Conserv."},{"key":"ref_67","unstructured":"ASHRAE (2022). Air Filtration Guidelines for Museums, Galleries, Archives and Libraries. ASHRAE Handbook: HVAC Applications, ASHRAE. Chapter 24."},{"key":"ref_68","first-page":"110236","article-title":"Fungal Aerosolisation from HVAC Systems: A Review","volume":"235","author":"Reponen","year":"2023","journal-title":"Build. Environ."},{"key":"ref_69","first-page":"89","article-title":"Organic Dust Composition in Heritage Collections and Its Impact on Fungal Colonisation","volume":"64","author":"Gallo","year":"2023","journal-title":"J. Cult. Herit."},{"key":"ref_70","first-page":"345","article-title":"Viability and Persistence of Airborne Fungal Spores in Indoor Environments","volume":"31","author":"Wessel","year":"2022","journal-title":"Indoor Built Environ."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"119762","DOI":"10.1016\/j.jenvman.2023.119762","article-title":"Microclimate, airborne particles, and microbiological monitoring protocol for conservation of rock-art caves: The case of the world-heritage site La Garma cave (Spain)","volume":"351","author":"Cuezva","year":"2024","journal-title":"J. Environ. Manag."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1081\/ASR-200054370","article-title":"Spectroscopic Techniques in Cultural Heritage Conservation: A Survey","volume":"40","author":"Bitossi","year":"2005","journal-title":"Appl. Spectrosc. Rev."},{"key":"ref_73","unstructured":"Heracles Project Consortium (2022). HERACLES: Heritage Resilience Against Climate Events\u2014Integrated Methodologies for Risk Assessment. Environ. Sci. Policy, 137, 56\u201371."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Tringa, E., Georgoulias, A.K., Akritidis, D., Feidas, H., and Zanis, P. (2025). Assessing the Future Risk of Damage to European Cultural Heritage Due to Climate Change. Heritage, 8.","DOI":"10.3390\/heritage8050175"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"101136","DOI":"10.1016\/j.apr.2021.101136","article-title":"Indoor and Outdoor Microbiological Air Quality in Naturally and Mechanically Venti-lated University Libraries","volume":"12","author":"Hassan","year":"2021","journal-title":"Atmos. Pollut. Res."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"e00046","DOI":"10.1002\/ggn2.202500046","article-title":"Bio-Archive of Cultural Heritage Microbiomes for Sustainable Conservation in the Multi-Omics Era","volume":"6","author":"Fu","year":"2025","journal-title":"Adv. Genet."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"1601","DOI":"10.1007\/s42770-024-01323-z","article-title":"Metabarcode Insights into the Airborne Fungal Diversity in the Indoor and Outdoor Environments in Archives from Yunnan, Southwestern China","volume":"55","author":"Zhang","year":"2024","journal-title":"Braz. J. Microbiol."},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Dimabayao, J.J., Lara, J.L., Canoura, L.G., and Solheim, S. (2025). Integrating Climate Risk in Cultural Heritage: A Critical Review of Assessment Frameworks. Heritage, 8.","DOI":"10.3390\/heritage8080312"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"176","DOI":"10.1080\/00393630.2024.2375160","article-title":"Towards a National Heritage Climate Change Risk Assessment","volume":"69","author":"Lankester","year":"2024","journal-title":"Stud. Conserv."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1017\/aap.2024.18","article-title":"Exploring Climate Change Adaptations for Cultural Heritage: The ADAPT Framework","volume":"12","author":"Wright","year":"2024","journal-title":"Adv. Archaeol. Pract."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"506","DOI":"10.1128\/MMBR.66.3.506-577.2002","article-title":"Microbial Cellulose Utilization: Fundamentals and Biotechnology","volume":"66","author":"Lynd","year":"2002","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1186\/1754-6834-5-45","article-title":"Novel Enzymes for the Degradation of Cellulose","volume":"5","author":"Horn","year":"2012","journal-title":"Biotechnol. Biofuels"},{"key":"ref_83","first-page":"152","article-title":"Microscopic and Mechanical Alterations in Fungally Degraded Paper","volume":"41","author":"Kim","year":"2020","journal-title":"J. Cult. Herit."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"9637","DOI":"10.1007\/s00253-013-5283-1","article-title":"Microbial Deterioration of Cultural Heritage and Works of Art\u2014Tilting at Windmills?","volume":"97","author":"Sterflinger","year":"2013","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1038\/s40494-025-01785-w","article-title":"Protection of Paper Manuscripts by Antimicrobial Polyphenolic Compounds from Fungal Extracts and Antioxidant Nanoparticles","volume":"13","author":"Mahmoud","year":"2025","journal-title":"npj Herit. Sci."},{"key":"ref_86","unstructured":"Florian, M.-L.E. (2002). Fungal Facts: Solving Fungal Problems in Heritage Collections, Archetype Publications."},{"key":"ref_87","first-page":"88","article-title":"Organic Acids Produced by Fungal Contaminants and Their Effects on Cellulosic Materials","volume":"11","author":"Gallo","year":"2023","journal-title":"Herit. Sci."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1016\/j.culher.2024.03.002","article-title":"Isolation and Identification of Fungal Biodeteriogens from the Wall of a Cultural Heritage Church and Potential Applicability of Antifungal Proteins in Protection","volume":"67","author":"Farkas","year":"2024","journal-title":"J. Cult. Herit."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"1366","DOI":"10.3390\/heritage4030074","article-title":"Paper Foxing Stains on a Historic Manuscript from the Early Qajar Era: Abiotic or Biotic Foxing?","volume":"4","author":"Koochakzaei","year":"2021","journal-title":"Heritage"},{"key":"ref_90","doi-asserted-by":"crossref","unstructured":"Pavlovic, J., Farkas, Z., Krakov\u00e1, L., and Pangallo, D. (2022). Color Stains on Paper: Fungal Pigments, Synthetic Dyes and Their Hypothetical Removal by Enzymatic Approaches. Appl. Sci., 12.","DOI":"10.3390\/app12199991"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1016\/j.funbio.2021.11.001","article-title":"Biodeterioration and Cellulolytic Activity by Fungi Isolated from a Nineteenth-Century Painting at the National Theatre of Costa Rica","volume":"126","author":"Vieto","year":"2022","journal-title":"Fungal Biol."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1038\/s40494-025-02173-0","article-title":"Eco-Friendly Control of Biodeteriogenic Black Fungi on Cultural Heritage Stones Using Lamiaceae Essential Oils","volume":"13","author":"SERT","year":"2025","journal-title":"npj Herit. Sci."},{"key":"ref_93","first-page":"198","article-title":"Integrated Microchemical and Spectroscopic Approaches for Early Detection of Fungal Activity on Paper","volume":"57","author":"Bicchieri","year":"2022","journal-title":"J. Cult. Herit."},{"key":"ref_94","first-page":"105516","article-title":"Relative Humidity Thresholds for Fungal Activation on Paper Artefacts","volume":"187","author":"Sadek","year":"2024","journal-title":"Int. Biodeterior. Biodegrad."},{"key":"ref_95","doi-asserted-by":"crossref","unstructured":"Kuka, E., Cirule, D., Andersone, I., Andersons, B., and Fridrihsone, V. (2022). Conditions Influencing Mould Growth for Effective Prevention of Wood Deterioration Indoors. Appl. Sci., 12.","DOI":"10.3390\/app12030975"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"112548","DOI":"10.1016\/j.buildenv.2025.112548","article-title":"Revisiting the Joint Effect of Temper-ature and Relative Humidity on Airborne Mold and Bacteria Concentration in Indoor Environment: A Machine Learning Ap-proach","volume":"270","author":"Kim","year":"2025","journal-title":"Build. Environ."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1016\/j.culher.2022.02.009","article-title":"The Mysterious Mould Outbreak\u2014A Comprehensive Fungal Colonisation in a Climate-Controlled Museum Repository Challenges the Environmental Guidelines for Heritage Collections","volume":"55","author":"Bastholm","year":"2022","journal-title":"J. Cult. Herit."},{"key":"ref_98","first-page":"201","article-title":"Microclimatic Heterogeneity in Historic Buildings","volume":"67","author":"Mazzei","year":"2022","journal-title":"Stud. Conserv."},{"key":"ref_99","unstructured":"European Commission (2021). Forging a Climate-Resilient Europe: The New EU Strategy on Adaptation to Climate Change, European Union."},{"key":"ref_100","unstructured":"United Nations (2015). Transforming Our World: The 2030 Agenda for Sustainable Development, United Nations."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"100727","DOI":"10.1016\/j.crm.2025.100727","article-title":"Multi-Determinant Climate Change Risk Assessment for Heritage: A Review of Current Approaches and Future Needs","volume":"49","author":"Thomas","year":"2025","journal-title":"Clim. Risk Manag."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"e21928","DOI":"10.1016\/j.heliyon.2023.e21928","article-title":"Study of Kinetic Model for Fungal Spore Germination under Dynamic Conditions: Case Study on Germination of Penicillium Expansum Spores","volume":"9","author":"Boonruang","year":"2023","journal-title":"Heliyon"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"105730","DOI":"10.1016\/j.ibiod.2024.105730","article-title":"Xerophilic and Cellulolytic Fungi in the Indoor Air of Houses in Havana","volume":"188","author":"Almaguer","year":"2024","journal-title":"Int. Biodeterior. Biodegrad."},{"key":"ref_104","doi-asserted-by":"crossref","unstructured":"Cirone, M., Figoli, A., Galiano, F., La Russa, M.F., Macchia, A., Mancuso, R., Ricca, M., Rovella, N., Taverniti, M., and Ruffolo, S.A. (2023). Innovative Methodologies for the Conservation of Cultural Heritage against Biodeterioration: A Review. Coatings, 13.","DOI":"10.3390\/coatings13121986"},{"key":"ref_105","doi-asserted-by":"crossref","unstructured":"Dalla Mora, T., De Vivo, M.A., Scarpa, M., and Peron, F. (2025). Critical Review of the Application of the Principal International Standards and Guidelines on Indoor Microclimates for the Preventive Conservation of Cultural Heritage. Sustainability, 17.","DOI":"10.3390\/su17031189"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"113173","DOI":"10.1016\/j.jobe.2025.113173","article-title":"Preventive Conservation Guidelines versus Damage Risk Models: The Hygrothermal Environment for Organic Collections in Heritage Museums in a Mediterranean Climate","volume":"111","author":"Saraiva","year":"2025","journal-title":"J. Build. Eng."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"479","DOI":"10.26444\/aaem\/193292","article-title":"Distribution Characteristics and Analysis of Fungal Aerosol Concentration and Particle Size in Air-Conditioned Wards in Wuhan, China","volume":"31","author":"Song","year":"2024","journal-title":"Ann. Agric. Environ. Med."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"lxae258","DOI":"10.1093\/jambio\/lxae258","article-title":"Long-Term Monitoring of the Hypogeal Etruscan Tomba degli Scudi, Tarquinia, Italy: Early Detection of Black Spots, Investigation of Fungal Community, and Evaluation of Their Biodeterioration Potential","volume":"135","author":"Kratter","year":"2024","journal-title":"J. Appl. Microbiol."},{"key":"ref_109","doi-asserted-by":"crossref","unstructured":"De Leo, F., and Isola, D. (2022). The Role of Fungi in Biodeterioration of Cultural Heritage: New Insights for Their Control. Appl. Sci., 12.","DOI":"10.3390\/app122010490"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"7380","DOI":"10.1038\/s41598-025-92134-1","article-title":"Characterisation and Restoration of a Severely Damaged Paper Archive from the Qing Dynasty","volume":"15","author":"Liu","year":"2025","journal-title":"Sci. Rep."},{"key":"ref_111","first-page":"89","article-title":"Natural Substances as Biocides in the Fungi Treatment on Artistic Products to Protect the Environment and Health of Restoration Workers","volume":"52","author":"Casorri","year":"2023","journal-title":"Ital. J. Mycol."}],"container-title":["Microorganisms"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2607\/14\/3\/526\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,25]],"date-time":"2026-02-25T09:22:41Z","timestamp":1772011361000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2607\/14\/3\/526"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,25]]},"references-count":111,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2026,3]]}},"alternative-id":["microorganisms14030526"],"URL":"https:\/\/doi.org\/10.3390\/microorganisms14030526","relation":{},"ISSN":["2076-2607"],"issn-type":[{"value":"2076-2607","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,25]]}}}