Science of the Total Environment 493 (2014) 116–123

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Science of the Total Environment

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Diversity of an aerial phototrophic coating of historic buildings in theformer Auschwitz II-Birkenau concentration camp

Paulina Nowicka-Krawczyk a,⁎, Joanna Żelazna-Wieczorek a,1, Anna Otlewska b,2, Anna Koziróg b,2,Katarzyna Rajkowska b,2, Małgorzata Piotrowska b,2, Beata Gutarowska b,2, Agnieszka Żydzik-Białek c,3

a Department of Algology and Mycology, Faculty of Biology and Environmental Protection, University of Łódź, Banacha Str. 12/16, 90-237, Łódź, Polandb Institute of Fermentation Technology and Microbiology, Łódź University of Technology, Wólczańska Str. 171/173, 90-924 Łódź, Polandc Auschwitz-Birkenau State Museum in Oświęcim, Więźniów Oświęcimia Str. 20, 32-603 Oświęcim, Poland

H I G H L I G H T S

• We analysed diversity of phototrophs causing biodeterioration of the camp buildings.• We indicated preferences of phototrophs in colonizing varied substrate types.• We predicted the environmental factor that determines the growth of phototrophs.• We indicated the pioneer phototrophs in the prevailing climatic conditions.

⁎ Corresponding author. Tel.: +48 426354111.E-mail addresses: paulina_nowicka84@interia.pl (P. N

zelazna@biol.uni.lodz.pl (J. Żelazna-Wieczorek), anna.otleanna.kozirog@p.lodz.pl (A. Koziróg), katarzyna.rajkowskamalgorzata.piotrowska@p.lodz.pl (M. Piotrowska), beata.g(B. Gutarowska), agnieszka.bialek@auschwitz.org (A. Żyd

1 Tel.: +48 426354111.2 Tel.: +48 426313470.3 Tel.: +48 338448176.

http://dx.doi.org/10.1016/j.scitotenv.2014.05.1130048-9697/© 2014 Elsevier B.V. All rights reserved.

a b s t r a c t

a r t i c l e i n f o

Article history:Received 9 March 2014Received in revised form 8 May 2014Accepted 25 May 2014Available online 14 June 2014

Editor: Charlotte Poschenrieder

Keywords:Aerial algaeBiodeteriorationCyanobacteriaDiversitySeasonal changesHistoric buildings

Aerial phototrophs colonize materials of anthropogenic origin, thus contributing to their biodeterioration. Struc-tures preserved at the former Auschwitz II-Birkenau concentration and extermination camp show signs of deg-radation by cyanobacteria and algae. In order to protect the Auschwitz-BirkenauMemorial Site, diversity of aerialphototrophs growing on the historic buildings has been studied. Analyses of cyanobacterial and algal biofilmsgrowing on various construction substrates were carried out in summer and winter. Multivariate data analyseswere used to: characterize the diversity of cyanobacteria and algae growing in brick andwooden camp buildingsdepending on the research season, indicate preferences of cyanobacteria and algae in colonizing substrates, andto predict the environmental factor that most determines the growth of phototrophs. The biofilms were formedmainly by cyanobacteria, green algae and diatoms. The amount of cyanobacteria and algae in the biofilms wasvaried, which resulted from changes in climatic conditions, the type of substrate and the height at which thebiofilms developed. In the summer, the ratio of cyanobacteria and algae groupswas balanced,while in thewinter,green algae and diatomswere dominant. Green algae showed a preference for colonizing plaster, wood and con-crete, of which the walls and doors of the buildings were made. Their participation was correlated with a heightgradient. Cyanobacteria and diatoms grew on bricks and soil on the floor of the buildings and temperature andrelative humidity were the factors that modified their amount. Green algae were more cosmopolitan—occurredin dry places, potentially inaccessible to other organisms; therefore, they have been identified as the pioneergroup in the prevailing climatic conditions.

© 2014 Elsevier B.V. All rights reserved.

owicka-Krawczyk),wska@p.lodz.pl (A. Otlewska),@p.lodz.pl (K. Rajkowska),utarowska@p.lodz.plzik-Białek).

1. Introduction

Cyanobacteria and algae include a group of organisms characterizedby a specific ecology. These are aerial phototrophs, which have devel-oped various adaptation mechanisms, yielding the ability to colonizeand grow in the terrestrial environment. They can modify the propor-tion of pigments in cells, which protects the photosynthetic apparatusagainst excessive UV radiation and produce envelope membranes thatprotect cells against water loss (Johansen, 2001; Żelazna-Wieczorek,2011). These organisms grow on tree trunks, bedrock, soil (Kaweckaand Eloranta, 1994; Samad and Adhikary, 2008) as well as all kinds of

117P. Nowicka-Krawczyk et al. / Science of the Total Environment 493 (2014) 116–123

artificial substrates produced by human activities (Rindi and Guiry,2004). Aerial cyanobacteria and algae communities develop in all cli-mate zones (Gaylarde and Gaylarde, 2000; Samad and Adhikary,2008). In the temperate zone, these communities are largely formedby unicellular and filamentous forms of green algae (Chlorophyta)(Rindi andGuiry, 2004; Samad and Adhikary, 2008), while in thewarm-er, temperate to tropical climate, by blue–green algae (Cyanobacteria)that produce various kinds of mucilaginous envelopes protecting cellsfrom excessive drying (Crispim et al., 2004; Samad andAdhikary, 2008).

Man does not play a significant role in the spread of terrestrial algae.The cells of algae and their spores are carried by the wind (Brown et al.,1964; Barberousse et al., 2007). After reaching the substrate in goodcondition, they divide and grow to form various kinds of biofilms. Basi-cally, the type of substrate does not play a decisive role in the develop-ment of natural strains, because cyanobacteria and algae are so welladapted that the growth biofilms is possible on wood, brick or concretesubstrates. There are, however, trends in colonization,which are relatedto the physical properties of materials such as roughness and porosity(Barberousse et al., 2007). The decisive role in the growth and develop-ment of cyanobacteria and algae is played by appropriate light condi-tions, temperature and humidity, which are associated largely with adistance from larger aquatic ecosystems and vegetation (Barberousseet al., 2006). Access to mineral compounds and adequate substrate pHare also important (Grbić et al., 2010). Seasonal modifications in thestructure of phototrophic communities growing on substrates are dueto seasonal variation in temperature and humidity (Ress and Lowe,2013).

Terrestrial cyanobacteria and algae are pioneer organisms, which col-onize habitats potentially unavailable for living organisms and transformthem, allowing other groups of organisms to colonize there (Grahamet al., 2009). Formation of the phototrophic biocenosis allows the growthof more complex communities, which are then formed from heterotro-phic microbiota (Tomaselli et al., 2000). Phototrophs inhabiting anthro-pogenic substrates thereby contribute to their rapid biodeterioration(Tomaselli et al., 2000; Crispim and Gaylarde, 2004; Samad andAdhikary, 2008). They produce photosynthetic pigments, which changethe colour of the substrates on which the cyanobacteria and algaegrow. This adversely affects the aesthetic value of buildings and culturalmonuments (Grbić et al., 2010; Hauer, 2010; Stupar et al., 2012). Whenhumidity changes, the hydration and volume of algal cells are also mod-ified, causing structural microdamages to substrates (Gorbushina, 2007;Hauer, 2010). Many phototrophs are capable of dissolving compoundscontained in a substrate and penetrating into it, causing mechanical ero-sion (Brehmet al., 2005; Crispim andGaylarde, 2004; Hauer, 2010). Dur-ing themetabolic activity of the algal cells, various types of inorganic andorganic acids are produced and algae secrete them into the external en-vironment, causing chemical deterioration of substrates (Gaylarde andMorton, 1999; Samad and Adhikary, 2008; Stupar et al., 2012).

For the protection of buildings of historic interest, it is crucial toidentify the organisms that cause biodeterioration of materials andlearn their autecology. Buildings in the former Auschwitz II-Birkenauconcentration and extermination camp in Brzezinka the site of theNazi German concentration and extermination camp Auschwitz wereinscribed on the UNESCO World Heritage List in 1979. It is one of themost important symbols of the atrocities of World War II. One of thegoals of the Auschwitz-Birkenau Preservation Plan is to protect thestructures of the Auschwitz-Birkenau Memorial Site against biodeterio-ration, because signs of biological degradation caused by bacteria, fungiand algae have been observed on them. The results of research on mi-croorganisms that cause biodegradation of historic structures areshown in Rajkowska et al. (2013) and Koziróg et al. (2014), whereasthis work presents the results of the study on the diversity of terrestrialcyanobacteria and algae.

The aim of this paper is to characterize the taxonomic diversity of ae-rial phototrophs on the historic buildings at the former Auschwitz II-Birkenau concentration and extermination camp. The scope of the

research includes the qualitative and quantitative analyses of photo-trophic communities growing on various substrates in summer andwin-ter, an analysis of the dominance of phototrophic groups depending onthe season, examination of preferences in colonizing different types ofsubstrates and characterization of the effect of temperature and relativehumidity on the growth of cyanobacteria and algae. The paper also pre-sents a research hypothesis: is it possible to specify a pioneer groupamong the identified phototrophs in the given climatic conditions?

2. Material and methods

2.1. Study area

The former Auschwitz II-Birkenau concentration and exterminationcamp inOświęcim covers an area of about 171 hectares. Currently, thereare 76 brick buildings and 22 wooden buildings (Fig. 1). The area of theformer camp, where the historic housing buildings are situated, is de-void of trees and the structures are exposed to constant light radiation.The camp is surrounded by forest from the north-western side, while awatercourse runs on its south-western side. During heavy downpours,the water level rises in the watercourse, soil moisture goes up and thebuildings located on the eastern side of the camp are partially flooded.

The study was conducted in ten buildings: eight brick barracks (inv.no 66, 70, 110, 113, 114, 124, 125 and 138) and two wooden barracks(inv. no 159 and 169). The barracks were in different technical condi-tions. Some were more damaged and damp (70, 113, 114, 124, 125,138), while others were characterized by a better state and lowermois-ture content (66, 110, 159, 169) (Table 1). In placeswhere the visual as-sessment showed the development of phototrophic biofilms (Fig. 2),environmental parameters were measured: air temperature and rela-tive humidity were measured using the thermohygrometer ElmetronPWT-401 and samples were taken for phycological analysis. Sampleswere collected from locations inside and outside the buildings: floors,walls, foundations of buildings and doors, from various heights (from0 to 200 cm). Biofilms were taken frommineral substrates (bricks, con-crete and plaster) and organic substrates (indoor soil and wood). Sam-ples of moss thalli were also collected from two locations aroundbuilding 124, where algae grow in the macroscopic visible form, inorder to check whether these sites are the source from which thealgae get onto the building substrates.

2.2. Phycological analysis

Phycological material was collected during the summer (06/2012)and winter (12/2012). Biofilms of approximately 9 cm2 were removedfrom substrates with a hard brush and placed in 4% aqueous solutionof formaldehyde, in order to fix them. The moss surface was wipedwith a soft brush and then the thallus was shaken in distilled water.The resulting precipitate was placed as above in the solution of formal-dehyde. Samples, where diatoms were detected, were subjected toetching in a mixture of sulphuric VI and chromic VII acids (Singh et al.,2006; Żelazna-Wieczorek, 2011) to obtain silica cell walls, whose ap-pearance is a diagnostic feature of diatoms.

The qualitative analysis of samples was conducted on the basis ofmorphological features of the observed taxa to the genus or specieslevel, followed by Ettl (1978, 1983), Krammer and Lange-Bertalot(1986, 1988, 1991a,b), Ettl and Gärtner (1988), Komárek andAnagnostidis (1989, 1999, 2007), Lange-Bertalot (2001), Samad andAdhikary (2008), Hofmann et al. (2011), and Komárek (2013).Cyanobacteria and algae were observed with a Nikon YS 100 light mi-croscope under magnification of 400× and 1000×.

The amount of individual taxa in samples was estimated. For thispurpose, the frequencies at which different taxa occurred in 30 consec-utive fields of microscopic view under a magnification of 200×, in trip-licates, were observed (Kawecka and Eloranta, 1994).

Fig. 1. The former camp of Auschwitz II-Birkenau, (fot. Paweł Sawicki) [1 column fitting].

118 P. Nowicka-Krawczyk et al. / Science of the Total Environment 493 (2014) 116–123

2.3. Multivariate data analysis

Phototrophic communities and the effect of habitat conditions onthe structure of the communities were characterized by multivariateanalyses using the software CANOCO for Windows 5.0. The analyseddata were different in nature—linear and unimodal. Therefore the anal-ysis was carried out using four different techniques. A principal compo-nent analysis (PCA) and a redundancy analysis (RDA) were used forlinear data, while a detrended correspondence analysis (DCA) and a ca-nonical correspondence analysis (CCA) were used for unimodal data(Lepš and Šmilauer, 2003). The PCA and DCA techniques were used toinvestigate the diversity of samples based on the qualitative and quan-titative analyses of algae and to show the dissimilarity of communitiesstructure in samples depending on the research season. The CCA tech-nique was applied to show preferences in settling on different types ofsubstrates, while the RDA was used to indicate the impact of environ-mental factors on the growth of algae.

3. Results

A total of 51 cyanobacterial and algal taxa were identified in thesamples, belonging to four taxonomic groups: blue–green algae(Cyanobacteria), diatoms (Bacillariophyta), chrysophytes (Chrysophyta)and green algae (Chlorophyta) (Table 2). Among the identifiedphototrophs, 13 taxa were connected only with bryophytes developingaroundbarracks and their presencewasnot recorded in samples collectedfrom construction substrates. Due to the high specificity of the communi-ty associatedwith bryophytes, the sampleswere excluded frommultivar-iate analyses.

It was observed that one or two taxawere dominant in the amount ofbiofilm-forming cyanobacteria and algae. The communities were domi-nated by: cyanobacteria of the genera Chroococcus, Leptolyngbya andScytonema drilosiphon; diatoms Diadesmis contenta and Achnanthidiumminutissimum and green algae of the genera Chlorella and Apatococcuslobatus. In the PCA analysis, the vertical ordination axis differentiated be-tween samples frombarracks in a better technical condition and samplestaken from barracks in a poor state of repair (Fig. 3a–b). Algal biofilmsgrowing on substrates in the first type of buildings were mostly formedby single-celled green algae:A. lobatus, Chlorococcum infusionum,Chlorel-la vulgaris, Chlorella sp. and Coenochloris sp. (Fig. 3a: I). The more

damaged buildings were coated by the biofilms of cyanobacteria and di-atoms (Fig. 3a: II). On the brick floor of brick bath buildings, 70, 113, 114,and 124, the thallus of S. drilosiphon strongly developed during the sum-mer period and covered locallymore than 70% of the substrate surface. Inthe winter, this thallus completely disappeared. The horizontal ordina-tion axis (Fig. 3b) subsequently identified samples from the winter peri-od. The structures of phototrophic communities emerging in the winter(W) were less diversified compared to samples from the summer (S)(Fig. 3b).

There was a seasonal change in the amount of biofilm-forming taxa.In the summer, the amount of blue–green algae, diatoms and greenalgaewas relatively similar (Fig. 4a). However, a significant quantitativeadvantage of single-cell green algae and diatoms was observed in sam-ples collected in the winter. Cyanobacteria occurred less frequently inthat period (Fig. 4b).

The biofilms developed on the studied mineral substrates (bricks,concrete and plaster) and organic substrates (wood and soil inside thebuildings). The CCA analysis made it possible to observe preferencesof particular systematic groups of phototrophs in colonizing differentsubstrates (Fig. 5). The vertical ordination axes divided substrates intotwo groups, depending on their location. The first group, including sub-strates such as concrete, plaster and wood, was located vertically in thebuildings and was mostly inhabited by green algae. The second group,located horizontally in the buildings, included brick and soil substratesand was mostly inhabited by cyanobacteria and diatoms (Fig. 5).Among the green algae, Trentepohlia sp., Trebouxia sp., Coenochloris sp.,C. infusionum, A. lobatus and green algae of the genus Chlorella werecharacteristic of concrete, plaster and brick. The analysis also identifieda group exclusively associatedwith soil samples: Chroococcus sp.,Nostocsp. and cyanobacteria of the genus Leptolyngbya (Fig. 5). Horizontal or-dination axes divided substrates intomineral and organic ones, depend-ing on their origin. However, no colonization patterns relating to thiscriterion were observed.

Samples from concrete, plaster and wood were taken from differentheights, while those from bricks and soil were collected only from thefloors of the buildings. The RDA analysis showed that the height atwhich biofilm develops affects the development and amount of greenalgae (Fig. 6). The height gradient correlated most strongly with thequantitative participation of A. lobatus in biofilms. Air temperature in-fluences the development of green algae to a lesser extent. Air humidityfor the species of the genus Chlorella is a variable that is negatively

Table 1Technical state of historic buildings and characteristic of the microhabitats of the sampling sites.

Inv. no. Technical state of buildings Substrate for phycol.analysis

No. of samples Temperature [°C] Air microhumidity [%]

Summer Winter Summer Winter Summer Winter

B-66 The building surrounded by soil and debris, damagedconcrete floor inside the building, soil exposedbetween concrete cracks, no signs of damp woodenfloor in the barrack master's room and warehouse,damaged plaster on the walls.

Plaster, soil;wooda

31a

31a

21 – 2325a

−5 – −4−5a

41 – 5157a

74 – 7768a

B-70 The building surrounded by soil and rubble, brickfloor inside the building, cracked in some places, soilexposed where the floor is bulging, locally damp,damaged plaster on the walls.

Wood, soil 3 3 22 – 23 −4 – −3 44 – 47 70 – 75

B-110 The building surrounded by a band of concrete, theconcrete floor without damage, no signs of humidity,slightly damaged plaster on the walls.

Concrete, plaster 3 3 22 – 23 −4 – −3 42 – 43 63 – 66

B-113 The building is surrounded by soil and debris,damaged and damp wooden floor in the barrackmaster's room and warehouse, soil exposed betweenconcrete cracks, damaged plaster on the walls.

Brick, soil 5 5 20 – 21 −6 – −5 51 – 64 72 – 73

B-114 The building surrounded by a damaged band ofconcrete, damaged and damp wooden floor in thebarrack master's room and warehouse, soil exposedbetween concrete cracks, damaged plaster on thewalls.

Brick, soil 5 5 21 – 23 −5 – −4 51 – 55 71 – 72

B-124 The building surrounded by soil and debris, damagedfloor inside the building and the wooden floor in thebarrackmaster's room, visible signs of high humidity,soil exposed between concrete cracks, damagedplaster on the walls.

Plaster, brick, soilbriophytesa

82a

82a

18 – 2124a

−6 – −5−5a

47 – 6158a

78 – 7970a

B-125 The building surrounded by soil and rubble, damagedwooden floor inside the building, visible signs of highhumidity, undamaged concrete floor, damagedplaster on the walls.

Plaster, brick 2 2 20 – 21 −4 – −3 48 – 52 61 – 63

B-138 The building surrounded by a damaged band ofconcrete, damaged wooden floor in the barrackmaster's room and warehouse, visiblesigns of high humidity, damaged concrete floorinside, soil exposed between concrete cracks,damaged plaster on the walls.

Wood, brick 3 3 20 – 22 −6 – −5 49 – 51 55 – 68

B-159 The building surrounded by a damaged band ofconcrete, damaged brick and concretefloor inside thebuilding, locally damp wooden walls.

Brick, concretewooda, concretea

42a

42a

22 – 2325a

−4 – −3−5a

40 – 4157a

69 – 7468a

B-169 The building surrounded by a band of stones, brickfloor inside the building, locally soil, locally dampwooden walls.

Concrete, brick, soilwooda, bricka

42a

42a

22 – 2325a

−4 – −3−5a

46 – 4757a

67 – 7168a

a Details for samples taken outside the buildings.

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correlated with their quantitative participation. In the case of diatomsand cyanobacteria, temperature is a variable which differentiates thestructure of the communities more strongly than air humidity (Fig. 6).

4. Summary and discussion

Thirty-eight taxa of cyanobacteria and algae growing on the historicbuildings in the former Auschwitz II-Birkenau concentration and exter-mination camp were observed during the study. Aerial phototrophsformed biofilms on different types of building substrates. Given therelatively small size of the study area, such a taxonomic diversity ofphototrophs appears to be large. In Europe, Rindi and Guiry (2004) iden-tified only 17 taxa on concretewalls and the ground adjacent to thewalls,while in France, Barberousse et al. (2006) found47 taxa ondifferent typesof organic and mineral substrate. The study at the former Auschwitz II-Birkenau concentration camp confirms that types of substrates are im-portant for measuring the taxonomic diversity of phototrophs.

The study of aerial phototrophs indicates that the representativesof blue–green algae (Cyanobacteria) and green algae (Chlorophyta)are the main components of communities growing on substrates inthe terrestrial environment (Davey, 1988; Tomaselli et al., 2000;Gärtner and Stoyneva, 2003; Barberousse et al., 2007; Stupar et al.,2012). Diatoms (Bacillariophyta) (Ress and Lowe, 2013), chryso-phytes (Chrysophta), euglenoids (Euglenophyta) and dinoflagellates

(Dinophyta) (Khaybullina et al., 2010) are less numerous and they ac-company the communities. The presence of euglenophytes anddinophytes was not recorded in the study. Xanthonema was the onlygenus found of chrysophytes and it occurred in a very small numberof samples collected exclusively from thewooden doors of the barracks.At the same time, there were many blue–green algae, diatoms andgreen algae. The proportion of their participation in the communitiesvaried depending on the technical condition of the barracks and wassubject to seasonal change. In the summer, their participation remainedin relative equilibrium. Differences occurred in the case of individualsamples from the barracks in a good state of repair which were betterventilated and less damp than the barracks in a poor technical condi-tion, which were stuffy and the floor bore the signs of prolonged mois-ture. A. lobatus, C. vulgaris, Chlorella sp., C. infusionum, Coenochloris sp.and Trentepohlia cf. odoratawere dominant in the first type of barracks.Due to a more favourable microclimate for the development ofcyanobacteria and algae (smaller range of temperatures and higher hu-midity), cyanobacterial biofilms and thalli, with a dominance ofS. drilosiphon and taxa of the genera Leptolyngbya and Chroococcus, de-veloped on the brick floor and soil in the second type of buildings. Inthe winter, the quantitative participation of cyanobacteria decreasedsignificantly at low temperature and only representatives of the genusChroococcus were most often recorded. In the winter, the unicellulargreen algae significantly increased their amount in the communities.

Fig. 2. Biofilms developing on the buildingmaterials in the former Auschwitz II-Birkenau concentration camp. a= biofilm on the brick floor in a brick barrack; b= biofilm on the soil in abrick barrack; c = biofilm on a patch in a brick barrack; d = biofilm on the concrete and wood of a wooden barrack [each part 1 column fitting; all 2 columns fitting].

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Diatoms were recorded in samples throughout the study period and agreater diversity of species was observed under the conditions of highhumidity, which, as noted by Ress and Lowe (2013), is due to theirlower adaptive capacity to over-drying. Casamatta et al. (2002) statethat a lack of diatoms in dry places is a common phenomenon, but inthe study these algae commonly occurred in dry places, but in theform of almost single-species cultures of D. contenta. This species is typ-ical for terrestrial habitats. D. contenta are able to successfully developeven after little and short-term contact with water (Hofmann et al.,

Table 2Cyanobacteria and algae identified inside (I) and outside (O) the buildings and on bryophytes

Taxa Code I76a

O10a

B4

CyanobacteriaChroococcopsis cf. fluviatilis Chpflu 28 – –

Chroococcus minor Chrmin 41 3 –

Chroococcus sp. Chrcsp 1 2 –

Chroococcus varius Chrvar 44 3 –

Cyanobium cf. parvum Cyafpv – – 1Gloeocapsa sp. Glcasp 5 1 –

Gloeothece palea Gltpal 11 1 –

Gloeothece sp. Gltcsp 13 3 –

Leptolyngbya foveolarum Lepfov 14 – –

Leptolyngbya notata Lepnot 4 – –

Pseudanabaena frigida (=Leptolyngbya frigida) Leptsp 11 2 –

Microcoleus vaginatus Mclvag – – 2Nostoc commune Noscom – – 2Nostoc microscopicum Nocmic – – 2Nostoc punctiforme Nospun – – 1Nostoc sp. Nostsp 1 – –

Oscillatoria cf. princeps Oscfpr – – 1Phormidium aerugineo-caeruleum Phoaec 4 1 –

Phormidium breve Phobre 2 2 –

Phormidium tergestinum Photer 4 – –

Scytonema drilosiphon Scydri 14 – –

Tolypothrix sp. Tolysp 8 1 –

BacillariophytaAchnanthidium minutissimum Achmin 51 4 –

Anomoeoneis vitrea Anovit 9 – –

Cocconeis placentula var. lineata Cocpll – – 1Cocconeis pseudothumensis Cocpst 3 4 –

a The total number of samples.

2011). Their presence in extremely dry habitats on the studied struc-tures and in highly humid places, on the border of the water and air en-vironment, where they were often observed, suggests a wide range oftolerance to humidity conditions, as confirmed by Veselá andJohansen (2009). Cyanobacteria S. drilosiphon growing on the bricks ofbath barracks clearly preferred warmer and humid microclimate.When the temperature dropped below 20°C, the cyanobacteria startedto die, whichwas accompanied by discolouration of the thallus from in-tense turquoise to pale yellow. This species is identified in terrestrial

(B) around the studied structures—the number of samples with the presence of taxa.

aTaxa Code I

76aO10a

B4a

Bacillariophyta [cn.]Denticula kuetzingii Denkue 5 – –

Diadesmis contenta Diacon 74 9 –

Epithemia turgida var. granulata Epitug – – 1Gomphonema parvulum Gompar 19 2 –

Luticola mutica Lutmut 4 – –

Navicula cryptocephala Navcpt – – 2Neidium ampliatum Neiamp 6 1 –

Nitzschia debilis Ntzdeb 4 – –

Nitzschia fonticola Ntzfon – – 2Nitzschia palea Ntzpal – – 2Nitzschia vitrea Ntzvit 9 – –

Pinnularia borealis Pinbor – – 1Planothidium lanceolatum Planlan 2 3 –

Trybionella hungarica Tryhun 14 1 –

ChrysophytaXanthonema montanum Xanmon 1 – –

Xanthonema sp. Xantsp 6 – –

ChlorophytaApatococcus lobatus Apalob 41 10 –

Chlorella vulgaris Chlvul 41 6 –

Chlorella sp. Chlosp 48 9 –

Chlorococcum infusionum Chlinf 1 5 –

Coenochloris sp. Coensp 8 8 –

Euastrum sp. Euassp 4 2 –

Phacotus sp. Phacsp – – 1Trebouxia sp. Trebsp 1 – –

Trentepohlia cf. odorata Trensp 1 1 –

Fig. 3. The PCA ordination plot based on the qualitative and quantitative analyses ofphototrophs in all collected samples. a= taxa frombuildings in a good (I) and abad (II) tech-nical condition; b = differentiation between samples collected in winter (W) and summer(S): marked winter samples [each 1 column fitting, all 2 columns fitting].

Fig. 4. The DCA ordination plot displaying participation of cyanobacteria and algae groups(in squares) in collected samples. a = in the summer season; b = in the winter season[each 1 column fitting; all 2 columns fitting].

Fig. 5. The CCA ordination biplot displaying preferences of phototrophs to substrate types[1.5 columns fitting].

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ecosystems, where it often develops on bricks. It is observed mainly intropical climates, but at elevated temperature and humidity; it also oc-curs in temperate climates (Starmach, 1966; Guiry, 2013; Komárek,2013).

Green algae (Chlorophyta) can be considered a characteristic andpermanent group of aerophytic algae that occurs on the historic build-ings in the former Auschwitz II-Birkenau concentration camp. Their tax-onomic diversity is much smaller than that of cyanobacteria anddiatoms; however, they occur more frequently than other phototrophsand their quantities are almost equal both in summer and winter.A. lobatus, which occurred very often and in large numbers, markedlyincreased their amount in the communities located above the ground,whichwas associatedwith a decrease in humidity. It is a species charac-teristic of terrestrial algal communities and it often develops on mate-rials of anthropogenic origin. The occurrence of this species appears tobe independent of the type of substrate, because it can be found bothon a mineral substrate – granite, limestone, concrete (Tomaselli et al.,2000; Gärtner and Stoyneva, 2003; Barberousse et al., 2006) – and onan organic substrate such as wood.

Sunlight, humidity and temperature are the most significant factorsin the development of natural phototrophic biofilms on building mate-rials (Barberousse et al., 2006; Sethi et al., 2012). In the area of the for-mer Auschwitz II-Birkenau concentration camp, cyanobacteria andalgae appeared inside the barracks only in areas exposed to sunlight.Biofilms occurred both in dry and humid places and at temperaturesfrom above 25 °C in the summer to about −6 °C in the winter. Thesefactors did not affect the phenomenon of phototrophic colonization ofsubstrates. Temperature and humidity were important, but from thepoint of view of particular groups of phototrophs. Khaybullina et al.(2010) also stress that a substrate pH is an important colonization

parameter, as it mainly affects the qualitative composition ofcyanobacteria and algae.

Cyanobacterial and algal preference for substrates identified inthe studywas related to their location. Green algae inhabited verticalsubstrates – walls made of wood, concrete or plaster – whilecyanobacteria preferred horizontal substrates—floors made of bricks

Fig. 6. The RDA ordination biplot displaying the influence of environmental variables onthe presence of phototrophs [1 column fitting].

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or soil. Barberousse et al. (2007) indicate that porosity and roughnessare the most important physical characteristics of materials in termsof the rate of photorophic colonization on substrates, because they in-crease the ability forwater absorption and retention, which then causesan increase in microhumidity. However, differences in microhumiditydo not affect the weight of cyanobacteria and algae growing on struc-tures, but modify their taxonomic composition, since the occurrenceof certain species such as Leptolyngbya foveolarum can have a direct re-lationship with increased microhumidity (Barberousse et al., 2006). Inthe study, air humidity represented an environmental factor, which af-fected the growth of phototrophs to a lesser extent than temperature.During the study, relative humidity was a parameter that did notchange between microhabitats as significantly as temperature rangingup to 5 °C.

On the basis of the observations, it can be confirmed that single-celled green algae are better adapted to lower humidity conditions.They were found in large quantities in both research seasons and theyare the first to appear in the better preserved buildings. Only an increasein humidity affects colonization of substrates by cyanobacteria, whosedevelopment is recorded primarily on ground substrates. Can it thus beconcluded that green algae are the pioneer group for the study area?The study in different latitudes of Europe by Rindi and Guiry (2004) sug-gests that this group of organisms is themost widespread and shows pi-oneer tendencies in colonizing building structures. It is green algae thatbegin the succession in substrates in unfavourable conditions aftersnow melting (Davey, 1988; Barberousse et al., 2006). The successionof aerial phototrophic groups is different for warm and tropical climates,where cyanobacteria form themain biomass in communities and are thefirst to colonize an inaccessible environment (Mulec et al., 2008; Ressand Lowe, 2013). It can therefore be concluded that under the conditionsof higher temperature and humidity, cyanobacteria show pioneer ten-dencies in settling in microhabitats, while green algae are the pioneergroup in temperate and cold climate as they are adapted to drierconditions.

The pioneer green algae indicated in the study are a biological degra-dative factor, which primarily affects the aesthetic qualities of buildings.The activity of green algae is less degradative than that of cyanobacteriaas the latter change the hydration of cells and their membranes, pene-trate the surfaces of substrates (even if they are coated with a layer ofprotective paint) and thus damage them (Hauer, 2010). For this reason,no study has been conducted so far to identify the role and examine thedegradative activity of green algae.

In further studies of aerial phototrophs growing on the buildings inthe former Auschwitz II-Birkenau concentration camp, investigation isplanned into the effect of different compounds on the development ofbiofilms. The study will allow for the selection of appropriate methodsto control and prevent degradation caused by autotrophic organismsin order to protect the monuments of the Auschwitz-Birkenau StateMuseum in Oświęcim.

Acknowledgements

This work was conducted as a part of the Auschwitz-Birkenau Pres-ervation Plan and supported by the Auschwitz-Birkenau Foundationunder the grant number 5/2012/GPK.

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