Chapter 4 : Fungi in Polar Environments

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Although some fungi that are endemic to the extreme polar regions show psychrophilic behavior, the majority are instead psychrotolerant and globally distributed, ranging from the Arctic to Antarctica. It is important to note that although their natural ecological niches are in the polar environments, such fungi can also grow in human proximity; they can inhabit freezers and cold-storage rooms, and refrigerated and even frozen food. The methods used for fungal detection have been time appropriate, from the classical early microscope visualization to the more recent sophisticated DNA-based techniques, which have been complemented lately by metagenomic studies, although these have generally not been focused on fungi. Permafrost in polar regions covers more than 25% of the land surface and significant parts of the coastal sea shelves. Fungal diversity in the Arctic and Antarctic permafrost has been studied intensively over the last decade. The dominant species in Arctic glaciers environments was , which represented on average half of all of the isolated strains from the glaciers studied. Penicillia were the most frequently occurring filamentous fungi in all of our samples, including seawater, sea ice, snow/coastal ice in tidal zones, puddles on snow, subglacial ice, and glacial meltwater.

Citation: Zalar P, Sonjak S, Gunde-Cimerman N. 2012. Fungi in Polar Environments, p 79-94. In Miller R, Whyte L (ed), Polar Microbiology: Life in a Deep Freeze. ASM Press, Washington, DC. doi: 10.1128/9781555817183.ch4

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Microbial Ecology
Debaryomyces hansenii
Green Algae
Man-Made Environment
Epifluorescence Microscopy
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Micromorphological structures of the most common fungi in polar regions. (A) ; (B) ; (C) a sp.; (D) ; (E) a sp.

Citation: Zalar P, Sonjak S, Gunde-Cimerman N. 2012. Fungi in Polar Environments, p 79-94. In Miller R, Whyte L (ed), Polar Microbiology: Life in a Deep Freeze. ASM Press, Washington, DC. doi: 10.1128/9781555817183.ch4
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1. Abyzov, S. S., 1993. Microorganisms in the Antarctic ice, p. 265295. In E. I. Friedmann (ed.), Antarctic Microbiology. Wiley-Liss, New York, NY.
2. Abyzov, S. S.,, R. B. Hoover,, S. Imura,, I. N. Mitskevich,, T. Naganuma,, M. N. Poglazova,, and M. V. Ivanov. 2004. Use of different methods for discovery of ice-entrapped microorganisms in ancient layers of the Antarctic glacier. Adv. Space Res. 33:12221230.
3. Azmi, O. R.,, and R. D. Seppelt. 1998. The broad-scale distribution of microfungi in the Windmill Islands region, continental Antarctica. Polar Biol. 19:92100.
4. Babjeva, I.,, and I. Reshetova. 1998. Yeast resources in natural habitats at polar circle latitude. Food Technol. Biotechnol. 36:15.
5. Baublis, J. A.,, R. A. Wharton,, and P. A. Volz. 1991. Diversity of micro-fungi in an Antarctic dry valley. J. Basic Microbiol. 31:112.
6. Belzile, C.,, W. F. Vincent,, J. A. Gibson,, and P. V. Hove. 2001. Bio-optical characteristics of the snow, ice, and water column of a perennially ice-covered lake in the High Arctic. Can. J. Fish. Aquat. Sci. 58:24052418.
7. Bergero, R.,, M. Girlanda,, G. C. Varese,, D. Intili,, and A. M. Luppi. 1999. Psychrooligotrophic fungi from Arctic soils of Franz Joseph Land. Polar Biol. 21:361368.
8. Blehert, D. S.,, A. C. Hicks,, M. Behr,, C. U. Meteyer,, B. M. Berlowski-Zier,, E. L. Buckles,, J. T. H. Coleman,, S. R. Darling,, A. Gargas,, R. Niver,, J. C. Okoniewski,, R. J. Rudd,, and W. B. Stone. 2009. Bat white-nose syndrome: an emerging fungal pathogen? Science 323:227.
9. Bodiselitsch, B.,, C. Koeberl,, S. Master,, and W. U. Reimold. 2005. Estimating duration and intensity of Neoproterozoic snowball glaciations from Ir anomalies. Science 308:239242.
10. Branda, E.,, B. Turchetti,, G. Diolaiuti,, M. Pecci,, C. Smiraglia,, and P. Buzzini. 2010. Yeast and yeast-like diversity in the southernmost glacier of Europe (Calderone Glacier, Apennines, Italy). FEMS Microbiol. Ecol. 72:354369.
11. Brown, M. V.,, and J. P. Bowman. 2001. A molecular phylogenetic survey of sea-ice microbial communities (SIMCO). FEMS Microbiol. Ecol. 35:267275.
12. Burford, E. P.,, M. Fomina,, and G. M. Gadd. 2003. Fungal involvement in bioweathering and biotransformation of rocks and minerals. Mineral. Mag. 67:11271155.
13. Butinar, L.,, I. Spencer-Martins,, and N. Gunde-Cimerman. 2007. Yeasts in high Arctic glaciers: the discovery of a new habitat for eukaryotic microorganisms. Antonie van Leeuwenhoek 91:277289.
14. Butinar, L.,, T. Strmole,, and N. Gunde-Cimerman. 2011. Relative incidence of ascomycetous yeasts in Arctic coastal environments. Microbiol Ecol. 61:832843.
15. Cavicchioli, R.,, K. S. Siddiqui,, D. Andrews,, and K. R. Sowers. 2002. Low-temperature extremophiles and their applications. Curr. Opin. Biotechnol. 13:253261.
16. Chi, Z. M.,, F. Wang,, Z. Chi,, L. X. Yue,, G. L. Liu,, and T. Zhang. 2009. Bioproducts from Aureobasidium pullulans, a biotechnologically important yeast. Appl. Microbiol. Biotechnol. 82:793804.
17. Christner, B. C.,, E. Mosley-Thompson,, L. G. Thompson,, V. Zagorodnov,, K. Sandman,, and J. N. Reeve. 2000. Recovery and identification of viable bacteria immured in glacial ice. Icarus 144:479485.
18. de García, V.,, S. Brizzio,, D. Libkind,, P. Buzzini,, and M. van Broock. 2007. Biodiversity of cold-adapted yeasts from glacial meltwater rivers in Patagonia, Argentina. FEMS Microbiol. Ecol. 59:331341.
19. de García, V.,, S. Brizzio,, D. Libkind,, C. A. Rosa,, and M. van Broock. 2010a. Wickerhamomyces patagonicus sp. nov., an ascomycetous yeast species from Patagonia, Argentina. Int. J. Syst. Evol. Microbiol. 60:16931696.
20. de García, V.,, S. Brizzio,, G. Russo,, C. A. Rosa,, T. Boekhout,, B. Theelen,, D. Libkind,, and M. van Broock. 2010b. Cryptococcus spencermartinsiae sp. nov., a basidiomycetous yeast isolated from glacial waters and apple fruits. Int. J. Syst. Evol. Microbiol. 60:707711.
21. de Hoog, G. S.,, E. Göttlich,, G. Platas,, O. Genilloud,, G. Leotta,, and J. van Brummelen. 2005. Evolution, taxonomy and ecology of the genus Thelebolus in Antarctica. Stud. Mycol. 51:3373.
22. de la Torre, J. R.,, B. M. Goebel,, E. I. Friedmann,, and N. R. Pace. 2003. Microbial diversity of cryptoendolithic communities from the McMurdo Dry Valleys, Antarctica. Appl. Environ. Microbiol. 69:38583867.
23. D'Elia, T.,, R. Veerapaneni,, V. Theraisnathan,, and S. O. Rogers. 2009. Isolation of fungi from Lake Vostok accretion ice. Mycologia 101:751763.
24. de los Ríos, A.,, J. Wierzchos,, L. G. Sancho,, and C. Ascaso. 2003. Acid microenvironments in microbial biofilms of Antarctic endolithic microecosystems. Environ. Microbiol. 5:231237.
25. de los Ríos, A.,, L. G. Sancho,, M. Grube,, J. Wierzchos,, and C. Ascaso. 2005. Endolithic growth of two lecidea lichens in granite from Continental Antarctica detected by molecular and microscopy techniques. New Phytol. 165:181189.
26. Deming, J. W. 2002. Psychrophiles and polar regions. Curr. Opin. Microbiol. 5:301309.
27. De Wit, R.,, P. Dyer,, O. Genilloud,, E. Goetlich,, D. Hodgson,, G. S. de Hoog,, B. Jones,, J. Laybourn-Parry,, F. Marinelli,, E. Stackebrandt,, J. Swings,, B. J. Tindall,, W. Vyverman,, and A. Wilmotte,. 2003. Antarctic lakes—‘hot spots’ for microbial diversity and biotechnological screening, p. 228. In T. Avšič-Županc,, A. van Belkum,, C. Bruschi,, I. Chet,, J. Cole,, D. Farr,, W. Holzapfel,, R. J. Koerner,, A. Netrusov,, J.-C. Piffaretti,, E. Z. Ron,, R. Rosselló-Mora,, B. Schink,, S. Spiro,, B. J. Tindall,, and H. G. Trüper (ed.), 1st FEMS Congress of European Microbiologists, Slovenia, Ljubljana, June 29-July 3. Federation of European Microbiological Societies, Delft, The Netherlands.
28. Dmitriev, V. V.,, D. A. Gilichinsky,, R. N. Faizutdinova,, N. V. Ostroumova,, W. I. A. Golubev,, and V. I. Duda. 1997a. Yeasts in late Pleistocene-early Pleistocene Siberian permafrost. Cryosphera Zemli 1:6770. (In Russian.)
29. Dmitriev, V. V.,, D. A. Gilichinsky,, R. N. Faizutdinova,, I. N. Shershunov,, W. I. A. Golubev,, and V. I. Duda. 1997b. Occurrence of viable yeasts in 3-million-year-old permafrost in Siberia. Mikrobiologiya 66:655660. (In Russian.)
30. Domsch, K. H.,, W. Gams,, and T. H. Anderson. 1980. Compendium of Soil Fungi. Academic Press, London, United Kingdom.
31. Dugan, F.,, K. Schubert,, and U. Braun. 2004. Checklist of Cladosporium names. Schlechtendalia 11:1103.
32. Ekstrom, G.,, M. Nettles,, and G. A. Abers. 2003. Glacial earthquakes. Science 302:622624.
33. Ellis-Evans, J. C. 1985. Fungi from maritime Antarctic freshwater environments. Br. Antarct. Surv. Bull. 68:3745.
34. Fahnestock, M. 2003. Geophysics: glacial flow goes seismic. Science 302:578579.
35. Faizutdinova, R. N.,, N. E. Suzina,, V. I. Duda,, L. E. Petrovskaya,, and D. A. Gilichinsky,. 2005. Yeasts isolated from ancient permafrost, p. 118126. In J. D. Castello, and S. O. Rogers (ed.), Life in Ancient Ice. Princeton University Press, Princeton, NJ.
36. Fell, J. W.,, G. Scorzetti,, L. Connell,, and S. Craig. 2006. Biodiversity of micro-eukaryotes in Antarctic Dry Valley soils with <5% soil moisture. Soil Biol. Biochem. 38:31073119.
37. Foght, J.,, J. Aislabie,, S. Turner,, C. E. Brown,, J. Ryburn,, D. J. Saul,, and W. Lawson. 2004. Culturable bacteria in subglacial sediments and ice from two Southern Hemisphere glaciers. Microb. Ecol. 47:329340.
38. Freeman, K. R.,, A. P. Martin,, D. Karki,, R. C. Lynch,, M. S. Mitter,, A. F. Meyer,, J. E. Longcore,, D. R. Simmons,, and S. K. Schmidt. 2009. Evidence that chytrids dominate fungal communities in high-elevation soils. Proc. Natl. Acad. Sci. USA 106:1831518320.
39. Friedmann, E. I. 1982. Endolithic microorganisms in the Antarctic cold desert. Science 215:10451053.
40. Friedmann, E. I.,, and A. M. Koriem. 1989. Life on Mars: how it disappeared (if it was ever there). Adv. Space Res. 9:167172.
41. Frisvad, J. C., 2008. Fungi in cold ecosystems, p. 137156. In R. Margesin,, F. Schinner,, J.-C. Marx,, and C. Gerday (ed.), Psychrophiles: from Biodiversity to Biotechnology. Springer, Berlin, Germany.
42. Frisvad, J. C.,, T. O. Larsen,, P. W. Dalsgaard,, K. A. Seifert,, G. Louis-Seize,, E. K. Lyhne,, B. B. Jarvis,, J. C. Fettinger,, and D. P. Overy. 2006. Four psychrotolerant species with high chemical diversity consistently producing cycloaspeptide A, Penicillium jamesonlandense sp nov., Penicillium ribium sp nov., Penicillium soppii and Penicillium lanosum. Int. J. Syst. Evol. Microbiol. 56:14271437.
43. Gadd, G. M. 2007. Geomycology: biogeochemical transformations of rocks, minerals, metals and radionuclides by fungi, bioweathering and bioremediation. Mycol. Res. 111:349.
44. Gaidos, E.,, B. Lanoil,, T. Thorsteinsson,, A. Graham,, M. Skidmore,, S. K. Han,, T. Rust,, and B. Popp. 2004. A viable microbial community in a subglacial volcanic crater lake, Iceland. Astrobiology 4:327344.
45. Gilichinsky, D.,, E. Rivkina,, C. Bakermans,, V. Shcherbakova,, L. Petrovskaya,, S. Ozerskaya,, N. Ivanushkina,, G. Kochkina,, K. Laurinavichuis,, S. Pecheritsina,, R. Fattakhova,, and J. M. Tiedje. 2005. Biodiversity of cryopegs in permafrost. FEMS Microbiol. Ecol. 53:117128.
46. Gilichinsky, D. A.,, G. S. Wilson,, E. I. Friedmann,, C. P. McKay,, R. S. Sletten,, E. M. Rivkina,, T. A. Vishnivetskaya,, L. G. Erokhina,, N. E. Ivanushkina,, G. A. Kochkina,, V. A. Shcherbakova,, V. S. Soina,, E. V. Spirina,, E. A. Vorobyova,, D. G. Fyodorov-Davydov,, B. Hallet,, S. M. Ozerskaya,, V. A. Sorokovikov,, K. S. Laurinavichyus,, A. V. Shatilovich,, J. P. Chanton,, V. E. Ostroumov,, and J. M. Tiedje. 2007. Microbial populations in Antarctic permafrost: biodiversity, state, age, and implication for astrobiology. Astrobiology 7:275311.
47. Golubev, W. I. 1998. New species of basidiomycetous yeasts, Rhodotorula creatinovora and R. yakutica, isolated from permafrost soils of Eastern-Siberian Arctic. Mykol. Phytopathol. 32:813. (In Russian.)
48. Golubic, S.,, E. I. Friedmann,, and J. Schneider. 1981. The lithobiontic ecological niche, with special reference to microorganisms. J. Sediment. Res. 51:475478.
49. Gorbushina, A. A.,, and W. J. Broughton. 2009. Microbiology of the atmosphere-rock interface: how biological interactions and physical stresses modulate a sophisticated microbial ecosystem. Annu. Rev. Microbiol. 63:431450.
50. Gosink, J. J.,, R. L. Irgens,, and J. T. Staley. 1993. Vertical distribution of bacteria in Arctic sea ice. FEMS Microbiol. Lett. 102:8590.
51. Gostinčar, C.,, M. Grube,, S. de Hoog,, P. Zalar,, and N. Gunde-Cimerman. 2010. Extremotolerance in fungi: evolution on the edge. FEMS Microbiol. Ecol. 71:211.
52. Grabińska-Łoniewska, A.,, T. Koniłłowicz-Kowalska,, G. Wardzyńska,, and K. Boryn. 2007. Occurrence of fungi in water distribution system. Pol. J. Environ. Stud. 16:539547.
53. Gunde-Cimerman, N.,, L. Butinar,, S. Sonjak,, M. Turk,, V. Uršič,, P. Zalar,, and A. Plemenitaš,. 2005. Halotolerant and halophilic fungi from coastal environments in the Arctics, p. 397423. In N. Gunde-Cimerman,, A. Orenand,, and A. Plemenitaš (ed.), Adaptation to Life at High Salt Concentrations in Archaea, Bacteria, and Eukarya. Springer, Dordrecht, The Netherlands.
54. Gunde-Cimerman, N.,, S. Sonjak,, P. Zalar,, J. C. Frisvad,, B. Diderichsen,, and A. Plemenitaš. 2003. Extremophilic fungi in arctic ice: a relationship between adaptation to low temperature and water activity. Phys. Chem. Earth B 28:12731278.
55. Gunde-Cimerman, N.,, P. Zalar,, S. de Hoog,, and A. Plemenitaš. 2000. Hypersaline waters in salterns—natural ecological niches for halophilic black yeasts. FEMS Microbiol. Ecol. 32:235240.
56. Hölker, U.,, J. Bend,, R. Pracht,, L. Tetsch,, T. Müller,, M. Höfer,, and G. S. de Hoog. 2004. Hortaea acidophila, a new acid-tolerant black yeast from lignite. Antonie van Leeuwenhoek 86:287294.
57. Hughes, K. A.,, and B. Lawley. 2003. A novel Antarctic microbial endolithic community within gypsum crusts. Environ. Microbiol. 5:555565.
58. Ivanushkina, N. E.,, G. A. Kochkina,, and S. M. Ozerskaya,. 2005. Fungi in ancient permafrost sediments of the Arctic and Antarctic regions, p. 127139. In J. D. Castello, and S. O. Rogers (ed.), Life in Ancient Ice. Princeton University Press, Princeton, NJ.
59. Jones, G. E. B. 1976. Recent Advances in Aquatic Mycology. The Gresham Press, Old Woking, Surrey, United Kingdom.
60. Junge, K.,, F. Imhoff,, T. Staley,, and W. Deming. 2002. Phylogenetic diversity of numerically important Arctic sea-ice bacteria cultured at subzero temperature. Microb. Ecol. 43:315328.
61. Kennedy, J.,, B. Flemer,, S. A. Jackson,, D. P. H. Lejon,, J. P. Morrissey,, F. O’Gara,, and A. D. W. Dobson. 2010. Marine metagenomics: new tools for the study and exploitation of marine microbial metabolism. Mar. Drugs 8:608628.
62. Kriss, A. E.,, I. N. Mitskevich,, E. P. Rozanova,, and L. K. Osnitskaia. 1976. Microbiological studies of the Wanda Lake (Antarctica). Mikrobiologiya 45:10751081. (In Russian.)
63. Kurek, E.,, T. Korniłłowicz-Kowalska,, A. Słomka,, and A. J. Melke. 2007. Characteristics of soil filamentous fungi communities isolated from various micro-relief forms in the high Arctic tundra (Bellsund region, Spitsbergen). Pol. Polar Res. 28:5773.
64. Libkind, D.,, S. Brizzio,, A. Ruffini,, M. Gadanho,, M. van Broock,, and J. P. Sampaio. 2003. Molecular characterization of carotenogenic yeasts from aquatic environments in Patagonia, Argentina. Antonie van Leeuwenhoek 84:313322.
65. Libkind, D.,, M. Gadanho,, M. van Broock,, and J. P. Sampaio. 2005. Sporidiobolus longiusculus sp. nov. and Sporobolomyces patagonicus sp. nov., novel yeasts of the Sporidiobolales isolated from aquatic environments in Patagonia, Argentina. Int. J. Syst. Evol. Microbiol. 55:503509.
66. Libkind, D.,, M. Moline,, J. P. Sampaio,, and M. van Broock. 2009. Yeasts from high-altitude lakes: influence of UV radiation. FEMS Microbiol. Ecol. 69:353362.
67. López-García, P.,, F. Rodríguez-Valera,, C. Pedrós-Alió,, and D. Moreira. 2001. Unexpected diversity of small eukaryotes in deep-sea Antarctic plankton. Nature 409:603607.
68. Loque, C. P.,, A. O. Medeiros,, F. M. Pellizzari,, E. C. Oliveira,, C. A. Rosa,, and L. H. Rosa. 2010. Fungal community associated with marine macroalgae from Antarctica. Polar Biol. 33:641648.
69. Ludley, K. E.,, and C. H. Robinson. 2008. “Decomposer” Basidiomycota in Arctic and Antarctic ecosystems. Soil Biol. Biochem. 40:1129.
70. Lydolph, M. C.,, J. Jacobsen,, P. Arctander,, M. T. Gilbert,, D. A. Gilichinsky,, A. J. Hansen,, E. Willerslev,, and L. Lange. 2005. Beringian paleoecology inferred from permafrost-preserved fungal DNA. Appl. Environ. Microbiol. 71:10121017.
71. Ma, L. J.,, C. M. Catranis,, W. T. Starmer,, and S. O. Rogers. 1999. Revival and characterization of fungi from ancient polar ice. Mycologist 13:7073.
72. Ma, L. J.,, S. O. Rogers,, C. M. Catranis,, and W. T. Starmer. 2000. Detection and characterization of ancient fungi entrapped in glacial ice. Mycologia 92:286295.
73. Margesin, R. 2009. Effect of temperature on growth parameters of psychrophilic bacteria and yeasts. Extremophiles 13:257262.
74. Margesin, R.,, and J. W. Fell. 2008. Mrakiella cryoconiti gen. nov., sp. nov., a psychrophilic, anamorphic, basidiomycetous yeast from alpine and arctic habitats. Int. J. Syst. Evol. Microbiol. 58:29772982.
75. Margesin, R.,, P. A. Fonteyne,, F. Schinner,, and J. P. Sampaio. 2007. Rhodotorula psychrophila sp. nov., Rhodotorula psychrophenolica sp. nov. and Rhodotorula glacialis sp. nov., novel psychrophilic basidiomycetous yeast species isolated from alpine environments. Int. J. Syst. Evol. Microbiol. 57:21792184.
76. Margesin, R.,, G. Zacke,, and F. Schinner. 2002. Characterization of heterotrophic microorganisms in alpine glacier cryoconite. Arct. Antarct. Alp. Res. 34:8893.
77. McLoughlin, N.,, M. D. Brasier,, D. Wacey,, O. R. Green,, and R. S. Perry. 2007. On biogenicity criteria for endolithic microborings on early Earth and beyond. Astrobiology 7:1026.
78. McRae, C. F.,, A. D. Hocking,, and R. D. Seppelt. 1999. Penicillium species from terrestrial habitats in the Windmill Islands, East Antarctica, including a new species, Penicillium antarcticum. Polar Biol. 21:97111.
79. Montes, M. J.,, C. Belloch,, M. Galiana,, M. D. Garcia,, C. Andrés,, S. Ferrer,, J. M. Torres-Rodriguez,, and J. Guinea. 1999. Polyphasic taxonomy of a novel yeast isolated from Antarctic environment; description of Cryptococcus victoriae sp. nov. Syst. Appl. Microbiol. 22:97105.
80. Morozova, D.,, D. Möhlmann,, and D. Wagner. 2007. Survival of methanogenic archaea from Siberian permafrost under simulated Martian thermal conditions. Orig. Life Evol. Biosph. 37:189200.
81. Mueller, D. R.,, W. F. Vincent,, W. H. Pollard,, and C. H. Fritsen. 2001. Glacial cryoconite ecosystems: a bipolar comparison of algal communities and habitats. Nova Hedwig. Beih. 123:173197.
82. Nagano, Y.,, T. Nagahama,, Y. Hatada,, T. Nunoura,, H. Takami,, J. Miyazaki,, K. Takai,, and K. Horikoshi. 2010. Fungal diversity in deep-sea sediments—the presence of novel fungal groups. Fungal Ecol. 3:316325.
83. Nienow, J. A.,, and E. I. Friedmann,. 1993. Terrestrial lithophytic (rock) communities, p. 343412. In E. I. Friedmann (ed.), Antarctic Microbiology. Wiley-Liss, New York, NY.
84. Omelon, C. R.,, W. H. Pollard,, and F. G. Ferris. 2006. Environmental controls on microbial colonization of high Arctic cryptoendolithic habitats. Polar Biol. 30:1929.
85. Omelon, C. R.,, W. H. Pollard,, and F. G. Ferris. 2007. Inorganic species distribution and microbial diversity within high Arctic cryptoendolithic habitats. Microb. Ecol. 54:740752.
86. Onofri, S.,, D. Barreca,, L. Selbmann,, D. Isola,, E. Rabbow,, G. Horneck,, J. P. P. de Vera,, J. Hatton,, and L. Zucconi. 2008. Resistance of Antarctic black fungi and cryptoendolithic communities to simulated space and Martian conditions. Stud. Mycol. 61:99109.
87. Onofri, S.,, M. Fenice,, A. R. Cicalini,, S. Tosi,, A. Magrino,, S. Pagano,, L. Selbmann,, L. Zucconi,, H. S. Vishniac,, R. Ocampo-Friedmann,, and E. I. Friedmann. 2000. Ecology and biology of microfungi from Antarctic rocks and soils. Ital. J. Zool. 67:163167.
88. Onofri, S.,, S. Pagano,, L. Zucconi,, and S. Tosi. 1999. Friedmanniomyces endolithicus (Fungi, Hyphomycetes), anam.-gen. and sp. nov., from continental Antarctica. Nova Hedwig. Beih. 68:175181.
89. Onofri, S.,, L. Selbmann,, L. Zucconi,, and S. Pagano. 2004. Antarctic microfungi as models for exobiology. Planet. Space Sci. 52:229237.
90. Ozerskaya, S.,, G. Kochkina,, N. Ivanushkina,, and D. A. Gilichinsky,. 2009. Fungi in permafrost, p. 8595. In R. Margesin (ed.), Permafrost Soils. Springer, Berlin, Germany.
91. Ozerskaya, S. M.,, G. A. Kochkina,, N. E. Ivanushkina,, E. V. Knyazeva,, and D. A. Gilichinskii. 2008. The structure of micromycete complexes in permafrost and cryopegs of the Arctic. Microbiology 77:482489.
92. Panikov, N. S., 2009. Microbial activity in frozen soils, p. 119147. In R. Margesin (ed.), Permafrost Soils. Springer, Berlin, Germany.
93. Pathan, A.,, B. Bhadra,, Z. Begum,, and S. Shivaji. 2010. Diversity of yeasts from puddles in the vicinity of Midre Lovénbreen Glacier, Arctic and bioprospecting for enzymes and fatty acids. Curr. Microbiol. 60:307314.
94. Pennisi, E. 2003. Neither cold nor snow stops tundra fungi. Science 301:1307.
95. Pitt, J. I.,, and A. D. Hocking. 1999. Fungi and Food Spoilage. Aspen Publishers, Gaithersburg, Maryland.
96. Pitt, J. I.,, R. A. Samson,, and J. C. Frisvad,. 2000. List of accepted species and their synonyms in the family Trichocomaceae, p. 947. In R. A. Samson, and J. I. Pitt (ed.), Integration of Modern Taxonomic Methods for Penicillium and Aspergillus Classification. Harwood Academic Publishers, Amsterdam, The Netherlands.
97. Poglazova, M. N.,, I. N. Mitskevich,, S. S. Abyzov,, and M. V. Ivanov. 2001. Microbiological characterization of the accreted ice of subglacial Lake Vostok, Antarctica. Microbiology 70:723730.
98. Price, P. B. 2000. A habitat for psychrophiles in deep Antarctic ice. Proc. Natl. Acad. Sci. USA 97:12471251.
99. Price, P. B.,, and T. Sowers. 2004. Temperature dependence of metabolic rates for microbial growth, maintenance, and survival. Proc. Natl. Acad. Sci. USA 101:46314636.
100. Priscu, J. C.,, C. H. Fritsen,, E. E. Adams,, S. J. Giovannoni,, H. W. Paerl,, C. P. McKay,, P. T. Doran,, D. A. Gordon,, B. D. Lanoil,, and J. L. Pinckney. 1998. Perennial Antarctic lake ice: an oasis for life in a polar desert. Science 280:20952098.
101. Reeve, J. N.,, B. C. Christner,, B. H. Kvitko,, E. Mosley-Thompson,, and L. G. Thompson,. 2002. Life in glacial ice, p. 27. In M. Rossi,, S. Bartolucci,, M. Ciaramellaand,, and M. Moracci (ed.), Extremophiles 2002, The Fourth International Congress on Extremophiles, Naples, Italy. Institute of Protein Biochemistry and University of Naples “Federico II,” Naples, Italy.
102. Rivkina, E. M.,, E. I. Friedmann,, C. P. McKay,, and D. A. Gilichinsky. 2000. Metabolic activity of permafrost bacteria below the freezing point. Appl. Environ. Microbiol. 66:32303233.
103. Rivkina, E.,, K. Laurinavichius,, J. McGrath,, J. Tiedje,, V. Shcherbakova,, and D. Gilichinsky. 2004. Microbial life in permafrost. Adv. Space Res. 33:12151221.
104. Robinson, C. H. 2001. Cold adaptation in Arctic and Antarctic fungi. New Phytol. 151:341353.
105. Rohde, R. A.,, and P. B. Price. 2007. Diffusion-controlled metabolism for long-term survival of single isolated microorganisms trapped within ice crystals. Proc. Natl. Acad. Sci. USA 104:1659216597.
106. Ruibal, C.,, C. Gueidan,, L. Selbmann,, A. A. Gorbushina,, P. W. Crous,, J. Z. Groenewald,, L. Muggia,, M. Grube,, D. Isola,, C. L. Schoch,, J. T. Staley,, F. Lutzoni,, and G. S. de Hoog. 2009. Phylogeny of rock-inhabiting fungi related to Dothideomycetes. Stud. Mycol. 64:123133.
107. Ruisi, S.,, D. Barreca,, L. Selbmann,, L. Zucconi,, and S. Onofri. 2007. Fungi in Antarctica. Rev. Environ. Sci. Biotechnol. 6:127141.
108. Russell, N. J.,, P. Harrisson,, I. A. Johnston,, R. Jaenicke,, M. Zuber,, F. Franks,, and D. Wynn-Williams. 1990. Cold adaptation of microorganisms. Philos. Trans. R. Soc. London B 326:595611.
109. Samson, R. A.,, E. S. Hoekstra,, J. C. Frisvad,, and O. Filtenborg. 2002. Introduction to Food- and Airborne Fungi. Centralbureau voor Schimmelcultures, Utrecht, The Netherlands.
110. Schadt, C. W.,, A. P. Martin,, D. A. Lipson,, and S. K. Schmidt. 2003. Seasonal dynamics of previously unknown fungal lineages in tundra soils. Science 301:13591361.
111. Schmidt, N.,, and M. Bölter. 2002. Fungal and bacterial biomass in tundra soils along an arctic transect from Taimyr Peninsula, central Siberia. Polar Biol. 25:871877.
112. Schubert, K.,, J. Z. Groenewald,, U. Braun,, J. Dijksterhuis,, M. Starink,, C. F. Hill,, P. Zalar,, G. S. de Hoog,, and P. W. Crous. 2007. Biodiversity in the Cladosporium herbarum complex (Davidiellaceae, Capnodiales), with standardisation of methods for Cladosporium taxonomy and diagnostics. Stud. Mycol. 58:105156.
113. Scorzetti, G.,, I. Petrescu,, D. Yarrow,, and J. W. Fell. 2000. Cryptococcus adeliensis sp. nov., a xylanase producing basidiomycetous yeast from Antarctica. Antonie van Leeuwenhoek 77:153157.
114. Selbmann, L.,, G. S. de Hoog,, A. Mazzaglia,, E. I. Friedmann,, and S. Onofri. 2005. Fungi at the edge of life: cryptoendolithic black fungi from Antarctic desert. Stud. Mycol. 51:132.
115. Selbmann, L.,, G. S. de Hoog,, L. Zucconi,, D. Isola,, S. Ruisi,, A. H. van den Ende,, C. Ruibal,, F. De Leo,, C. Urzi,, and S. Onofri. 2008. Drought meets acid: three new genera in a dothidealean clade of extremotolerant fungi. Stud. Mycol. 61:120.
116. Siegert, M. J.,, J. C. Ellis-Evans,, M. Tranter,, C. Mayer,, J.-R. Petit,, A. Salamatin,, and J. C. Priscu. 2001. Physical, chemical and biological processes in Lake Vostok and other Antarctic subglacial lakes. Nature 414:603609.
117. Siegert, M. J.,, M. Tranter,, J. C. Ellis-Evans,, J. C. Priscu,, and W. Berry Lyons. 2003. The hydrochemistry of Lake Vostok and the potential for life in Antarctic subglacial lakes. Hydrol. Processes 17:795814.
118. Simon, C.,, A. Wiezer,, A. W. Strittmatter,, and R. Daniel. 2009. Phylogenetic diversity and metabolic potential revealed in a glacier ice metagenome. Appl. Environ. Microbiol. 75:75197526.
119. Skidmore, M. L.,, J. M. Foght,, and M. J. Sharp. 2000. Microbial life beneath a high Arctic glacier. Appl. Environ. Microbiol. 66:32143220.
120. Sonjak, S.,, J. C. Frisvad,, and N. Gunde-Cimerman. 2005. Comparison of secondary metabolite production by Penicillium crustosum strains, isolated from Arctic and other various ecological niches. FEMS Microbiol. Ecol. 53:5160.
121. Sonjak, S.,, J. C. Frisvad,, and N. Gunde-Cimerman. 2006. Penicillium mycobiota in Arctic subglacial ice. Microb. Ecol. 52:207216.
122. Sonjak, S.,, J. C. Frisvad,, and N. Gunde-Cimerman. 2007a. Genetic variation among Penicillium crustosum isolates from Arctic and other ecological niches. Microb. Ecol. 54:298305.
123. Sonjak, S.,, V. Uršič,, J. C. Frisvad,, and N. Gunde-Cimerman. 2007b. Penicillium svalbardense, a new species from Arctic glacial ice. Antonie van Leeuwenhoek 92:4351.
124. Stakhov, V.,, S. Gubin,, S. Maksimovich,, D. Rebrikov,, A. Savilova,, G. Kochkina,, S. Ozerskaya,, N. Ivanushkina,, and E. Vorobyova. 2008. Microbial communities of ancient seeds derived from permanently frozen Pleistocene deposits. Microbiology 77:348355.
125. Starmer, W.,, J. Fell,, C. Catranis,, V. Aberdeen,, L. Ma,, S. Zhou,, and S. Rogers,. 2005. Yeasts in the genus Rhodotorula recovered from the Greenland ice sheet, p. 181195. In J. D. Castello, and S. O. Rogers (ed.), Life in Ancient Ice. Princeton University Press, Princeton, NJ.
126. Sterflinger, K.,, G. S. de Hoog,, and G. Haase. 1999. Phylogeny and ecology of meristematic ascomycetes. Stud. Mycol. 43:522.
127. Sterflinger, K.,, and W. E. Krumbein. 1997. Dematiaceous fungi as a major agent for biopitting on Mediterranean marbles and limestones. Geomicrobiol. J. 14:219230.
128. Steven, B.,, R. Leveille,, W. H. Pollard,, and L. G. Whyte. 2006. Microbial ecology and biodiversity in permafrost. Extremophiles 10:259267.
129. Takano, Y.,, K. Kobayashi,, K. Marumo,, and Y. Ishikawa,. 2004. Biochemical indicators and enzymatic activity below permafrost environment, p. 84. In F. Robb,, M. W. Adams,, K. Horikoshi,, R. M. Kelly,, J. Littlechild,, K. E. Nelson,, J. Reeve,, R. Roberts,, K. R. Sowers,, and K. Stetter (ed.), Extremophiles 2004, 5th International Conference on Extremophiles, September 19-23, Cambridge, Maryland. American Society for Microbiology, Washington, DC.
130. Thomas, D. N.,, and G. S. Dieckmann. 2002. Antarctic sea ice—a habitat for extremophiles. Science 295:641644.
131. Thomas-Hall, S. R.,, B. Turchetti,, P. Buzzini,, E. Branda,, T. Boekhout,, B. Theelen,, and K. Watson. 2010. Cold-adapted yeasts from Antarctica and the Italian Alps—description of three novel species: Mrakia robertii sp. nov., Mrakia blollopis sp. nov. and Mrakiella niccombsii sp. nov. Extremophiles 14:4759.
132. Thomas-Hall, S.,, and K. Watson. 2002. Cryptococcus nyarrowii sp. nov., a basidiomycetous yeast from Antarctica. Int. J. Syst. Evol. Microbiol. 52:10331038.
133. Thomas-Hall, S.,, K. Watson,, and G. Scorzetti. 2002. Cryptococcus statzelliae sp. nov. and three novel strains of Cryptococcus victoriae, yeasts isolated from Antarctic soils. Int. J. Syst. Evol. Microbiol. 52:23032308.
134. Tosi, S.,, B. Casado,, R. Gerdol,, and G. Caretta. 2002. Fungi isolated from Antarctic mosses. Polar Biol. 25:262268.
135. Tosi, S.,, S. Onofri,, M. Brusoni,, L. Zucconi,, and H. Vishniac. 2005. Response of Antarctic soil fungal assemblages to experimental warming and reduction of UV radiation. Polar Biol. 28:470482.
136. Turchetti, B.,, P. Buzzini,, M. Goretti,, E. Branda,, G. Diolaiuti,, C. D'Agata,, C. Smiraglia,, and A. Vaughan-Martini. 2008. Psychrophilic yeasts in glacial environments of Alpine glaciers. FEMS Microbiol. Ecol. 63:7383.
137. Vincent, W. F. 2000. Evolutionary origins of Antarctic microbiota: invasion, selection and endemism. Antarct. Sci. 12:374385.
138. Vincent, W. F.,, D. R. Mueller,, and S. Bonilla. 2004. Ecosystems on ice: the microbial ecology of Markham Ice Shelf in the high Arctic. Cryobiology 48:103112.
139. Vishniac, H. S., 1993. The microbiology of Antarctic soils, p. 297342. In E. I. Friedmann (ed.), Antarctic Microbiology. Wiley-Liss, New York, NY.
140. Vishniac, H. S. 2002. Cryptococcus tephrensis, sp. nov., and Cryptococcus heimaeyensis, sp. nov.; new anamorphic basidiomycetous yeast species from Iceland. Can. J. Microbiol. 48:463467.
141. Vishniac, H. S., 2006. Yeast biodiversity in the Antarctic, p. 419440. In C. A. Rosaand, and G. Péter (ed.), Biodiversity and Ecophysiology of Yeasts. Springer, Berlin, Germany.
142. Vishniac, H. S.,, and S. Onofri. 2003. Cryptococcus antarcticus var. circumpolaris var. nov., a basidiomycetous yeast from Antarctica. Antonie van Leeuwenhoek 83:231233.
143. Vorobyova, E.,, V. Soina,, M. Gorlenko,, N. Minkovskaya,, N. Zalinova,, A. Mamukelashvili,, D. Gilichinsky,, E. Rivkina,, and T. Vishnivetskaya. 1997. The deep cold biosphere: facts and hypothesis. FEMS Microbiol. Rev. 20:277290.
144. Wagner, D., 2008. Microbial communities and processes in Arctic permafrost environments, p. 133154. In P. Dionand, and C. S. Nautiyal (ed.), Microbiology of Extreme Soils. Springer, Berlin, Germany.
145. Wagner, D.,, A. Lipski,, A. Embacher,, and A. Gattinger. 2005. Methane fluxes in permafrost habitats of the Lena Delta: effects of microbial community structure and organic matter quality. Environ. Microbiol. 7:15821592.
146. Wallenstein, M. D.,, S. McMahon,, and J. Schimel. 2007. Bacterial and fungal community structure in Arctic tundra tussock and shrub soils. FEMS Microbiol. Ecol. 59:428435.
147. Wicklow, D.,, and D. Malloch. 1971. Studies in the genus Thelebolus: temperature optima for growth and ascocarp development. Mycologia 63:118131.
148. Xiao, N.,, K. Suzuki,, Y. Nishimiya,, H. Kondo,, A. Miura,, S. Tsuda,, and T. Hoshino. 2010. Comparison of functional properties of two fungal antifreeze proteins from Antarctomyces psychrotrophicus and Typhula ishikariensis. FEBS J. 277:394403.
149. Xin, M. X.,, and P. J. Zhou. 2007. Mrakia psychrophila sp. nov., a new species isolated from Antarctic soil. J. Zhejiang Univ. Sci. B 8:260265.
150. Zalar, P.,, G. S. de Hoog,, H.-J. Schroers,, P. W. Crous,, J. Z. Groenewald,, and N. Gunde-Cimerman. 2007. Phylogeny and ecology of the ubiquitous saprobe Cladosporium sphaerospermum, with descriptions of seven new species from hypersaline environments. Stud. Mycol. 58:157183.
151. Zalar, P.,, C. Gostinčar,, G. S. de Hoog,, V. Uršič,, M. Sudhadham,, and N. Gunde-Cimerman. 2008. Redefinition of Aureobasidium pullulans and its varieties. Stud. Mycol. 61:2138.

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