Black mold on a white limestone: the role of Stachybotrys chartarum in stone heritage deterioration
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Abstract
Abstract Fungal deterioration of historical stone objects can significantly impact their mineralogy and chemistry, leading to irreplaceable cultural heritage losses. Ançã stone, a unique type of limestone widely used in some of the most emblematic Portuguese monuments, due to its soft, porous calcium carbonate nature, is known to have a high intrinsic bioreceptivity, being highly susceptible to fungal colonization and subsequent biodeterioration. In a prior experimental study aimed at characterizing the fungal diversity thriving on a Ançã stone funerary art piece showing visible signs of alteration at the Lemos Pantheon, a national monument in Águeda, Portugal, several isolates of an unexpected black mold were obtained. The detection of Stachybotrys chartarum across the multiple approaches used during that investigation suggested that its occurrence was not solely due to environmental contamination, prompting us to conduct a comprehensive analysis, as this species had never been previously associated with this lithotype. Our results proved the presence of S. chartarum, representing the first confirmed occurrence of this species from Ançã stone. Using a multi-locus phylogeny combined with morphological, physiological, and ecological analyses, we characterized these isolates. We also demonstrated the biodeteriorative potential of S. chartarum through plate assays, along with its in vitro ability to actively colonize limestone, with and without organic matter deposition, and solely under the natural fluctuations in relative humidity, causing changes to the substrate. We were able to observe hyphae penetration into calcite layers of inoculated stone probes, potential mineral desegregation and displacement, as well as putative mineralization phenomena around its hyphae after 10 months. This study provides critical new insights into the biodeteriorative potential of S. chartarum, which, along with its well-known toxigenic attributes, adds to its infamous reputation.
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OpenAlex topics: Building materials and conservation · Cultural Heritage Materials Analysis · Lichen and fungal ecology
SciVX discovery tags are terminology-based. OpenAlex topics are source-provided classifications.
References (100+)
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1. Wang, Y. et al. Overview of Stachybotrys (Memnoniella) and current species status. Fungal Divers. 71, 17–83 (2015).10.1007/s13225-014-0319-0 ↗
2. Corda, A. C. I. Icones Fungorum Hucusque Cognitorum. https://archive.org/details/per_cabimicrofilm_aci-corda_1837_1 (1837).
3. Etzel, R. A. et al. Acute pulmonary haemorrhage in infants associated with exposure to Stachybotrys atra and other fungi. Arch. Pediatr. Adolesc. Med. 152, 757–762 (1998).10.1001/archpedi.152.8.757 ↗
4. Vesper, S. J., Magnuson, M. L., Dearborn, D. G., Yike, I. & Haugland, R. A. Initial characterization of the hemolysin stachylysin from Stachybotrys chartarum. Infect. Immun. 69, 912–916 (2001).10.1128/IAI.69.2.912-916.2001 ↗
5. Vesper, S. J. & Vesper, M. J. Stachylysin may be a cause of hemorrhaging in humans exposed to Stachybotrys chartarum. Infect. Immun. 70, 2065–2069 (2002).10.1128/IAI.70.4.2065-2069.2002 ↗
6. Piecková, E. et al. Inflammatory and haematotoxic potential of indoor Stachybotrys chartarum (Ehrenb.) Hughes metabolites. Arh. Hig. Rada Toksikol. 60, 401–409 (2009).10.2478/10004-1254-60-2009-1971 ↗
7. Al-Ahmad, M. et al. Symptoms after mould exposure including Stachybotrys chartarum, and comparison with darkroom disease. Allergy 65, 245–255 (2010).10.1111/j.1398-9995.2009.02157.x ↗
8. Yike, I. & Dearborn, D. Guest editorial—novel insights into the pathology of Stachybotrys chartarum. Mycopathologia 172, 1–3 (2011).10.1007/s11046-011-9426-6 ↗
9. Bhan, U. et al. Stachybotrys chartarum-induced hypersensitivity pneumonitis is TLR9 dependent. Am. J. Pathol. 179, 2779–2787 (2011).10.1016/j.ajpath.2011.08.019 ↗
10. Hyde, K. D. et al. The world’s ten most feared fungi. Fungal Divers. 93, 161–194 (2018).10.1007/s13225-018-0413-9 ↗
11. Zyska, B. Fungi isolated from library materials: a review of the literature. Int. Biodeterior. Biodegrad. 40, 43–51 (1997).10.1016/S0964-8305(97)00061-9 ↗
12. Hodgson, M. J. et al. Building-associated pulmonary disease from exposure to Stachybotrys chartarum and Aspergillus versicolor. J. Occup. Environ. Med. 40, 241–249 (1998).10.1097/00043764-199803000-00006 ↗
13. Korpia, A., Pasanen, A.-L. & Viitanenb, H. Volatile metabolites of Serpula lacrymans, Coniophora puteana, Poria placenta, Stachybotrys chartarum and Chaetomium globosum. Build Environ. 34, 205–211 (1998).10.1016/S0360-1323(98)00011-0 ↗
14. Nielsen, K. F., Hansen, M. Ø., Larsen, T. O. & Thrane, U. Production of trichothecene mycotoxins on water damaged gypsum boards in Danish buildings. Int. Biodeterior. Biodegrad. 42, 1–7 (1998).10.1016/S0964-8305(98)00035-3 ↗
15. Kuhn, D. M. & Ghannoum, M. A. Indoor mould, toxigenic fungi, and Stachybotrys chartarum: infectious disease perspective. Clin. Microbiol. Rev. 16, 144–172 (2003).10.1128/CMR.16.1.144-172.2003 ↗
16. Szostak-Kotowa, J. Biodeterioration of textiles. Int. Biodeterior. Biodegrad. 53, 165–170 (2004).10.1016/S0964-8305(03)00090-8 ↗
17. Pournou, A. & Bogomolova, E. Fungal colonization on excavated prehistoric wood: implications for in-situ display. Int. Biodeterior. Biodegrad. 63, 371–378 (2009).10.1016/j.ibiod.2008.11.001 ↗
18. Andersen, B., Dosen, I., Lewinska, A. M. & Nielsen, K. F. Pre-contamination of new gypsum wallboard with potentially harmful fungal species. Indoor Air 27, 6–12 (2017).10.1111/ina.12298 ↗
19. Dyląg, M., Spychała, K., Zielinski, J., Łagowski, D. & Gnat, S. Update on Stachybotrys chartarum—black mould perceived as toxigenic and potentially pathogenic to humans. Biology 11, 352 (2022).10.3390/biology11030352 ↗
20. Loukou, E., Jensen, N. F., Rohde, L. & Andersen, B. Damp buildings: associated fungi and how to find them. J. Fungi 10, 108 (2024).10.3390/jof10020108 ↗
21. Andersen, B. & Nissen, A. T. Evaluation of media for detection of Stachybotrys and Chaetomium species associated with water-damaged buildings. Int. Biodeterior. Biodegrad. 46, 111–116 (2000).10.1016/S0964-8305(00)00092-5 ↗
22. Frazer, S., Magan, N. & Aldred, D. The influence of water activity and temperature on germination, growth and sporulation of Stachybotrys chartarum strains. Mycopathologia 172, 17–23 (2011).10.1007/s11046-011-9394-x ↗
23. Ayerst, G. The effects of moisture and temperature on growth and spore germination in some fungi. J. Stored Prod. Res. 5, 127–141 (1969).10.1016/0022-474X(69)90055-1 ↗
24. Grant, C., Hunter, C. A., Flannigan, B. & Bravery, A. F. The moisture requirements of moulds isolated from domestic dwellings. Int. Biodeterior. 25, 259–284 (1989).10.1016/0265-3036(89)90002-X ↗
25. Johansson, P., Svensson, T. & Ekstrand-Tobin, A. Validation of critical moisture conditions for mould growth on building materials. Build Environ. 62, 201–209 (2013).10.1016/j.buildenv.2013.01.012 ↗
26. Hossain, M. A., Ahmed, M. S. & Ghannoum, M. A. Attributes of Stachybotrys chartarum and its association with human disease. J. Allergy Clin. Immunol. 113, 200–208 (2004).10.1016/j.jaci.2003.12.018 ↗
27. Cooley, J. D., Wong, W. C., Jumper, C. A. & Straus, D. C. Correlation between the prevalence of certain fungi and sick building syndrome. Occup. Environ. Med. 55, 579–584 (1998).10.1136/oem.55.9.579 ↗
28. Gravesen, S., Nielsen, P. A., Iversen, R. & Nielsen, K. F. Microfungal contamination of damp buildings—examples of risk constructions and risk materials. Environ. Health Perspect. 107, 505–508 (1999).10.1289/ehp.99107s3505 ↗
29. Tuomi, T. et al. Mycotoxins in crude building materials from water-damaged buildings. Appl. Environ. Microbiol. 66, 1899–1904 (2000).10.1128/AEM.66.5.1899-1904.2000 ↗
30. Øya, E. et al. Characterization and pro-inflammatory responses of spore and hyphae samples from various mould species. Indoor Air 28, 28–39 (2017).10.1111/ina.12426 ↗
31. Wilkins, K., Nielsen, K. F. & Din, S. U. Patterns of volatile metabolites and non-volatile trichothecenes produced by isolates of Stachybotrys, Fusarium, Trichoderma, Trichothecium and Memnoniella. Environ. Sci. Pollut. Res. Int. 10, 162–166 (2003).10.1065/espr2002.05.118 ↗
32. Betancourt, D. A., Krebs, K., Moore, S. A. & Martin, S. M. Microbial volatile organic compound emissions from Stachybotrys chartarum growing on gypsum wallboard and ceiling tile. BMC Microbiol. 13, 283 (2013).10.1186/1471-2180-13-283 ↗
33. Paiva, D. S. et al. Uncovering the fungal diversity colonizing limestone walls of a forgotten monument in the central region of Portugal by high-throughput sequencing and culture-based methods. Appl. Sci. 12, 10650 (2022).10.3390/app122010650 ↗
34. Paiva, D. S. et al. Exploring differences in culturable fungal diversity using standard freezing incubation—a case study in the limestones of Lemos Pantheon (Portugal). J. Fungi 9, 501 (2023).10.3390/jof9040501 ↗
35. Abdelhafez, A. A. M., El-Wekeel, F. M., Ramadan, E. M. & Abed-Allah, A. A. Microbial deterioration of archaeological marble: identification and treatment. Ann. Agric. Sci. 57, 137–144 (2012).10.1016/j.aoas.2012.08.007 ↗
36. Li, Y., Huang, Z., Petropoulos, E., Ma, Y. & Shen, Y. Humidity governs the wall-inhabiting fungal community composition in a 1600-year tomb of Emperor Yang. Sci. Rep. 10, 8421 (2020).10.1038/s41598-020-65478-z ↗
37. Mascaro, M. E., Pellegrino, G. & Palermo, A. M. Analysis of biodeteriogens on architectural heritage. An approach of applied botany on a Gothic building in southern Italy. Sustainability 14, 34 (2021).10.3390/su14010034 ↗
38. Scheerer, S., Ortega-Morales, O. & Gaylarde, C. Microbial deterioration of stone monuments—an updated overview. Adv. Appl. Microbiol. 66, 97–139 (2009).10.1016/S0065-2164(08)00805-8 ↗
39. Sterflinger, K. Fungi: their role in deterioration of cultural heritage. Fungal Biol. Rev. 24, 47–55 (2010).10.1016/j.fbr.2010.03.003 ↗
40. Dakal, T. C. & Cameotra, S. S. Microbially induced deterioration of architectural heritages: routes and mechanisms involved. Environ. Sci. Eur. 24, 36 (2012).10.1186/2190-4715-24-36 ↗
41. Gadd, G. M. Geomycology: biogeochemical transformations of rocks, minerals, metals and radionuclides by fungi, bioweathering and bioremediation. Mycol. Res. 111, 3–49 (2007).10.1016/j.mycres.2006.12.001 ↗
42. Gadd, G. M. Geomicrobiology of the built environment. Nat. Microbiol. 2, 16275 (2017).10.1038/nmicrobiol.2016.275 ↗
43. Gadd, G. M. Fungi, rocks, and minerals. Elements 13, 171–176 (2017).10.2113/gselements.13.3.171 ↗
44. Agrawal, S., Khumlianlal, J. & Devi, S. I. Uncovering the fungal diversity and biodeterioration phenomenon on archaeological carvings of the Badami cave temples: a microcosm study. Life 14, 28 (2024).10.3390/life14010028 ↗
45. Gadd, G. M., Fomina, M. & Pinzari, F. Fungal biodeterioration and preservation of cultural heritage, artwork, and historical artifacts: extremophily and adaptation. Microbiol. Mol. Biol. Rev. 88, e00200-22 (2024).10.1128/mmbr.00200-22 ↗
46. Miller, A. Z. et al. Primary bioreceptivity of limestones used in southern European monuments. Geol. Soc. Spec. Publ. 331, 79–92 (2010).10.1144/SP331.6 ↗
47. Miller, A., Dionísio, A. & Macedo, M. F. Primary bioreceptivity: a comparative study of different Portuguese lithotypes. Int. Biodeterior. Biodegrad. 57, 136–142 (2006).10.1016/j.ibiod.2006.01.003 ↗
48. Pinheiro, A. C. et al. Limestone biodeterioration: a review on the Portuguese cultural heritage scenario. J. Cult. Herit. 36, 275–285 (2019).10.1016/j.culher.2018.07.008 ↗
49. Trovão, J. et al. A contribution to understand the Portuguese emblematic Ançã limestone bioreceptivity to fungal colonization and biodeterioration. J. Cult. Herit. 49, 305–312 (2021).10.1016/j.culher.2021.04.003 ↗
50. Paiva, D. S. et al. Expanding the microcolonial black fungi Aeminiaceae family: Saxispiralis lemnorum gen. et sp. nov. (Mycosphaerellales), isolated from deteriorated limestone in the Lemos Pantheon, Portugal. J. Fungi 9, 916 (2023).10.3390/jof9090916 ↗
51. White, T. J., Bruns, T., Lee, S. & Taylor, J. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics in PCR Protocols, 315–322 (Elsevier, 1990).10.1016/B978-0-12-372180-8.50042-1 ↗
52. Vilgalys, R. & Hester, M. Rapid genetic identification and mapping of enzymatically amplified ribosomal DNA from several Cryptococcus species. J. Bacteriol. 172, 4238–4246 (1990).10.1128/jb.172.8.4238-4246.1990 ↗
53. Rehner, S. A. & Samuels, G. J. Molecular systematics of the Hypocreales: a teleomorph gene phylogeny and the status of their anamorphs. Can. J. Bot. 73, 816–823 (1995).10.1139/b95-327 ↗
54. Liu, Y. J., Whelen, S. & Hall, B. D. Phylogenetic relationships among ascomycetes: evidence from an RNA polymerse II subunit. Mol. Biol. Evol. 16, 1799–1808 (1999).10.1093/oxfordjournals.molbev.a026092 ↗
55. Sung, G.-H., Sung, J.-M., Hywel-Jones, N. L. & Spatafora, J. W. A multi-gene phylogeny of Clavicipitaceae (Ascomycota, Fungi): identification of localized incongruence using a combinational bootstrap approach. Mol. Phylogenet. Evol. 44, 1204–1223 (2007).10.1016/j.ympev.2007.03.011 ↗
56. Hong, S.-B., Cho, H.-S., Shin, H.-D., Frisvad, J. C. & Samson, R. A. Novel Neosartorya species isolated from soil in Korea. Int. J. Syst. Evol. Microbiol. 56, 477–486 (2006).10.1099/ijs.0.63980-0 ↗
57. Crous, P. W. et al. Phylogenetic lineages in Pseudocercospora. Stud. Mycol. 75, 37–114 (2013).10.3114/sim0005 ↗
58. Hunter, G. C., Wingfield, B. D., Crous, P. W. & Wingfield, M. J. A multi-gene phylogeny for species of Mycosphaerella occurring on Eucalyptus leaves. Stud. Mycol. 55, 147–161 (2006).10.3114/sim.55.1.147 ↗
59. Houbraken, J. & Samson, R. A. Phylogeny of Penicillium and the segregation of Trichocomaceae into three families. Stud. Mycol. 70, 1–51 (2011).10.3114/sim.2011.70.01 ↗
60. Altschul, S. F. et al. Gapped BLAST and PSI-BLAST: a new generation of protein database search programs. Nucleic Acids Res. 25, 3389–3402 (1997).10.1093/nar/25.17.3389 ↗
61. Katoh, K. & Standley, D. M. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Mol. Biol. Evol. 30, 772–780 (2013).10.1093/molbev/mst010 ↗
62. Okonechnikov, K., Golosova, O. & Fursov, M. UGENE team. Unipro UGENE: a unified bioinformatics toolkit. Bioinformatics 28, 1166–1167 (2012).10.1093/bioinformatics/bts091 ↗
63. Gouy, M., Guindon, S. & Gascuel, O. SeaView version 4: a multiplatform graphical user interface for sequence alignment and phylogenetic tree building. Mol. Biol. Evol. 27, 221–224 (2010).10.1093/molbev/msp259 ↗
64. Kalyaanamoorthy, S., Minh, B. Q., Wong, T. K. F., von Haeseler, A. & Jermiin, L. S. ModelFinder: fast model selection for accurate phylogenetic estimates. Nat. Methods 14, 587–589 (2017).10.1038/nmeth.4285 ↗
65. Trifinopoulos, J., Nguyen, L.-T., von Haeseler, A. & Minh, B. Q. W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Res. 44, W232–W235 (2016).10.1093/nar/gkw256 ↗
66. Edler, D., Klein, J., Antonelli, A. & Silvestro, D. raxmlGUI 2.0: a graphical interface and toolkit for phylogenetic analyses using RAxML. Methods Ecol. Evol. 12, 373–377 (2021).10.1111/2041-210X.13512 ↗
67. Rambaut, A. & Drummond, A. J. FigTree: tree figure drawing tool, v. 1.2.2. http://tree.bio.ed.ac.uk/software/figtree/ (2008).
68. Crous, P. W., Verkley, G. J. M., Groenewald, J. Z. & Samson, R. A. Fungal Biodiversity. CBS Laboratory Manual Series (Westerdijk Fungal Biodiversity Institute, 2009).
69. Hammer, O., Harper, D. A. T. & Ryan, P. D. Palaeontological statistics software package for education and data analysis. Palaeontol. Electron. 4, 1–9 (2001).
70. Tiago, I., Chung, A. P. & Veríssimo, A. Bacterial diversity in a nonsaline alkaline environment: heterotrophic aerobic populations. Appl. Environ. Microbiol. 70, 7378–7387 (2004).10.1128/AEM.70.12.7378-7387.2004 ↗
71. Albertano, P. & Urzì, C. Structural interactions among epilithic cyanobacteria and heterotrophic microorganisms in Roman hypogea. Micro. Ecol. 38, 244–252 (1999).10.1007/s002489900174 ↗
72. Guggiari, M. et al. Experimental calcium-oxalate crystal production and dissolution by selected wood rot fungi. Int. Biodeterior. Biodegrad. 65, 803–809 (2011).10.1016/j.ibiod.2011.02.012 ↗
73. Savković, Ž. et al. Diversity and biodeteriorative potential of fungal dwellers on ancient stone stela. Int. Biodeterior. Biodegrad. 115, 212–223 (2016).10.1016/j.ibiod.2016.08.027 ↗
74. Trovão, J., Tiago, I., Catarino, L., Gil, F. & Portugal, A. In vitro analyses of fungi and dolomitic limestone interactions: bioreceptivity and biodeterioration assessment. Int. Biodeterior. Biodegrad. 155, 105107 (2020).10.1016/j.ibiod.2020.105107 ↗
75. Paiva, D. S., Fernandes, L., Portugal, A. & Trovão, J. First genome sequence of the microcolonial black fungus Saxispiralis lemnorum MUM 23.14: insights into the unique genomic traits of the Aeminiaceae family. Microorganisms 12, 104 (2024).10.3390/microorganisms12010104 ↗
76. Liu, B., Fu, R., Wu, B., Liu, X. & Xiang, M. Rock-inhabiting fungi: terminology, diversity, evolution and adaptation mechanisms. Mycology 13, 1–31 (2022).10.1080/21501203.2021.2002452 ↗
77. De Leo, F., Marchetta, A. & Urzì, C. Black fungi on stone-built heritage: current knowledge and future outlook. Appl. Sci. 12, 3969 (2022).10.3390/app12083969 ↗
78. Coutinho, M. L. et al. Biodeterioration of majolica glazed tiles by the fungus Devriesia imbrexigena. Constr. Build Mater. 212, 49–56 (2019).10.1016/j.conbuildmat.2019.03.268 ↗
79. Macedo, M. F., Miller, A. Z., Dionísio, A. & Saiz-Jimenez, C. Biodiversity of cyanobacteria and green algae on monuments in the Mediterranean Basin: an overview. Microbiology 155, 3476–3490 (2009).10.1099/mic.0.032508-0 ↗
80. Burgio, L. & Clark, R. J. Library of FT-Raman spectra of pigments, minerals, pigment media and varnishes, and supplement to existing library of Raman spectra of pigments with visible excitation. Spectrochim. Acta A Mol. Biomol. Spectrosc. 57, 1491–1521 (2001).10.1016/S1386-1425(00)00495-9 ↗
81. Lombard, L. et al. Generic hyper-diversity in Stachybotriaceae. Persoonia 36, 156–246 (2016).10.3767/003158516X691582 ↗
82. Bao, D.-F. et al. Taxonomy, phylogeny and evolution of freshwater Hypocreomycetidae (Sordariomycetes). Fungal Divers. 121, 1–94 (2023).10.1007/s13225-023-00521-8 ↗
83. Luo, Z.-L. et al. Freshwater Sordariomycetes. Fungal Divers. 99, 451–660 (2019).10.1007/s13225-019-00438-1 ↗
84. Andersen, B. et al. Molecular and phenotypic descriptions of Stachybotrys chlorohalonata sp. nov. and two chemotypes of Stachybotrys chartarum found in water-damaged buildings. Mycologia 95, 1227–1238 (2003).10.1080/15572536.2004.11833031 ↗
85. Steinert, K. et al. Tools to make Stachybotrys chartarum genetically amendable: key to unlocking cryptic biosynthetic gene clusters. Fungal Genet. Biol. 172, 103892 (2024).10.1016/j.fgb.2024.103892 ↗
86. Frazer, S. et al. Impact of environmental factors on growth and satratoxin G production by strains of Stachybotrys chartarum. World Mycotoxin J. 5, 37–43 (2012).10.3920/WMJ2011.1329 ↗
87. Miller, A. Z., Dionísio, A., Laiz, L., Macedo, M. F. & Saiz-Jimenez, C. The influence of inherent properties of building limestones on their bioreceptivity to phototrophic microorganisms. Ann. Microbiol. 59, 705–713 (2009).10.1007/BF03179212 ↗
88. Benavente, D. et al. Salt damage and microclimate in the Postumius Tomb, Roman Necropolis of Carmona, Spain. Environ. Earth Sci. 63, 1529–1543 (2011).10.1007/s12665-010-0815-9 ↗
89. Sena da Fonseca, B., Ferreira Pinto, A. P., Rodrigues, A., Piçarra, S. & Montemor, M. F. The role of properties on the decay susceptibility and conservation issues of soft limestones: contribution of Ançã stone (Portugal). J. Build. Eng. 44, 102997 (2021).10.1016/j.jobe.2021.102997 ↗
90. Barnés-Guirado, M., Stchigel, A. M. & Cano-Lira, J. F. A new genus of the Microascaceae (Ascomycota) family from a hypersaline lagoon in Spain and the delimitation of the genus Wardomyces. J. Fungi 10, 236 (2024).10.3390/jof10040236 ↗
91. Unković, N. et al. Biodegradative potential of fungal isolates from sacral ambient: in vitro study as risk assessment implication for the conservation of wall paintings. PLoS ONE 13, e0190922 (2018).10.1371/journal.pone.0190922 ↗
92. Warscheid, T. & Braams, J. Biodeterioration of stone: a review. Int. Biodeterior. Biodegrad. 46, 343–368 (2000).10.1016/S0964-8305(00)00109-8 ↗
93. Gadd, G. M. Metals, minerals and microbes: geomicrobiology and bioremediation. Microbiology 156, 609–643 (2010).10.1099/mic.0.037143-0 ↗
94. Burford, E. P., Kierans, M. & Gadd, G. M. Geomycology: fungi in mineral substrata. Mycologist 17, 98–107 (2003).10.1017/S0269915X03003112 ↗
95. Gadd, G. M. & Raven, J. A. Geomicrobiology of eukaryotic microorganisms. Geomicrobiol. J. 27, 491–519 (2010).10.1080/01490451003703006 ↗
96. Balland-Bolou-Bi, C. et al. Effect of cultivable bacteria and fungi on the limestone weathering used in historical buildings. Diversity 15, 587 (2023).10.3390/d15050587 ↗
97. Boniek, D., de Abreu, C. S., dos Santos, A. F. B. & de Resende Stoianoff, M. A. Filamentous fungi in Brazilian indoor cultural heritage as potential risk to human health and biodeterioration of artworks. Air Qual. Atmos. Health 15, 339–346 (2022).10.1007/s11869-021-01108-5 ↗
98. Rojas, T. I., Aira, M. J., Batista, A., Cruz, I. L. & González, S. Fungal biodeterioration in historic buildings of Havana (Cuba). Grana 51, 44–51 (2012).10.1080/00173134.2011.643920 ↗
99. Pinna, D. Fungal physiology and the formation of calcium oxalate films on stone monuments. Aerobiologia 9, 157–167 (1993).10.1007/BF02066257 ↗
100. Fomina, M., Burford, E. P., Hillier, S., Kierans, M. & Gadd, G. M. Rock-building fungi. Geomicrobiol. J. 27, 624–629 (2010).10.1080/01490451003702974 ↗