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Black Fungi are one of the main group of microorganisms responsible for the biodeterioration of stone cultural heritage artifacts. The term “black fungi” refers to a very huge group of dematiaceous fungi, unrelated phylogenetically, which have in common the presence of melanin in the cell wall that confers an olive brown appearance to the colony. Another common characteristic is the ability to withstand hostile environments such as scarcity of nutrients, high solar irradiation, scarcity of water, high osmolarity, and low pH.
Class/Order | Genera * | Substrate | Environmental and Climatic Features | Alterations Associated to Fungal Colonization | Refs |
---|---|---|---|---|---|
Dothideomycetes incertae sedis | Coniosporium | Calcarenite, granite, limestone, marble | Mediterranean climate, urban environment | Grayish-black patina, pitting, black spots, greenish to dark green patina, crater shaped lesions, chipping, exfoliation, sugaring, crumbling, superficial deposit, and biofilm | [36][38][43][44][45][46][47][48][49] |
Dothideomycetes/Capnodiales incertae sedis | Capnobotryella | Limestone, marble | Mediterranean climate, continental climate, and urban environment |
Black spots, crater shaped lesions, chipping, exfoliation, sugaring, crumbling, pitting, superficial deposit, and biofilm formation | [45][48][50][51][52] |
Constantinomyces | Sandstone | Urban environment, temperate climate | Discolorations, patina | [53] | |
Pseudotaeniolina | Marble, sandstone | Mediterranean climate, arid and desert climate | Biological green patina | [54][55][56] | |
Dothideomycetes/Capnodiales | Aeminium | Limestone | Temperate climate | Black discoloration with salt efflorescence | [57] |
Dothideomycetes/Cladosporiales | Cladosporium | Calcarenite, granite, limestone, marble, plaster, sandstone, tufa | Ubiquitous worldwide distribution in indoor environments and outdoor | Dark alterations, black spots, black patinas, detachment of marble grains, light grayish patina, crater shaped lesions, chipping, exfoliation, sugaring, crumbling, pitting, superficial deposit, biofilm, black crusts, green biofilm with salt efflorescence, stone erosion and disintegration, and discoloration | [26][39][46][48][49][58][59][60][61][62][63][64][65][66][67] |
Verrucocladosporium | Limestone, marble, sandstone | Mediterranean climate, temperate climate, and urban environment | Black patina, discoloration | [36][53] | |
Dothideomycetes/Dothideales | Aureobasidium | Granite, limestone, marble, plaster, sandstone |
Urban environment, Mediterranean climate, temperate climate, indoor environment, and urban environment | Black patina, black spots, detachments, superficial deposit, biofilm, discolorations with or without salt efflorescence, black crusts, and stone erosion and disintegration | [36][39][45][49][53][63][64][65][68] |
Dothideomycetes/Mycosphaerellales | Salinomyces | Marble, sandstone | Mediterranean climate | Black patina | [36] |
Neocatenulostroma | Limestone, sandstone | Temperate climate, urban environment | Discolorations and/or patina, structural damage | [53] | |
Neodevresia | Limestone, marble, plaster, tufa | Mediterranean climate | Black patina, discolorations, structural damage | [36][53][55][63] | |
Saxophila | Marble | Mediterranean climate | Black patina | [36] | |
Vermiconidia | Limestone, marble, travertine | Mediterranean climate, urban environment | Black patina | [36] | |
Dothideomycetes/Neophaeothecales | Neophaeotheca | Marble | Mediterranean climate | Black patina | [36] |
Dothideomycetes/Pleosporales | Alternaria | Calcarenite, granite, limestone, marble, plaster, tufa | Ubiquitous worldwide distribution in indoor environments and outdoor | Black spots, black patina, detachment of marble grains, greenish to dark green patina, biofilm, black crusts, green-black patina; and blackish patina | [39][46][49][58][59][60][63][64][66][67] |
Epicoccum | Granite, limestone, marble | Urban environment, mediterranean climate, and temperate climate | Black spots, black patinas, detachment, superficial deposit, biofilm, blackish patina, green biofilm, and dark and green biofilm with salt efflorescence | [39][45][49][60][64] | |
Phoma | Calcarenite, granite, limestone, marble, plaster, tufa | Mediterranean climate, temperate climate, urban environment, continental-cold climate, and indoor and outdoor environments | Black spots, black patinas, detachment of marble grains; color changes, crater shaped lesions, chipping and exfoliation, sugaring, crumbling, pitting, superficial deposit, biofilm, and black crusts | [39][46][48][49][58][63] | |
Dothideomycetes/Venturiales | Ochroconis | Calcarenite | Subterranean environment | Black patina | [69] |
Eurotiomycetes incertae sedis | Sarcinomyces | Marble | Mediterranean climate | Black spots | [70] |
Eurotiomycetes/Chaetothyriales | Cyphellophora sp. | Plaster | Mediterranean climate | Black/grayish patina | [63] |
Exophiala | Calcarenite, limestone, marble, sandstone |
Mediterranean climate, urban environment, temperate climate, and hypogean environment | Dark alterations, black spots, black patinas, detachment of marble grains, discolorations, and visible structural damage | [26][36][39][45][53][71] | |
Lithophila | Limestone, marble | Mediterranean climate, urban environment, and dry continental climate |
Black spots, black patinas, detachment of marble grains | [36][39][72] | |
Knufia | Limestone, marble, sandstone travertine |
Mediterranean climate, urban environment, continental temperate climate, and dry continental climate | Black and grey spots, dark macropitting, biopitting, crater shaped lesions, chipping, exfoliation, sugaring, crumbling, discolorations, patina, and visible structural damage | [36][40][43][45][48][53][72][73][74] | |
Rhinocladiella | Marble | Mediterranean climate | Black spots, crater shaped lesions, chipping and exfoliation, sugaring, crumbling, and pitting | [48] | |
Eurotiomycetes/Mycocaliciales | Mycocalicium | Marble | Mediterranean climate, urban environment | Black spots, crater shaped lesions, chipping and exfoliation, sugaring, crumbling, and pitting | [45][48] |
Taxon | Strain | ITS rDNA |
---|---|---|
Capnobotryella antalyensis | MA 4615 | AJ972858 |
Capnobotryella antalyensis | MA 4624 | AJ972850 |
Capnobotryella antalyensis | MA 4766 | AJ972851 |
Capnobotryella antalyensis | MA 4775 | AJ972860 |
Capnobotryella isilogui | MA 3619 | AM746201 |
Capnobotryella erdogani | MA 4625 | AJ972857 |
Capnobotryella kirizoglui | MA 4899 | AJ972859 |
Capnobotryella sp. | MA 4701 | AJ972856 |
Capnobotryella sp. | MA 4697 | AJ972855 |
Capnobotryella sp. | MA 3615 | AM746203 |
Neodevriesia modesta | CCFEE 5672 | KF309984 |
Neodevriesia simplex | CCFEE 5681 | KF309985 |
Neodevriesia sardiniae | CCFEE 6202 | KP791765 |
Neodevriesia sardiniae | CCFEE 6210 | KP791766 |
Saxophila tyrrhenica | CCFEE 5935 | KP791764 |
Aeminium ludgeri | E12 | MG938054 |
Aeminium ludgeri | E16 | MG938061 |
Neocatenulostroma sp. | CR1 | KY111907 |
Constantinomyces sp. | CR21 | KY111911 |
Pseudaeniolina globosa | DPS10 | MH396690 |
Pseudotaeniolina globosa | CBS109889 | NR136960 |
Pseudotaeniolina globosa | CCFEE5734 | KF309976 |
Vermiconidia calcicola | CBS 140080 | NR_145012 |
Vermiconidia calcicola | CCFEE 5780 | KP791761 |
Vermiconidia flagrans | CCFEE 5922 | KP791753 |
Coniosporium uncinatum | CBS 100219 | AJ244270 |
Coniosporium apollinis | CBS 100213 | AJ244271 |
Coniosporium apollinis | CBS 352.97 | NR159787 |
Coniosporium apollinis | CBS 100216 | AJ244272 |
Coniosporium apollinis | QIIIa | MH023395 |
Lithophila catenulata | BJ10118 | JN650519 |
Lithophila guttulata | M1 | MW361305 |
Lithophila guttulata | CCFEE 5884 | KP791768 |
Lithophila guttulata | CCFEE 5907 | KP791773 |
Knufia mediterranea | CCFEE 5738 | KP791791 |
Knufia mediterranea | CCFEE 6211 | KP791793 |
Knufia vaticanii | CCFEE 5939 | KP791780 |
Knufia calcarecola | SL11033 | JQ354925 |
Knufia calcarecola | CGMCC 3.17222 | KP174862 |
Knufia marmoricola | CCFEE 5895 | KP791775 |
Knufia marmoricola | CCFEE 5716 | KP791786 |
Knufia perforans | CBS 885.95 | AJ244230 |
Knufia karalitana | CCFEE 5732 | KP791782 |
Knufia karalitana | CCFEE 5929 | KP791783 |
Knufia petricola | CCFEE 726.95 | KC978746 |
Knufia petricola | CBS 600.93 | KC978744 |
Knufia petricola | IMI38917 | AJ507323 |
Knufia petricola | D1 | JF749183 |
Knufia petricola | M4 | FJ556910 |
Knufia sp. | QIIa | MH023393 |
Knufia sp. | QIIb | MH023394 |
Very little is said regarding the effectiveness of treatments against black fungi. Black fungi, especially meristematic ones, are very difficult to eradicate and tend to be one of the first colonizers after cleaning procedures. In order to achieve protection of an artifact, both indirect and direct methods should be implemented. Direct treatments aiming to kill/reduce black fungi on the stone should be different on the basis of their colonization pattern (diffuse patina, spot-like colonization, or intercrystalline growth) and on the characteristics of the environment. Among the potential methods commonly used to control biodeterioration, physical methods such as mechanical removal and UV and heat shock treatments[81][82], are not very effective against black fungi[83][84]. Regarding chemical methods, in laboratory conditions, classical biocides (e.g., Preventol RI 50, Biotin R, RocimaTM 103) are still the most effective [83][85] and in the field they produce efficient results during cleaning procedures. Plant based extracts show a scarce effectiveness against fungi, and this difficult group of microorganisms is not even taken into account to assess their activity[86]. Nanoparticles are commonly used as biocides due to their activity against algae, cyanobacteria, and most bacteria, but they are not really satisfactory against black fungi. Protective coatings with antifouling properties may have various effects. In fact, TiO2 based coatings, pure or doped with Ag, show a good effect but are limited to a short/medium term after application [86]. However, in both laboratory and field conditions, after treatments with titania-based coatings, black fungi are the first to recolonize the stone surface in dry environments, while algae first appears in damping walls[63]. Very recently, in laboratory conditions, cholinium@Il based coatings have shown that the use of Il’s with a 12 C chains and DBS as anion in combination with nanosilica coatings (e.g., Nano Estel) could be effective against the colonization of black fungi for a period of time over 30 months [87]. One possible explanation of this scarce effectiveness of most treatments against black fungi is that they possess a genetic resistance to environmental stresses, as reported the previous paragraphs. Therefore, the different mechanisms concurring to the stress protection response may interfere to the biocidal treatments. Understanding the cause of their resilience could improve the strategies for their control.