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Complete Issue J. Mulec and G. Kosi – Lampenflora algae and methods of growth control. Journal of Cave and Karst Studies, v. 71, no. 2, p. 109–115. LAMPENFLORA ALGAE AND METHODS OF GROWTH CONTROL JANEZ MULEC1 AND GORAZD KOSI2 Abstract: Karst caves are unique natural features and habitats where specialized organisms live. Some caves are also important as cultural heritage sites. In recent decades, many caves have experienced intensified tourist visits. To attract visitors, artificial illumination was installed that changed conditions in the caves. As a result, communities of organisms called lampenflora develop in close and remote proximity to lights. These phototrophic organisms are inappropriate from an aesthetic point of view and cause the degradation of colonized substrata, which is a particular problem in caves with prehistoric art. Key factors that allow lampenflora to grow are light and moisture. Illuminated spots in caves can be quickly colonized by algae, some of which have broad tolerances for different substrata. Several phototrophs can survive in caves even at photon flux densities lower than the photosynthetic compensation point. In this paper, the pros and cons of physical, chemical, and biological methods to control phototrophic growth are reviewed and discussed. Experiences in show caves can be helpfulin controlling undesirable algal growth in other environments. INTRODUCTION show no oscillations in light intensity. Dobat (1972) named spots with growing lampenflora ecosystems in formation. Caves have a special place in human history. Early in One of the characteristics of the natural cave environ- prehistory, humans discovered that caves can provide ment is low nutrient input (Simon et al., 2007) that is suitable temporary or permanent shelters. Later, man changed with the introduction of light energy. Such drastic developed a different relation with caves, not only as changes to the cave ecosystem directly and indirectly shelter but also for their natural beauty and inspiration. In influence cave fauna. Higher nutrient input in cave many caves around the globe, remnants of prehistoric man environments enables newcomers to be more competitive are found. Especially interesting are those caves with than the originally present troglomorphic organisms. paintings. Many caves of natural and cultural importance Consequently obligate cave-dwelling organisms are threat- are listed on the United Nations Educational, Scientific, ened and may become extinct without restoration of and Cultural Organization (UNESCO) World Heritage previous natural conditions (Pipan, 2005). List. Cave tourism is considered to be one of the oldest In the last few years, many different views about forms of tourism. unwanted phototrophs in caves have appeared, but the In recent decades, many caves have experienced main question was not what these green cave dwellers are, intensified tourist visits. To attract visitors, artificial but how to prevent their growth (Planina 1974; Ash et al., illumination was installed. Illuminated areas such as rocky 1975; Caumartin, 1977; Caumartin, 1986; Iliopoulou- surfaces, sediments, and artificial materials around lamps Georgoudaki et al., 1993; Gurnee, 1994; Byoung-woo, quickly become colonized by phototrophic organisms. This 2002; Hazslinszky, 2002; Lochner, 2002; Olson, 2002; complex community of autotrophic photosynthetic organ- Merdenisianos, 2005). An important problem occurs when isms is called lampenflora and develops in natural and lampenflora becomes covered with CaCO3, irrespective of artificial caves around artificial light sources (Dobat, whether this carbonate is a result of abiotic or biotic 1998). In this lampenflora community, various aerophytic precipitation. Such an amorphous mix of dead photo- algae, as well as some mosses and ferns dominate, and are trophs and CaCO3 irreversibly destroys the natural usually strongly adhered to the substratum. Mosses and heritage of speleothems or other objects of cultural value ferns, also part of lampenflora, are not discussed further (Mulec, 2005). Loss of historic paintings and objects in because in the early phase of colonization and succession, caves due to biological activities is becoming an important algae, both prokaryotic cyanobacteria and eukaryotic problem. The purpose of this paper is to review various algae, usually play the most important role, while mosses methods to control lampenflora growth and to select the and ferns appear later in the succession. Vascular plants are most appropriate one. sometimes found, but almost always as germinating shoots (Martincˇicˇ et al., 1981). Lampenflora is, relative to the 1 Karst Research Institute, Scientific Research Centre of the Slovenian Academy aerophytic phototrophs from the cave entrances, complete- of Sciences and Arts, Titov trg 2, SI-6230 Postojna, Slovenia, janez.mulec@ guest.arnes.si ly independent of sunlight and other external climatic 2 National Institute of Biology, Vecˇna pot 111, SI-1000 Ljubljana, Slovenia, factors. In comparison with sunlight, artificial light sources [email protected] Journal of Cave and Karst Studies, August 2009 N 109 LAMPENFLORA ALGAE AND METHODS OF GROWTH CONTROL ALGAE OF THE LAMPENFLORA COMMUNITY stage of species succession in the lampenflora community, cyanobacteria become much more abundant, and thus, Biodiversity of lampenflora is low compared to the flora community composition becomes more similar to the from cave entrances where cyanobacteria are dominant community from a cave entrance where cyanobacteria (Palik, 1964; Golubic´, 1967; Buczko´ and Rajczy, 1989; dominate (Mulec et al., 2008). Vinogradova et al., 1995; Vinogradova et al., 1998; Asencio In another experiment, limestone disks placed around and Aboal, 2000a, b; Uher and Kova´cˇik, 2002; Mulec et lamps in show caves were colonized by the following al., 2008). Mulec et al. (2008) demonstrated that the phototrophs: Apatococcus lobatus, Chlorella sp., Lyngbya concentration of chlorophyll a per unit surface area of sp., Nostoc sp., Navicula mutica, and Trentepohlia aurea. lampenflora algae was slightly higher (max. 2.44 mgcm22) Similar composition of the algal community was observed compared to the epilithic algae from the cave entrance when limestone disks were covered with Jaworski agar (max. 1.71 mgcm22). This difference can be explained due (Mulec, 2005), a culture medium commonly used in to the different light regimes in both microhabitats (i.e., algology (Warren et al., 1997). Organisms identified on changing light quality and irradiance levels during the day disks covered with agar were: Apatococcus cf. lobatus, in the cave entrance), different periods of illumination, Chlorella sp., Lyngbya sp., Stichococcus bacillaris, and different in situ moisture levels, and different species Trentepohlia aurea. This later experiment confirmed that composition. Generally speaking, areas around lamps have eukaryotic microalgae successfully and quickly colonize more stable conditions because they are deeper in the new stony surfaces around lamps. Fast-growing photo- constant zone of the cave (Mulec et al., 2008). trophs can quickly re-colonize former niches, even with a As reported by Garbacki et al. (1999) in Belgian caves, high frequency of cleaning of rocky surfaces with biocides. Cyanobacteria made up 54% of the phototrophic organ- Microscopic observations of scrapes from disk surfaces isms, Chrysophyta 30%, and Chlorophyta 16%. Dobat showed that disks covered with agar were colonized to a (1970) and Chang and Chang-Schneider (1991) recorded greater extent by bacteria and fungi when compared to the the percentages of phototrophs in German caves; Cyano- disks without agar covering (Mulec, 2005). Nevertheless, bacteria 55%, Chrysophyta 14%, and Chlorophyta 31%. lampenflora algae are usually ubiquitous, fast reproducing, Cyanobacteria also prevailed in Slovenian caves with 51%, and adaptable soil algae (Rajczy, 1989). Existence of algae followed by Chrysophyta 27%, and Chlorophyta 22% and other higher plants (i.e., ferns and mosses deep in show (Dobat, 1973; Martincˇicˇ et al., 1981; Krivograd-Klemencˇicˇ caves) demonstrates effective transport of viable propa- and Vrhovsˇek, 2005; Mulec, 2005; Mulec et al., 2008). It gules from outside the caves. was shown that cyanobacteria, Aphanothece castagnei and Gloeocapsa sanguinea, and green alga, Stichococcus bacil- KEY ENVIRONMENTAL FACTORS THAT INFLUENCE laris, were part of lampenflora in all three countries. Other LAMPENFLORA GROWTH species that appeared with high frequency included Aphanocapsa grevillei, A. muscicola, Chondrocystis dermo- Three modes of transport of viable propagules into the chroa, Chroococcus varius, C. westii, Gloeocapsa atrata, G. karst underground can be distinguished: air currents, water compacta, G. rupestris, Gloeothece rupestris, Leptolyngbya flow, and introduction by animals and humans. (Dobat, lurida, L. perelegans, Phormidium retzii, Schizothrix calci- 1970; Vegh, 1989). Conditions deep in the cave are usually cola, Scytonema julianum (Cyanobacteria), Achnanthes sp., very stable, but that can be changed when high amounts of A. lanceolata, Elipsoidon oocystoides, Navicula contenta, energy are introduced. An important factor in the Pinnularia borealis (Chrysophyta), Chlorella miniata, spreading of lampenflora and other organisms is local air Gloeocystis polydermatica, Klebshormidium flaccidum, currents caused by warm air in the proximity of strong Pleurococus vulgaris, and Trentepohlia aurea (Chloro- lamps, especially halogen ones (500–1000
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