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International Biodeterioration & Biodegradation 60 (2007) 231–237 www.elsevier.com/locate/ibiod
Aeromycological study in the Cathedral of Santiago de Compostela (Spain) M.J. Airaa, V. Jatob, A.M. Stchigelc, F.J. Rodrı´ guez-Rajob,, E. Piontellid a
Department of Botany, Faculty of Pharmacy, University of Santiago de Compostela, Campus South, Santiago de Compostela, Spain b Department of Plant Biology and Soil Sciences, Sciences Faculty Ourense, University of Vigo, Campus As Lagoas, Spain c Microbiology Unit, Medicine and Health Sciences Faculty, Rovira y Virgili University, Tarragona, Spain d Mycology Chair, Medicine School, Valparaı´so University, Valparaı´so, Chile Received 26 June 2006; received in revised form 27 January 2007; accepted 28 February 2007 Available online 23 May 2007
Abstract A study of airborne fungi was carried out in the architectural complex of the Cathedral of Santiago de Compostela (Spain) during 2002, by using viable volumetric sampling methods. This resulted in a total of 35 identified taxa, of which the most abundant were: Alternaria, Aspergillus, Cladosporium and Penicillium. Sampling was completed with data from the outdoor atmosphere and swab samples in specific places. In general there were no statistically significant indoor/outdoor differences and in both cases the highest CFU m-3 were obtained during the spring-summer. Similar relatively low numbers of the same fungi were likewise detected at different points in the Cathedral nave, while up to nearly 6500 CFU m-3 were recorded in the Corticela Chapel. The study of intradiurnal levels carried out in the Cathedral nave reveals greater abundance of fungal concentrations at 13:00 h, the moment of massive influx of visitors in the Cathedral, with 406 CFU m-3 compared to the 380 CFU m-3 sampled at 9:00 h and the 350 CFU m-3 at 21:00 h. The whole investigation is the first study of the atmospheric fungal content of the Cathedral of Santiago de Compostela. r 2007 Elsevier Ltd. All rights reserved. Keywords: Aerobiology; Biodeterioration; Fungal spores; Santiago of Compostela Cathedral; Volumetric sampling
1. Introduction Different microorganisms such as fungi, algae and bacteria may have a negative effect on the preservation of artistic-historical heritage, especially when microclimatic conditions favour their development. Numerous museum complexes therefore employ systematic temperature and humidity monitoring, with the aim to prevent or slow down their growth (De Nuntiis et al., 2004). Some authors also report interaction between microscopic fungi and arthropods on the surface of wall paintings, which increases their alteration (Hoffland et al., 2004). These biological agents affect not only the aesthetical appearance but also the structure of materials. Contributing to their preservation therefore is the main Corresponding author. Tel.: +34 988387054; fax: +34 988387001.
E-mail address:
[email protected] (F.J. Rodrı´ guez-Rajo). 0964-8305/$ - see front matter r 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.ibiod.2007.02.007
goal of studies of the mycobiota carried out in heritage buildings. Studies on biodeterioration deal with different topics and substrates. There is great diversity in relation to the subjects of such studies since they encompass any type of work of art (Guglielminetti et al., 1994; Gorbushina and Palinska, 1999; Pitzurra et al., 1999; Maggi et al., 2000), as well as the materials of the buildings (Caretta and Piontelli, 1998; Petushkova and Kandyba, 1999). However, despite the interest of this type of study, those related to fungal biodeterioration are not very common; in the case of the Cathedral of Santiago de Compostela, no previous studies have been conducted. The main objective of this study was therefore to ascertain the mycobiota atmospheric content of the city’s most important architectural complex, which forms part of its World Heritage as pointed out by UNESCO, to identify the agents causing the proliferation of these
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microorganisms and its favouring factors. This will pave the way for a proposal regarding any necessary conservation work to the building, which is showing some evidence of alteration caused by fungi in certain areas. Indirectly, this type of study may have public health applications, since the calculation of atmospheric fungal content and the identification of certain human pathogens can show whether an atmosphere is healthy or not and point to potential allergy risks (Terr, 2004). In this regard, fungi such as Stachybotrys are a usual component of the mycobiota inside buildings with water-damaged walls (Terr, 2001). 2. Material and methods This study was carried out during the year 2002 in the main monumental complex of Santiago de Compostela (Spain), made up of the Cathedral and Museum. Atmospheric spore samples were collected by using two viable volumetric sampling systems. These are a six-stage Andersen sampler (A) and a single-stage Burkard portable air sampler for agar plates (BPC) including 1 sampling plate. The sampling lasted 10 min been the collectors placed 1 m above ground level. The results were expressed in CFU m-3 and the positive-hole correction in both samplers was applied (Mehta et al., 1996). Random swab samples were also collected, to identify the fungal types present on certain works of artistic value or ‘‘areas’’ showing fungal growth. The entire methodology used in the study was explained in detail by Aira et al. (2005). The preparation of culture media was conducted following the instructions of Hoog et al. (2000). With the aim to ascertain the fungal quantity and diversity throughout the year, sampling was carried out every four weeks at five points in the Cathedral nave, four at the extremes (identified as Point 1: Quintana, Point 2: Azabacheria, Point 3: Portico and Point 4: Platerias) and one at the centre (Point 5: Nave centre) of the Latin-cross ground plan. Sampling was also carried out outside the Cathedral (Point 6: Outdoors) as a reference (Fig. 1). Friday was chosen as a representative day of the Cathedral’s normal activity, i.e. without a massive influx of visitors that could affect atmospheric fungal content. At each sampling time and location, single samples were taken using the BPC and commercial Sabouraud agar amended with chloramphenicol to control bacteria. To evaluate efficiency of Sabouraud agar culture media in comparison with malt extract agar (MEA) and oatmeal agar (OA), sampling was carried out in the central area of the nave (Point 5: Nave centre) under the
same conditions as the previous ones, with the exception that the Sabouraud was replaced by MEA and oat agar (OA). The influence of the high quantity of people visiting the Cathedral on Sundays was studied, since religious services may be attended by more than 5000 people. To evaluate whether this variable influences fungal content, every fourth Sunday, air in the centre of the Nave (Point 5) and outdoors was sampled onto Sabouraud agar using the BPC. The samples were conducted when low numbers of visitors are in the Cathedral (09.00 h), in the moment of the high influx of visitors (13.00 h) and in an hour of the day with medium people presence (21.00 h), so as to assess its impact on the data obtained. Other seasonal samplings (Spring: April 21, Summer: August 11, Autumn: November 3 and Winter: December 29, January 27, February 1st) was carried out (Fig. 1) inside the Cathedral chapels (Point a: Corticela and Point b: Santı´ simo) and inside the Museum’s exhibition rooms (Point c: Library and Point d: Goya), where possible alterations produced by fungi were detected, to determine the particular atmospheric fungal content in each area. In this case both volumetric sampling systems were used, with Sabouraud agar as before as culture medium. Finally, swabbing was used to obtain samples directly from the walls inside of the Chapel of Corticela and the Chapel of Santı´ simo, the sides of the ‘‘Po´rtico de la Gloria’’ (which is located in the Cathedral nave) and from different artistic pieces on display inside the Museum’s exhibition rooms Library and Goya. The culture plates obtained were incubated at 25 1C during 7 d, after which the colonies were counted and isolated. Simultaneously, in all of the samples, temperature and humidity was measured using a weatherlink meteorological station; days with rainfall were also recorded due to the possible influence on the outdoor atmosphere’s fungal content. Finally, we applied Scheffe´’s test to study the homogeneity of the populations under study (Wassertheil-Smoller, 2004), which would enabled us to identify the presence of differences in the quantity of mycobiota content between the sampling points. Spearman’s correlation test was used to find a possible relationship between the fungal levels and meteorological factors (temperature and humidity) or visitors affluence.
3. Results 3.1. Quantitative data The total number of colony forming units in all of the samples taken in the Cathedral nave (38,137 CFU) was higher than that of outdoor air (20,509 CFU). However, despite this apparent predominance of fungi inside the Cathedral (Table 1), when applying Scheffe´’s test no general significant differences (po0.05) were revealed when comparing each sampling point inside the nave with the outside. In general, the results obtained throughout the year inside the nave of the Cathedral show that the maximum concentration is recorded during spring and summer months (with the exception of sampling in April), decreasing considerably in the first 3 months of the year. The
Table 1 Mean of CFU and range of counts for each sampling point in the nave of the Cathedral
Fig. 1. Sampling site points in the nave Cathedral (1—Quintana, 2— Azabacherı´ a, 3—Po´rtico, 4—Platerias, 5—Nave Centre, 6—Outdoors) and seasonal sampling site points (a—Corticela Chapel, b—Santı´ simo Chapel, c—Library Museum room, d—Goya Museum room).
Point 1 Point 2 Point 3 Point 4 Point 5 Quintana Azabacherı´ a Po´rtico Platerı´ as Nave center Range 135–560 Mean 327
145–615 350
70–610 85–570 388 339
105–560 357
Outdoors
55–540 309
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700
1
2
3
4
5
233
Out
600 CFU-s/m3
500 400 300 200 100 0 Jan 25 Feb 22 Mar 22 Apr 10 May 17 Jun 14 Jul 12
Aug 9
Sep 6
Oct 4
Nov 1 Nov 29 Dec 27
Fig. 2. Representation of number of CFU m-3 throughout the year in the Cathedral nave sampling points 1—Quintana, 2—Azabacherı´ a, 3—Po´rtico, 4— Platerias, 5—Nave Centre and 6—Outdoors.
minimum values were obtained in April at all sampling points, with the exception of point 2, which was recorded in October. The most marked differences among the five sampling points in the nave were recorded in September (points 1 and 2), January (points 3 and 4) and November (points 3 and 5); comparing the indoor and outdoor atmospheres, the greatest values occurred in January and August with amounts greater than 300 CFU m-3 (Fig. 2). In relation to culture media, the most effective medium from a quantitative point of view was MEA, in which a mean of 380 CFU m-3 was recorded, followed by Sabouraud agar with 357 CFU m-3 and finally OA with 308 CFU m-3. The greatest differences in relation to culture media were obtained between OA and MEA, although they were not statistically significant. Nevertheless, the highest individual value corresponded to the sampling carried out on March 24th with Sabouraud (560 CFU m-3). The study of intradiurnal levels carried out in the Cathedral nave reveals greater abundance of fungal concentrations around midday (at 13:00 h) in the moment of the high influx of visitors, with 406 CFU m-3, compared to the 380 CFU m-3 at 09:00 h and 350 CFU m-3 at 21:00 h. The levels recorded in the early morning are higher outdoors, with 427 CFU m-3 at 09:00 h compared to the 305 CFU m-3 sampled at 13.00 h and 385 CFU m-3 at 09.00 h (Table 2). In the nave of the Cathedral, the values obtained at 13:00 h were higher than those at 09:00 h in 9 of the 13 sampling days. The highest differences, greater than 100 CFU m-3, were obtained in May, January and October. The values recorded at 13:00 h were likewise higher than those at 21:00 h in 10 of the 13 sampling days. The most marked differences (greater than 100 CFU m-3) occurred in May, September and March. The comparison of indoor/outdoor results shows that the differences are greatest at 13:00 h, when the indoor concentrations are very high in relation to the outdoor ones. Even in the case of those days when the outdoor concentrations are highest, the difference decreases between 9:00 and 13:00 h. These differences seem to indicate the influence of the massive influx of visitors in the
Table 2 Number of CFU m-3 sampled at different hours of the day (09.00, 13.00 and 21.00 h) indoor (in the sampling point 5: Nave Centre) and outdoor Indoor
27 1 24 21 19 16 14 11 8 6 3 1 29
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Dec
Mean
Outdoor
09.00
13.00
21.00
09.00
13.00
21.00
385 530 505 195 245 460 580 455 470 285 405 210 210
510 575 530 220 455 495 630 490 465 395 95 200 220
550 540 380 215 260 495 575 410 275 320 165 145 215
255 550 390 105 355 575 650 445 530 470 460 435 335
255 85 130 95 315 505 645 515 205 505 185 275 255
465 115 295 320 245 530 610 480 535 430 480 260 245
380
406
350
427
305
385
Cathedral around mid-day, despite this differences were not significant according to Spearman’s correlation test. In the seasonal sampling (Table 3) carried out inside the Cathedral chapels and inside the two Museums rooms, the number of colony forming units was higher using the Andersen sampler, and the indoor levels were higher than the outdoor ones in both cases. This fact should be caused as a six-stage sampler achieves a better separation of spores than a one-stage sampler, and also because it sampled a larger volume of air. Some unexpected results such us, the winter BPC count inside Library Museum room were lower than outdoors, the summer BPC count in the Goya Museum room were higher than the Andersen count and the outdoor level in autumn were higher than inside the Santı´ simo Chapel, Library Museum room and Goya Museum room values, could be caused by the fact only that a single sample has been taken for each season. The highest levels inside the two chapels were obtained at different times of year, as it was the case of the two Museum rooms, while outdoors they were recorded during
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Table 3 Number of CFU m-3 obtained in the seasonal sampling of different Cathedral Chapels and rooms Spring
Summer
Autumn
Winter
Annual total
Corticela Chapel Andersen 2943 BPC 545 Temperature 13 Humidity 65
1547 505 21 58
1470 335 21 66
509 484 12 76
6469 1869 13
Santı´simo Chapel Andersen 539 BPC 480 Temperature 17 Humidity 57
373 525 21 64
360 300 25 60
966 490 14 80
2238 1795
Library Museum room Andersen 733 BPC 460 Temperature 15 Humidity 56
266 175 22 61
348 215 22 54
187 270 16 65
1534 1120
Goya Museum room Andersen 255 BPC 335 Temperature 15 Humidity 54
113 210 23 55
227 185 22 50
304 405 17 63
899 1135
Outdoors Andersen BPC Temperature Humidity
180 165 23 53
540 390 17 57
96 360 14 66
1051 920
235 5 21 45
The values of the meteorological data during the samples were show.
autumn, probably due to the influence of local mycobiota content and meteorological variables. Temperature in the indoor atmospheres sampled oscillated between 12 and 15 1C and humidity between 54% and 80%. Spearman correlation test was applied to ascertain the association degree between fungal levels and temperature and humidity, revealing a positive correlation with humidity (R ¼ 0.475) and a negative one with temperature (R ¼ 0.502), with 99% significance in both cases. 3.2. Qualitative data A total of 35 taxa were identified and the greatest fungal diversity was obtained in the samples taken in the Cathedral nave, with 31 taxa identified, followed by the outdoor atmosphere with 23 taxa and the Museum samples, in which a total of 14 taxa were identified (Table 4). The four most abundant genera, regardless of the sampling point, were: Alternaria, Aspergillus, Cladosporium and Penicillium, with the first three dominant in the nave while Alternaria stands out slightly outdoors (Table 5). The most abundant Alternaria species was Alternaria alternata (Fr.) Keissl., while in the case of Cladosporium, there was a codominance between cladosporioides-type
Cladosporium, which includes species such as Cladosporium cladosporioides (Fresen.) G.A. de Vries, Cladosporium avellaneum G.A. de Vries, Cladosporium elatum (Harz) Nannf., Cladosporium oxysporum Berk. and M.A. Curtis, among others, and herbarum-type Cladosporium, which encompasses Cladosporium herbarum (Pers.) Link, Cladosporium sphaerospermum Penz. and Cladosporium macrocarpum Preuss (Mediavilla et al., 1995). Within the Aspergillus genus, the most abundant species was Aspergillus fumigatus Fresen., representing up to 28% of the outdoor fungi count, while Penicillium purpurogenum Stoll was dominant within its genus, with a maximum representation of 48% (Table 6). In relation to the results obtained by swabbing the inside walls of the Santı´ simo Chapel, we identified Arthrobotrys oligospora Fresen., Aspergillus nidulans (Eidam) G. Winter, Aspergillus niger Tiegh., Aspergillus versicolor (Vuill.) Tirab., Aureobasidium pullulans (de Bary) G. Arnaud, Curvularia lunata (Wakker) Boedijn, Penicillium aspergilloides Rudakov, P. purpurogenum Stoll and Trichoderma viride Pers. in green and brownish-grey patinas, while A. pullulans(de Bary) G. Arnaud and Cunninghamella elegans Lendn. formed blackish powdery patinas on the sides of the ‘‘Po´rtico de la Gloria’’ and P. purpurogenum Stoll colonised several religious pieces of great artistic value in the inside walls of the Corticela Chapel. Different species of Penicillium, A. fumigatus and Absidia corymbifera (Cohn) Sacc. and Trotter were identified with the same technique in several display cabinets inside the Cathedral Museum’s library room; A. corymbifera (Cohn) Sacc. and Trotter was also isolated from a powdery sample from the Museum’s Choir Room. In the Goya Museum room, where mainly tapestries are on display, there were also different species of Penicillium and Chaetomium, A. fumigatus Fresen., A. versicolor (Vuill.) Tirab., C. cladosporioides (Fresen.) G.A. de Vries and Scopulariopsis brumptii Salvanet-Duvalin. 4. Discussion The study of the Cathedral nave’s fungal representation reveals the existence of mean fungal levels greater than 325 CFU m-3 at all sampling points. The application of Scheffe´’s statistical test to the results indicates that there are no significant differences, but to confirm this result a larger study by means a higher number of plates exposed should be done, as microorganisms in indoor air are not uniformly dispersed, and samples taken at the same location within minutes of each other could be markedly different. At all of the sampling points the mean levels were higher than those recorded outdoors (309 CFU m-3), although such differences were not significant. On the other hand, inside the Corticela and Santı´ simo chapels, using the data obtained with the BPC (which is, therefore, comparable to that of the similar points in the nave), the mean values are higher, and surpass those of the outdoor atmosphere.
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Table 4 Taxa identified by using volumetric sampling methods in the Cathedral nave, Chapels and Museum rooms, and outside Cathedral Nave
Acremonium sp. Alternaria sp. Arthrinium phaeospermum (Corda) M.B. Ellis Aspergillus candidus Link Aspergillus flavus Link Aspergillus fumigatus Aspergillus niger Fresen. Aspergillus oryzae (Ahlb.) E. Cohn Aspergillus terreus Thom Aspergillus versicolor (Vuill.) Tirab. Aspergillus sp. Beauveria bassiana (Bals.-Criv.) Vuill. Botrytis cinerea Pers. Chaetomium globosum Kunze Chaetomium indicum Corda Cladosporium sp. Cylindrocarpon sp. Microsphaeropsis pseudaspera B. Sutton Mucor mucedo L. Paecilomyces lilacinus (Thom) Samson Paecilomyces variotii Bainier Penicillum aurantiogriseum Dierckx Penicillium citrinum Sopp Penicillium crustosum Thom Penicillium funiculossum Thom Penicillium glabrum (Wehmer) Westling Penicillium pinophillum Thom Penicillium purpurogenum Stoll Penicillium spinulosum Thom Penicillium thomii Maire Penicillium sp. Rhizopus stolonifer (Ehrenb.) Vuill. Trichoderma sp. Stachybotrys atra Corda viride Pers.
+ + + + + + + + + + + + + + +
+ + +
+ + + + + + + + +
Cathedral Chapels
Museum rooms
Outdoors
Corticela
Santı´ simo
Library
Goya
+ + +
+ +
+ +
+
+
+ +
+ +
+
+ + + + + +
+ +
+
+
+
+ +
+ + +
+ + + +
+ +
+ +
+
+ +
+ + +
+ + + + + +
+ + +
+
+ +
+
+
+ +
+ +
+ +
+ + + + + +
+
Table 5 Number of colonies identified of the four most abundant genera Cathedral Nave
Alternaria Aspergillus. Cladosporium Penicillium
47 185 580 401
Cathedral Chapels
Museum rooms
Outdoors
Corticela
Santisimo
Library
Goya
15 114 12 81
7 40 17 59
18 33 39 48
12 19 25 28
Applying Scheffe´’s test to the results obtained with the Anderson method reveals the existence of significant differences between those inside the Corticela Chapel and those from the other sampled rooms and with those from outdoors, thereby indicating differential behaviour of its aeromycobiota. The levels attained indoors the Corticela Chapel are higher than 1400 CFU m-3 in three of the four seasonal samples. These quantitative differences may be due to the fact that the physical isolation favours a
53 74 512 186
microclimate that is especially suitable for fungal development. In fact, there is evident deterioration in some wooden beams inside of the Chapel of Corticela, while there is an abundance of patinas on the dampest walls indoors of the Santı´ simo Chapel. In view of the meteorological data, temperature does not seem to pose a risk to the artistic material’s preservation, unlike humidity, whose values exceed recommended levels, especially indoors the Cathedral’s chapels.
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Table 6 Representation of the most abundant species (in %) Cathedral Nave
Number of A. fumigatus isolations Number of Aspergillus sp. isolations A. fumigatus (%) Number of P. purpurogenum isolations Number of Penicillium sp. isolations P. purpurogenum (%)
29 185 16 47 401 12
Cathedral Chapel
Museum rooms
Outdoors
Corticela
Santisimo
Library
Goya
1 114 1 1 81 1
6 40 15 11 59 19
6 33 18 23 48 48
3 19 16 7 28 25
Similar values to those of this study were detected in Rome’s State Archives, where about 300 CFU m-3 were recorded, the most predominant genera being Cladosporium in winter and Penicillium in summer, along with others commonly found throughout the year such as Alternaria, Aspergillus and Chaetomium (Maggi et al., 2000). On the other hand, in India, a study carried out with an Andersen trap in several Delhi libraries showed a much higher level, since the Cladosporium genus alone amounted to 12,322 CFU m-3. Other important genera were Aspergillus, Penicillium and Alternaria (Singh et al., 1995). Qualitatively speaking, the taxa identified in Santiago are the same as those identified in other buildings of the same characteristics, such as Moscow Cathedral, where the most abundant genera were Acremonium and Penicillium (Petushkova and Kandyba, 1999). Studying wall paintings in different Italian monasteries has revealed the presence of fungi such as the genera Alternaria, Aspergillus, Cladosporium, Fusarium, Penicillium and Rhizopus, with the most common species being C. cladosporioides (Fresen.) G.A. de Vries, Alternaria alternata (Fr.) Keissl., Aspergillus niger Tiegh. and Penicillium expansum Link (Guglielminetti et al., 1994; Pitzurra et al., 1999). These results largely coincide with those of a study carried out in different historical buildings in Germany, where genera such as Acremonium, Aureobasidium, Beauveria, Chrysosporium, Engyodontium, Scopularopsis and Verticillium were also identified (Hoffland et al., 2004). The destructive activity of fungi is related to the type of material or substrate that is colonised; thus, many Chaetomium species show cellulolytic activity (Caretta and Piontelli, 1998), while Acremonium sp. has been cited as one of the fungi involved in the biodeterioration of glass pieces (Gorbushina and Palinska, 1999). It is important to point out that many of the species identified in this study are known to cause biodeterioration in cellulolytic material, being capable of damaging books or valuable archival material, which they use as substrate under conditions favouring the germination of their spores (Maggi et al., 2000). The results show an abundant representation of fungi in the architectural complex of the Cathedral of Santiago de Compostela. The whole investigation is only a preliminary
21 74 28 41 186 22
investigation and further research, by means other culture media and a higher number of plates exposed, should be conducted to complete and verify this results with the aim to generate more precise data for the preservation of artistic-historical heritage. Acknowledgements This study was subsidised by the Department of Culture, Social Communication and Tourism of the Xunta de Galicia (PGIDT01PAT38301PR). References Aira, M.J., Jato, V., Iglesias, I., 2005. Air Quality. Pollen and Spores in the Galician Region. Xunta of Galicia, Santiago de Compostela, Spain. Caretta, G., Piontelli, E., 1998. Preserved ascomatal and other fungal structures on the remains of a ninth century Longobard abbess exhumed from a Monastery in Pavia, Italia. Mycopathologia 140, 77–83. De Nuntiis, P., Bitelli, L., Guaraldi, P., Monco, A., Salvi, A., 2004. Intermuseum network for conservation of the artistic heritage: Musa project. In: Polen, A. (Ed.), Proceedings XI International Palynological Congress. University of Co´rdoba, Co´doba pp. 88. Gorbushina, A.A., Palinska, K.A., 1999. Biodeteriorative processes on glass: experimental proof of the role of fungi and cyanobacteria. Aerobiologia 15, 183–191. Guglielminetti, M., De Giuli, C., Radaelli, A., Bistini, F., Carruba, G., Spera, G., Caretta, G., 1994. Mycological and ultrastuctural studies to evaluate biodeterioration of mural paintings. Detection of fungi and mites in frescos of the Monastery of St. Damian in Assisi. International Journal of Biodeterioration and Biodegradation 33-3, 269–283. Hoffland, E., Kuyper, T.W., Wallander, H., Plassara, C., Gorbushina, A.A., Haselwandter, K., Holmstro¨m, S., Landeweert, R., Lundstro¨m, U.S., Rosling, A., Sen, R., Smits, M.M., van Hees, P.A.W., van Breemen, N., 2004. The role of fungi in weathering. Frontiers in Ecology and the Environment 2, 258–264. Hoog, G.S., Guarro, J., Gene´, J., Figueras, M.J., 2000. Clinical Fungi Atlas, second ed. University Roviri i Virgili, Reus, Spain. Mediavilla, A., Angulo, J., Infante, F. Domı´ nguez, E., 1995. Variation models of two Cladosporium conidials type (Dematiaceae, Deuteromycetes). In: Xunta de Galicia, A. (Ed.), Proceedings of the XI Simposio Nacional de Criptogamia. Santiago de Compostela, pp. 143–144. Maggi, O., Persiani, A.M., Gallo, F., Valenti, P., Pasquariello, G., Sclocchi, M.C., Scorrano, M., 2000. Airborne fungal spores in dust present in archives: proposal for a detection method, new for archival materials. Aerobiologia 16, 429–434.
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