190:(China),, also in cold deserts as Arctic and Antarctic, and deep subsoil and ocean trenches rocks. However, there are reports of endolithic microorganisms in inter-tropical zones, where humidity and solar radiation are significantly different from the above-mentioned biomes. Endoliths have been found in the rock down to a depth of 3 kilometers (1.9 mi), though it is unknown if that is their limit (due to the cost involved in drilling to such depths). The main threat to their survival seems not to result from the pressure at such depth, but from the increased temperature. Judging from
109:, living in places long considered inhospitable to life. The distribution, biomass, and diversity of endolith microorganisms are determined by the physical and chemical properties of the rock substrate, including the mineral composition, permeability, the presence of organic compounds, the structure and distribution of pores, water retention capacity, and the pH. Normally, the endoliths colonize the areas within lithic substrates to withstand intense solar radiation, temperature fluctuations, wind, and desiccation. They are of particular interest to
508:
under high water stress and oligotrophic conditions confer to endolithic microorganisms to survive in conditions similar to those found on Mars. There is evidence of the past existence of water on the red planet; perhaps, these microorganisms could develop adaptations found in current deserts on the Earth. Furthermore, The endolithic structures are a good way to find ancient or current biological activity (
412:, studies have suggested that the endolithic algae located in the skeleton of the coral may be aiding the survival of coral species by providing an alternative source of energy. Although the role that endolithic fungi play is important in coral reefs, it is often overlooked because much research is focused on the effects of coral bleaching as well as the relationships between
38:
507:
An excellent example of these adaptations is the non-hygroscopic but microporous translucent gypsum crusts, which are found as potential substrates that can mitigate exposure to UV radiation and desiccation and allow microbial colonization in hyper-arid deserts. In the same way, the ability to grow
475:
Fungus is seldom fossilized and even when it is preserved it can be difficult to distinguish endolithic hyphae from endolithic cyanobacteria and algae. Endolithic microbes can, however, be distinguished based on their distribution, ecology, and morphology. According to a 2008 study, the endolithic
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are still at an early stage of exploration. In some cases its biota can support simple invertebrates, most organisms are unicellular. Near-surface layers of rock may contain blue-green algae but most energy comes from chemical synthesis of minerals. The limited supply of energy limits the rates of
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As water and nutrients are rather sparse in the environment of the endolith, water limitation is a key factor in the capacity of survival of many endolithic microorganisms, many of those microorganisms have adaptations to survive in low concentrations of water. Besides, the presence of pigments,
189:
Endolithic microorganisms have been reported in many areas around the globe. There are reports in warm hyper-arid and arid deserts such as Mojave and Sonora (USA), Atacama (Chile), Gobi (China, Mongolia), Negev (Israel), Namib (Namibia Angola), Al-Jafr basin (Jordan) and the
Depression of Turpan
464:. The infestation of phototrophic endoliths resulted in lethal and sub-lethal effects such as the decrease in strength of the mussel shells. Although the rate of thickening of the shells were faster in more infested areas it is not rapid enough to combat the degradation of the mussel shells.
768:
Wierzchos, J.; Camara, B.; De Los Rios, A.; Davila, A. F.; Sanchaz Almazo, M.; Artieda, O.; Wierzchos, K.; Gomez-Silva, B.; McKay, C.; Ascaso, C. (2011). "Microbial colonization of Ca-sulfate crusts in the hyperarid core of the
Atacama Desert: Implications for the search for life on Mars".
504:, microorganisms find protection against thermal buffering, UV radiation, and desiccation while living inside pores and fissures of minerals and rocks. Life in these endolithic habitats might face similar stress due to the scarcity of water and high UV radiation that rule on modern Mars.
476:
fungi that formed on the eggshells would have resulted in the abnormal incubation of the eggs and may have killed the embryos in infected eggs of these dinosaurs. It may also have led to the preservation of dinosaur eggs, including some that contained embryos.
254:. In addition, whether they metabolize these directly from the surrounding rock, or rather excrete an acid to dissolve them first is yet undetermined. According to Meslier & DiRuggiero there are found genes in the endolithic community involved in
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Gaylarde, C.; Baptista-Neto, J. A.; Ogawa, A.; Kowalski, M.; Celikkol-Aydin, S.; Beech, I. (2017). "Epilithic and endolithic microorganisms and deterioration on stone church facades subject to urban pollution in a sub-tropical climate".
2370:
Warren-Rhodes, K. A.; Rhodes, K. L.; Pointing, S. B.; Ewing, S. A.; Lacap, D. C.; Gomez-Silva, B.; McKay, C. P. (2006). "Hypolithic cyanobacteria, dry limit of photosynthesis, and microbial ecology in the hyperarid
Atacama Desert".
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once every hundred years. In August 2013 researchers reported evidence of endoliths in the ocean floor, perhaps millions of years old and reproducing only once every 10,000 years. Most of their energy is spent repairing
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Schlesinger, W. H; Pippen, J. S.; Wallenstein, M. D.; Hofmockel, K. S.; Klepeis, D. M.; Mahall, B. E. (2003). "Community composition and photosynthesis by photoautotrophs under quartz pebbles, southern Mojave Desert".
2032:
Osterrothová, K; Culka, A; Němečková, K; Kaftan, D; Nedbalová, L; Procházková, L; Jehlička, J (2019). "Analyzing carotenoids of snow algae by Raman microspectroscopy and high-performance liquid chromatography".
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Stomeo, F; Valverde, A; Pointing, S. B.; McKay, C. P.; Warren-Rhodes, K. A.; Tuffin, M. I.; Cowan, D. A. (2013). "Hypolithic and soil microbial community assembly along an aridity gradient in the Namib Desert".
565:
Cockell, C. S.; Olsson, K.; Knowles, F.; Kelly, L.; Herrera, A.; Thorsteinsson, T.; Marteinsson, V. (2009). "Bacteria in weathered basaltic glass, Iceland. Bacteria in weathered basaltic glass, Iceland".
1124:
VĂtek, P.; Ascaso, C; Artieda, O; Wierzchos, J (2016). "Raman imaging in geomicrobiology: endolithic phototrophic microorganisms in gypsum from the extreme sun irradiation area in the
Atacama Desert".
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Endolithic fungi have been discovered in shells as early as the year 1889 by
Edouard Bornet and Charles Flahault. These two French phycologists specifically provided descriptions for two fungi:
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Inagaki, F.; Takai, K.; Komatsu, T.; Sakihama, Y.; Inoue, A.; Horikoshi, K. (2015). "Profile of microbial community structure and presence of endolithic microorganisms inside a deep-sea rock".
405:, has also been made in the beach sand of Australia by George Zembrowski. Findings have also been made in coral reefs and have been found to be, at times, beneficial to their coral hosts.
644:
Kelly, L. C.; Cockell, C. S.; Herrera-Belaroussi, A.; Piceno, Y.; Andersen, G.; DeSantis, T.; LeRoux, X. (2011). "Bacterial diversity of terrestrial crystalline volcanic rocks, Iceland".
339:, they can generate organic compounds essential for their survival on their own from inorganic matter. Some endoliths have specialized in feeding on their autotroph relatives. The micro-
914:
Bungartz, F; Garvie, L. A.; Nash, T. H. (2004). "Anatomy of the endolithic
Sonoran Desert lichen Verrucaria rubrocincta Breuss: implications for biodeterioration and biomineralization".
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Evidence of endolithic fungi were discovered within dinosaur eggshell found in central China. They were characterized as being “needle-like, ribbon-like, and silk-like.".
3124:
345:
3129:
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Meslier, V; DiRuggiero, J (2019). "7 Endolithic microbial communities as model systems for ecology and astrobiology". In
Seckbach, J.; Rampelotto, P.H. (eds.).
2538:— Compiled for professionals and advanced learners, this endolith collection includes online resources such as journal articles, academic reviews, and surveys.
2254:
GĂĽnther, Astrid (1990). "Distribution and bathymetric zonation of shell-boring endoliths in recent reef and shelf environments: Cozumel, Yucatan (Mexico)".
687:
Omelon, C. R.; Pollard, W. H.; Ferris, F. G. (2007). "Inorganic species distribution and microbial diversity within high Arctic cryptoendolithic habitats".
601:
Herrera, A.; Cockell, C. S.; Self, S.; Blaxter, M.; Reitner, J.; Thorsteinsson, T.; Tindle, A. G. (2009). "A cryptoendolithic community in volcanic glass".
3150:
3119:
2532:— This collection of online resources such as news articles, web sites, and reference pages provides a comprehensive array of information about endoliths.
1979:
Wierzchos, Jacek; Ascaso, Carmen; McKay, Christopher P. (2006). "Endolithic
Cyanobacteria in Halite Rocks from the Hyperarid Core of the Atacama Desert".
2289:
Kaehler, S.; McQuaid, C. D. (1999). "Lethal and sub-lethal effects of phototrophic endoliths attacking the shell of the intertidal mussel Perna perna".
1784:
Walker, Jeffrey J.; Spear, John R.; Pace, Norman R. (2005). "Geobiology of a microbial endolithic community in the
Yellowstone geothermal environment".
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Omelon, C. R.; Pollard, W. H.; Ferris, F. G. (2006). "Environmental controls on microbial colonization of high Arctic cryptoendolithic habitats".
484:
Endolithic microorganisms have been considered a model for the search for life on other planets by inquiring about what sort of microorganisms on
129:
The term "endolith", which defines an organism that colonizes the interior of any kind of rock, has been further classified into five subclasses:
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Colonizes structural cavities within natural pore spaces within the rocks. These pores are usually indirectly connected to the rock surface; (
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help them in the protection against dangerous radiation and is a way to obtain energy. Another characteristic is the presence of a very slow
156:
Penetrates actively into the interior of rocks forming channels and grooves that conform with the shape of its body, rock boring organism (
843:
1923:
1508:
Friedmann, E. I.; Weed, R. (1987). "Microbial trace-fossil formation, biogenous, and abiotic weathering in the
Antarctic cold desert".
496:. Several studies have been carried out in extreme places that serve as analogs for Mars's surface and subsurface, and many studies in
951:"Endolithic cyanobacteria in soil gypsum: Occurrences in Atacama (Chile), Mojave (United States), and Al-Jafr Basin (Jordan) Deserts"
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2882:
1226:
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Cowan, D. A.; Khan, N.; Pointing, S. B.; Cary, S. C. (2010). "Diverse hypolithic refuge communities in the McMurdo Dry Valleys".
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and its diversity even though there is a probability that endolithic fungi could perhaps play an important role in the health of
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3176:
995:"Cyanobacteria and chloroflexi-dominated hypolithic colonization of quartz at the hyper-arid core of the Atacama Desert, Chile"
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Cockell, C.; Osinski, G.; Lee, P. (2003). "The impact crater as a habitat: effects of impact processing of target materials".
3489:
2327:
Gong, YiMing; Xu, Ran; Hu, Bi (2008). "Endolithic fungi: A possible killer for the mass extinction of Cretaceous dinosaurs".
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Smith, M. C.; Bowman, J. P.; Scott, F. J.; Line, M. A. (2000). "Sublithic bacteria associated with Antarctic quartz stones".
118:
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The metabolism of endolithic microorganisms is versatile, in many of those communities have been found genes involved in
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3396:
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2556:
2459:"An endoevaporitic microbial mat within a gypsum crust: zonation of phototrophs, photopigments, and light penetration"
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endoliths were thought of as somewhat benign, but evidence has since surfaced that phototrophic endoliths (primarily
1736:"Molecular Characterization of an Endolithic Microbial Community in Dolomite Rock in the Central Alps (Switzerland)"
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2800:
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393:
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Golubic, Stjepko; Radtke, Gudrun; Campion-Alsumard, Therese Le (2005). "Endolithic fungi in marine ecosystems".
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492:, which helps to propose those lithologies as life detection targets on an extra-terrestrial surface such as
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growth and reproduction. In deeper rock layers microbes are exposed to high pressures and temperatures.
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The IUCN Global Ecosystem Typology 2.0: Descriptive profiles for biomes and ecosystem functional groups
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Makhalanyane, T. P.; Pointing, S. B.; Cowan, D. A. (2014). "Lithobionts: Cryptic and Refuge Niches".
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Colonizes the pore spaces located on the underside of the rock and that make contact with the soil (
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Keith, DA; Iliffe, TM; Gerovasileiou, V; Gonzalez, B; Brankovits, D; MartĂnez GarcĂa, A (2020).
859:
323:, and very little is available for reproduction or growth. It is thought that they weather long
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floor. Endolithic organisms have also been found in surface rocks in regions of low humidity (
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Advances in environmental microbiology. Their World: A Diversity of Microbial Environments
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can reproduce at 121 °C), which limits the possible depth to 4-4.5 km below the
550:
Omelon, CR (2016). "Endolithic Microorganisms and Their Habitats". In Hurst, C.J. (ed.).
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1992:
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430:(Mexico). The endoliths found there not only included algae and fungi but also included
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According to a study done by Astrid Gunther endoliths were also found in the island of
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2207:"Endolithic algae: an alternative source of photoassimilates during coral bleaching"
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1702:
1632:— Gold mines present "ideal environment" for geologists studying subsurface microbes
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1322:
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81:) that is able to acquire the necessary resources for growth in the inner part of a
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Lysnes, Kristine; Torsvik, Terje; Thorseth, Ingunn H.; Pedersen, Rolf B. (2004).
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1345:
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Bell, R. A. (1993). "Cryptoendolithic algae of hot semiarid lands and deserts".
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2183:
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1336:
Friedmann, E. I. (1980). "Endolithic Microbial Life in Hot and Cold Deserts".
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31:
2129:. In Keith, D.A.; Ferrer-Paris, J.R.; Nicholson, E.; Kingsford, R.T. (eds.).
1521:
3452:
3447:
3263:
2811:
2690:
2646:
2606:
1837:"Phylogenetic Composition of Rocky Mountain Endolithic Microbial Ecosystems"
844:"The lithobiotic ecological niche, with special reference to microorganisms"
336:
199:
62:
37:
2443:
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2240:
2222:
2191:
2086:"Soil beneath ocean found to harbor long-lived bacteria, fungi and viruses"
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2018:
2000:
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1720:
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1269:
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2302:
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2211:
Proceedings of the Royal Society of London. Series B: Biological Sciences
1860:
1673:
de la Torre, J. R.; Goebel, B. M.; Friedmann, E. I.; Pace, N. R. (2003).
975:
950:
343:
where these different endolithic species live together has been called a
207:
98:
78:
54:
1805:
1647:"Looking for life in all the wrong places — research on cryptoendoliths"
1218:
890:
454:) have infested 50 to 80% of midshore populations of the mussel species
3186:
2772:
2483:
2458:
2267:
2009:
1947:"Microbial Populations in Ocean Floor Basalt: Results from ODP Leg 187"
489:
435:
427:
340:
324:
102:
86:
58:
1313:
1165:
1163:
1093:
730:
Walker, J. J.; Pace, N. R. (2007). "Endolithic microbial ecosystems".
3160:
3050:
2762:
1206:
878:
842:
Golubic, Stjepko; Friedmann, E. Imre; Schneider, JĂĽrgen (June 1981).
263:
70:
66:
1260:
1243:
1056:
136:
Colonizes fissures and cracks in the rock connected to the surface (
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Spectrochimica Acta Part A: Molecular and Biomolecular Spectroscopy
376:
Only limited research has been done concerning the distribution of
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3026:
485:
358:
327:
in this fashion, emerging when the temperature in the area warms.
291:
203:
90:
74:
36:
1675:"Microbial Diversity of Cryptoendolithic Communities from the Mc
3213:
2541:
493:
227:
114:
2991:
2545:
993:; Warren-Rhodes, K. A.; McKay, C. P.; Pointing, S. B. (2011).
500:
on Earth's hot and cold deserts have been developed. In these
279:
2108:"Frequently Requested Information about the SLiME Hypothesis"
949:
Dong, H; Rech, J. A.; Jiang, H; Sun, H; Buck, B. J. (2007).
468:
Endolithic fungi found in the eggs of Cretaceous dinosaurs
194:
organisms, the temperature limit is at about 120 °C (
763:
761:
282:. Photosynthetic endoliths have also been discovered.
176:
Capable of rocks formation by mineral depositation (
3477:
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3364:
3308:
3297:
3277:
3204:
3169:
3143:
3105:
3074:
3043:
3034:
3025:
2860:
2810:
2771:
2708:
2699:
2579:
1734:Horath, Thomas; Bachofen, Reinhard (August 2009).
1892:
1890:
1888:
41:Endolith lifeform found inside an Antarctic rock
955:Journal of Geophysical Research: Biogeosciences
372:Endolithic fungi and algae in marine ecosystems
346:subsurface lithoautotrophic microbial ecosystem
113:, who theorize that endolithic environments on
306:cycle. Early data suggest some only engage in
3003:
2557:
1956:. Proceedings of the Ocean Drilling Program.
8:
1244:"Widespread colonization by polar hypoliths"
121:for extraterrestrial microbial communities.
3151:Mediterranean forests, woodlands, and scrub
1626:. Princeton Weekly Bulletin. Archived from
860:10.1306/212F7CB6-2B24-11D7-8648000102C1865D
816:"Visions of Life on Mars in Earth's Depths"
3305:
3040:
3031:
3010:
2996:
2988:
2705:
2564:
2550:
2542:
1207:"EcologĂa microbiana de sustratos lĂticos"
879:"EcologĂa microbiana de sustratos lĂticos"
2482:
2329:Science in China Series D: Earth Sciences
2230:
2008:
1868:
1710:
1640:
1638:
1312:
1259:
1092:
1018:
974:
397:. Discovery of endolithic fungi, such as
2457:Oren, A.; KĂĽhl, M.; Karsten, U. (1995).
1899:Model Ecosystems in Extreme Environments
2500:
2322:
2320:
542:
117:and other planets constitute potential
1841:Applied and Environmental Microbiology
1683:Applied and Environmental Microbiology
1379:
1369:
1242:Cockell, C. S.; Stokes, M. D. (2004).
1126:Analytical and Bioanalytical Chemistry
744:10.1146/annurev.micro.61.080706.093302
202:crust, and 7 or 7.5 km below the
2096:from the original on 29 October 2015.
7:
3135:Grasslands, savannas, and shrublands
3097:Grasslands, savannas, and shrublands
1622:Schultz, Steven (13 December 1999).
848:SEPM Journal of Sedimentary Research
814:Chang, Kenneth (12 September 2016).
1835:Walker, J. J.; Pace, N. R. (2007).
438:as well as many other microborers.
1477:Antarctic Terrestrial Microbiology
1340:. Vol. 10. pp. 223–235.
512:) on Mars or other rocky planets.
25:
3061:Montane grasslands and shrublands
2883:Acidophiles in acid mine drainage
2205:Fine, Maoz; Loya, Yossi (2002).
1966:10.2973/odp.proc.sr.187.203.2004
1184:10.1111/j.0022-3646.1993.00133.x
785:10.1111/j.1472-4669.2010.00254.x
3177:Flooded grasslands and savannas
1703:10.1128/AEM.69.7.3858-3867.2003
357:within the subterranean lithic
3490:Ecological land classification
3019:Biogeographic regionalisations
2463:Marine Ecology Progress Series
1924:"Glass Munchers Under the Sea"
480:Relationship with astrobiology
1:
2084:Yirka, Bob (29 August 2013).
1593:10.1080/08927014.2016.1269893
27:Organism living inside a rock
3156:Deserts and xeric shrublands
2536:Endolith Advanced Collection
2530:Endoliths General Collection
2133:. Gland, Switzerland: IUCN.
554:. Cincinnati, USA: Springer.
262:found microscopic trails in
105:grains of a rock. Many are
3397:Temperate Northern Atlantic
3087:Broadleaf and mixed forests
1928:NASA Astrobiology Institute
1485:10.1007/978-3-642-45213-0_9
1346:10.1007/978-94-009-9085-2_3
3544:
3500:Vegetation classifications
3402:Temperate Northern Pacific
2868:Abiogenic petroleum origin
2801:Thermococcus gammatolerans
2436:10.1089/153110703321632507
2139:10.2305/IUCN.CH.2020.13.en
29:
3412:Temperate Southern Africa
2385:10.1007/s00248-006-9055-7
2349:10.1007/s11430-008-0052-1
2184:10.1016/j.tim.2005.03.007
2127:"S1.2 Endolithic systems"
2055:10.1016/j.saa.2019.01.013
1755:10.1007/s00248-008-9483-7
1645:Hively, Will (May 1997).
1557:10.1080/01490450290098577
1454:10.1007/s00300-006-0155-0
1419:10.1017/S0954102000000237
1305:10.1017/S0954102010000507
1138:10.1007/s00216-016-9497-9
1085:10.1007/s00792-013-0519-7
1011:10.1007/s00792-010-0334-3
928:10.1017/S0024282904013854
701:10.1007/s00248-007-9235-0
658:10.1007/s00248-011-9864-1
580:10.1080/01490450903061101
408:In the wake of worldwide
394:Lithopythium gangliiforme
30:For the dental term, see
3422:Tropical Eastern Pacific
3372:Antarctic/Southern Ocean
2719:Chloroflexus aurantiacus
1679:Dry Valleys, Antarctica"
1522:10.1126/science.11536571
1211:Ciencia y Medio Ambiente
883:Ciencia y Medio Ambiente
3443:Biogeographic provinces
3407:Temperate South America
3125:Moist broadleaf forests
2842:Halicephalobus mephisto
2835:Paralvinella sulfincola
2821:Cyanidioschyzon merolae
2726:Deinococcus radiodurans
1624:"Two miles underground"
1545:Geomicrobiology Journal
568:Geomicrobiology Journal
378:marine endolithic fungi
238:Metabolism and survival
210:) and low temperature (
2223:10.1098/rspb.2002.1983
2172:Trends in Microbiology
2001:10.1089/ast.2006.6.415
389:Ostracoblabe implexis
335:As most endoliths are
260:Ocean Drilling Program
42:
3505:Zoogeographic regions
3463:Global 200 ecoregions
3392:Temperate Australasia
3130:Dry broadleaf forests
2828:Galdieria sulphuraria
2757:Spirochaeta americana
2303:10.1007/s002270050650
2114:on 30 September 2006.
623:10.1089/ast.2008.0278
442:Endolithic parasitism
40:
3485:Altitudinal zonation
3427:Western Indo-Pacific
3387:Eastern Indo-Pacific
3382:Central Indo-Pacific
2750:Thermus thermophilus
1954:Proc ODP Sci Results
1934:on 20 February 2013.
1861:10.1128/AEM.02656-06
1479:. pp. 163–179.
1172:Journal of Phycology
976:10.1029/2006JG000385
866:on 30 December 2010.
502:extreme environments
278:oceans that contain
18:Endolithic ecosystem
3458:Lists of ecoregions
3115:Tropical rainforest
2949:Radiotrophic fungus
2926:Helaeomyia petrolei
2873:Acidithiobacillales
2782:Pyrococcus furiosus
2475:1995MEPS..128..151O
2428:2003AsBio...3..181C
2341:2008ScChD..51..801G
2217:(1497): 1205–1210.
2047:2019AcSpA.212..262O
1993:2006AsBio...6..415W
1853:2007ApEnM..73.3497W
1806:10.1038/nature03447
1798:2005Natur.434.1011W
1792:(7036): 1011–1014.
1695:2003ApEnM..69.3858D
1630:on 13 January 2016.
1411:2000AntSc..12..177S
1297:2010AntSc..22..714C
967:2007JGRG..112.2030D
615:2009AsBio...9..369H
3495:Floristic kingdoms
3259:Hydrothermal vents
3120:Coniferous forests
3082:Coniferous forests
2484:10.3354/meps128151
2268:10.1007/bf02536953
1901:. Academic press.
1205:Ascaso, C (2002).
877:Ascaso, C (2002).
821:The New York Times
732:Annu Rev Microbiol
365:Endolithic systems
355:endolithic systems
244:sulphur metabolism
43:
3513:
3512:
3473:
3472:
3417:Tropical Atlantic
3293:
3292:
3200:
3199:
3092:Deciduous forests
2985:
2984:
2932:Hydrothermal vent
2856:
2855:
2794:Pyrolobus fumarii
2743:Thermus aquaticus
2373:Microbial Ecology
2148:978-2-8317-2077-7
1908:978-0-1281-2742-1
1847:(11): 3497–3504.
1743:Microbial Ecology
1516:(4802): 703–705.
1494:978-3-642-45212-3
1399:Antarctic Science
1355:978-94-009-9087-6
1285:Antarctic Science
1132:(15): 4083–4092.
1051:(12): 3222–3231.
916:The Lichenologist
689:Microbial Ecology
646:Microbial Ecology
488:inhabit specific
256:nitrogen fixation
214:), including the
16:(Redirected from
3535:
3306:
3041:
3032:
3012:
3005:
2998:
2989:
2888:Archaeoglobaceae
2861:Related articles
2706:
2686:Thermoacidophile
2681:Hyperthermophile
2657:Polyextremophile
2566:
2559:
2552:
2543:
2517:
2514:
2508:
2505:
2489:
2488:
2486:
2454:
2448:
2447:
2411:
2405:
2404:
2367:
2361:
2360:
2324:
2315:
2314:
2286:
2280:
2279:
2251:
2245:
2244:
2234:
2202:
2196:
2195:
2167:
2161:
2160:
2122:
2116:
2115:
2110:. Archived from
2104:
2098:
2097:
2081:
2075:
2074:
2029:
2023:
2022:
2012:
1976:
1970:
1969:
1951:
1942:
1936:
1935:
1930:. Archived from
1922:Mullen, Leslie.
1919:
1913:
1912:
1894:
1883:
1882:
1872:
1832:
1826:
1825:
1781:
1775:
1774:
1740:
1731:
1725:
1724:
1714:
1689:(7): 3858–3867.
1670:
1664:
1663:
1661:
1659:
1642:
1633:
1631:
1619:
1613:
1612:
1575:
1569:
1568:
1540:
1534:
1533:
1505:
1499:
1498:
1472:
1466:
1465:
1437:
1431:
1430:
1394:
1388:
1387:
1381:
1377:
1375:
1367:
1333:
1327:
1326:
1316:
1280:
1274:
1273:
1263:
1239:
1233:
1232:
1202:
1196:
1195:
1167:
1158:
1157:
1121:
1115:
1114:
1096:
1067:
1061:
1060:
1039:
1033:
1032:
1022:
987:
981:
980:
978:
946:
940:
939:
911:
905:
904:
874:
868:
867:
862:. Archived from
839:
833:
832:
830:
828:
811:
805:
804:
765:
756:
755:
727:
721:
720:
684:
678:
677:
641:
635:
634:
598:
592:
591:
562:
556:
555:
547:
446:Until the 1990s
399:Dodgella priscus
192:hyperthermophile
21:
3543:
3542:
3538:
3537:
3536:
3534:
3533:
3532:
3518:
3517:
3514:
3509:
3469:
3431:
3360:
3300:
3289:
3285:Endolithic zone
3273:
3206:
3196:
3165:
3139:
3107:
3101:
3070:
3036:
3021:
3016:
2986:
2981:
2972:Thermostability
2908:Grylloblattidae
2878:Acidobacteriota
2852:
2806:
2767:
2701:
2695:
2637:Metallotolerant
2575:
2570:
2526:
2521:
2520:
2515:
2511:
2506:
2502:
2497:
2492:
2456:
2455:
2451:
2422:(1): 3181–191.
2413:
2412:
2408:
2369:
2368:
2364:
2326:
2325:
2318:
2288:
2287:
2283:
2253:
2252:
2248:
2204:
2203:
2199:
2169:
2168:
2164:
2149:
2124:
2123:
2119:
2106:
2105:
2101:
2083:
2082:
2078:
2031:
2030:
2026:
1978:
1977:
1973:
1949:
1944:
1943:
1939:
1921:
1920:
1916:
1909:
1896:
1895:
1886:
1834:
1833:
1829:
1783:
1782:
1778:
1738:
1733:
1732:
1728:
1672:
1671:
1667:
1657:
1655:
1644:
1643:
1636:
1621:
1620:
1616:
1577:
1576:
1572:
1542:
1541:
1537:
1507:
1506:
1502:
1495:
1474:
1473:
1469:
1439:
1438:
1434:
1396:
1395:
1391:
1378:
1368:
1356:
1335:
1334:
1330:
1282:
1281:
1277:
1261:10.1038/431414a
1241:
1240:
1236:
1229:
1204:
1203:
1199:
1169:
1168:
1161:
1123:
1122:
1118:
1069:
1068:
1064:
1057:10.1890/02-0549
1041:
1040:
1036:
989:
988:
984:
948:
947:
943:
913:
912:
908:
901:
876:
875:
871:
841:
840:
836:
826:
824:
813:
812:
808:
767:
766:
759:
729:
728:
724:
686:
685:
681:
643:
642:
638:
600:
599:
595:
564:
563:
559:
549:
548:
544:
540:
518:
498:geomicrobiology
482:
470:
444:
410:coral bleaching
374:
333:
252:carbon fixation
248:iron metabolism
240:
232:Rocky Mountains
187:
127:
111:astrobiologists
35:
28:
23:
22:
15:
12:
11:
5:
3541:
3539:
3531:
3530:
3520:
3519:
3511:
3510:
3508:
3507:
3502:
3497:
3492:
3487:
3481:
3479:
3475:
3474:
3471:
3470:
3468:
3467:
3466:
3465:
3460:
3450:
3445:
3439:
3437:
3433:
3432:
3430:
3429:
3424:
3419:
3414:
3409:
3404:
3399:
3394:
3389:
3384:
3379:
3374:
3368:
3366:
3362:
3361:
3359:
3358:
3353:
3348:
3343:
3338:
3333:
3328:
3323:
3318:
3312:
3310:
3303:
3299:Biogeographic
3295:
3294:
3291:
3290:
3288:
3287:
3281:
3279:
3275:
3274:
3272:
3271:
3266:
3261:
3256:
3251:
3246:
3241:
3236:
3231:
3226:
3221:
3216:
3210:
3208:
3202:
3201:
3198:
3197:
3195:
3194:
3189:
3184:
3179:
3173:
3171:
3167:
3166:
3164:
3163:
3158:
3153:
3147:
3145:
3141:
3140:
3138:
3137:
3132:
3127:
3122:
3117:
3111:
3109:
3103:
3102:
3100:
3099:
3094:
3089:
3084:
3078:
3076:
3072:
3071:
3069:
3068:
3063:
3058:
3053:
3047:
3045:
3038:
3029:
3023:
3022:
3017:
3015:
3014:
3007:
3000:
2992:
2983:
2982:
2980:
2979:
2974:
2969:
2961:
2956:
2951:
2946:
2941:
2934:
2929:
2922:
2915:
2910:
2905:
2903:Thermoproteota
2900:
2895:
2890:
2885:
2880:
2875:
2870:
2864:
2862:
2858:
2857:
2854:
2853:
2851:
2850:
2845:
2838:
2831:
2824:
2816:
2814:
2808:
2807:
2805:
2804:
2797:
2790:
2785:
2777:
2775:
2769:
2768:
2766:
2765:
2760:
2753:
2746:
2739:
2734:
2729:
2722:
2714:
2712:
2703:
2697:
2696:
2694:
2693:
2688:
2683:
2674:
2672:Radioresistant
2669:
2664:
2659:
2654:
2649:
2644:
2639:
2634:
2629:
2624:
2622:Lithoautotroph
2619:
2614:
2609:
2604:
2599:
2594:
2589:
2583:
2581:
2577:
2576:
2571:
2569:
2568:
2561:
2554:
2546:
2540:
2539:
2533:
2525:
2524:External links
2522:
2519:
2518:
2509:
2499:
2498:
2496:
2493:
2491:
2490:
2449:
2406:
2379:(3): 389–398.
2362:
2335:(6): 801–807.
2316:
2297:(3): 497–503.
2291:Marine Biology
2281:
2262:(1): 233–261.
2246:
2197:
2178:(5): 229–235.
2162:
2147:
2117:
2099:
2076:
2024:
1987:(3): 415–422.
1971:
1937:
1914:
1907:
1884:
1827:
1776:
1749:(2): 290–306.
1726:
1665:
1634:
1614:
1587:(2): 113–127.
1570:
1551:(6): 535–552.
1535:
1500:
1493:
1467:
1432:
1405:(2): 177–184.
1389:
1380:|journal=
1354:
1338:Limits of Life
1328:
1291:(6): 714–720.
1275:
1234:
1227:
1197:
1178:(2): 133–139.
1159:
1116:
1079:(2): 329–337.
1062:
1034:
982:
941:
906:
899:
869:
854:(2): 475–478.
834:
806:
757:
722:
695:(4): 740–752.
679:
636:
609:(4): 369–381.
593:
574:(7): 491–507.
557:
541:
539:
536:
535:
534:
529:
524:
517:
514:
481:
478:
469:
466:
443:
440:
403:Conchyliastrum
373:
370:
332:
329:
296:beta carotenes
286:especially in
239:
236:
186:
183:
182:
181:
174:
171:
164:
161:
154:
151:
144:
143:Cryptoendolith
141:
134:
133:Chasmoendolith
126:
125:Subdefinitions
123:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
3540:
3529:
3526:
3525:
3523:
3516:
3506:
3503:
3501:
3498:
3496:
3493:
3491:
3488:
3486:
3483:
3482:
3480:
3476:
3464:
3461:
3459:
3456:
3455:
3454:
3451:
3449:
3446:
3444:
3441:
3440:
3438:
3434:
3428:
3425:
3423:
3420:
3418:
3415:
3413:
3410:
3408:
3405:
3403:
3400:
3398:
3395:
3393:
3390:
3388:
3385:
3383:
3380:
3378:
3375:
3373:
3370:
3369:
3367:
3363:
3357:
3354:
3352:
3349:
3347:
3344:
3342:
3339:
3337:
3334:
3332:
3329:
3327:
3324:
3322:
3319:
3317:
3314:
3313:
3311:
3307:
3304:
3302:
3296:
3286:
3283:
3282:
3280:
3276:
3270:
3269:Demersal zone
3267:
3265:
3262:
3260:
3257:
3255:
3252:
3250:
3247:
3245:
3242:
3240:
3237:
3235:
3232:
3230:
3227:
3225:
3222:
3220:
3217:
3215:
3212:
3211:
3209:
3203:
3193:
3190:
3188:
3185:
3183:
3180:
3178:
3175:
3174:
3172:
3168:
3162:
3159:
3157:
3154:
3152:
3149:
3148:
3146:
3142:
3136:
3133:
3131:
3128:
3126:
3123:
3121:
3118:
3116:
3113:
3112:
3110:
3104:
3098:
3095:
3093:
3090:
3088:
3085:
3083:
3080:
3079:
3077:
3073:
3067:
3066:Alpine tundra
3064:
3062:
3059:
3057:
3054:
3052:
3049:
3048:
3046:
3044:Polar/montane
3042:
3039:
3033:
3030:
3028:
3024:
3020:
3013:
3008:
3006:
3001:
2999:
2994:
2993:
2990:
2978:
2975:
2973:
2970:
2968:
2966:
2962:
2960:
2957:
2955:
2952:
2950:
2947:
2945:
2942:
2940:
2939:
2935:
2933:
2930:
2928:
2927:
2923:
2921:
2920:
2919:Halobacterium
2916:
2914:
2911:
2909:
2906:
2904:
2901:
2899:
2896:
2894:
2891:
2889:
2886:
2884:
2881:
2879:
2876:
2874:
2871:
2869:
2866:
2865:
2863:
2859:
2849:
2846:
2844:
2843:
2839:
2837:
2836:
2832:
2830:
2829:
2825:
2823:
2822:
2818:
2817:
2815:
2813:
2809:
2803:
2802:
2798:
2796:
2795:
2791:
2789:
2786:
2784:
2783:
2779:
2778:
2776:
2774:
2770:
2764:
2761:
2759:
2758:
2754:
2752:
2751:
2747:
2745:
2744:
2740:
2738:
2735:
2733:
2730:
2728:
2727:
2723:
2721:
2720:
2716:
2715:
2713:
2711:
2707:
2704:
2702:extremophiles
2698:
2692:
2689:
2687:
2684:
2682:
2678:
2675:
2673:
2670:
2668:
2665:
2663:
2660:
2658:
2655:
2653:
2650:
2648:
2645:
2643:
2640:
2638:
2635:
2633:
2630:
2628:
2625:
2623:
2620:
2618:
2615:
2613:
2610:
2608:
2605:
2603:
2600:
2598:
2595:
2593:
2590:
2588:
2585:
2584:
2582:
2578:
2574:
2573:Extremophiles
2567:
2562:
2560:
2555:
2553:
2548:
2547:
2544:
2537:
2534:
2531:
2528:
2527:
2523:
2513:
2510:
2504:
2501:
2494:
2485:
2480:
2476:
2472:
2468:
2464:
2460:
2453:
2450:
2445:
2441:
2437:
2433:
2429:
2425:
2421:
2417:
2410:
2407:
2402:
2398:
2394:
2390:
2386:
2382:
2378:
2374:
2366:
2363:
2358:
2354:
2350:
2346:
2342:
2338:
2334:
2330:
2323:
2321:
2317:
2312:
2308:
2304:
2300:
2296:
2292:
2285:
2282:
2277:
2273:
2269:
2265:
2261:
2257:
2250:
2247:
2242:
2238:
2233:
2228:
2224:
2220:
2216:
2212:
2208:
2201:
2198:
2193:
2189:
2185:
2181:
2177:
2173:
2166:
2163:
2158:
2154:
2150:
2144:
2140:
2136:
2132:
2128:
2121:
2118:
2113:
2109:
2103:
2100:
2095:
2091:
2087:
2080:
2077:
2072:
2068:
2064:
2060:
2056:
2052:
2048:
2044:
2040:
2036:
2028:
2025:
2020:
2016:
2011:
2006:
2002:
1998:
1994:
1990:
1986:
1982:
1975:
1972:
1967:
1963:
1959:
1955:
1948:
1941:
1938:
1933:
1929:
1925:
1918:
1915:
1910:
1904:
1900:
1893:
1891:
1889:
1885:
1880:
1876:
1871:
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1478:
1471:
1468:
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1447:
1443:
1442:Polar Biology
1436:
1433:
1428:
1424:
1420:
1416:
1412:
1408:
1404:
1400:
1393:
1390:
1385:
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1310:
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1302:
1298:
1294:
1290:
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1279:
1276:
1271:
1267:
1262:
1257:
1254:(7007): 414.
1253:
1249:
1245:
1238:
1235:
1230:
1228:9788469979723
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1220:
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1198:
1193:
1189:
1185:
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1173:
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1139:
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1127:
1120:
1117:
1112:
1108:
1104:
1100:
1095:
1090:
1086:
1082:
1078:
1074:
1073:Extremophiles
1066:
1063:
1058:
1054:
1050:
1046:
1038:
1035:
1030:
1026:
1021:
1016:
1012:
1008:
1004:
1000:
999:Extremophiles
996:
992:
986:
983:
977:
972:
968:
964:
960:
956:
952:
945:
942:
937:
933:
929:
925:
921:
917:
910:
907:
902:
900:9788469979723
896:
892:
888:
884:
880:
873:
870:
865:
861:
857:
853:
849:
845:
838:
835:
823:
822:
817:
810:
807:
802:
798:
794:
790:
786:
782:
778:
774:
773:
764:
762:
758:
753:
749:
745:
741:
737:
733:
726:
723:
718:
714:
710:
706:
702:
698:
694:
690:
683:
680:
675:
671:
667:
663:
659:
655:
651:
647:
640:
637:
632:
628:
624:
620:
616:
612:
608:
604:
597:
594:
589:
585:
581:
577:
573:
569:
561:
558:
553:
546:
543:
537:
533:
530:
528:
525:
523:
520:
519:
515:
513:
511:
510:biosignatures
505:
503:
499:
495:
491:
487:
479:
477:
473:
467:
465:
463:
459:
458:
453:
452:cyanobacteria
449:
441:
439:
437:
433:
432:cyanobacteria
429:
424:
422:
419:
418:endosymbiotic
415:
411:
406:
404:
400:
396:
395:
390:
385:
383:
379:
371:
369:
366:
362:
360:
356:
352:
348:
347:
342:
338:
330:
328:
326:
322:
318:
314:
309:
308:cell division
305:
301:
297:
293:
289:
288:cyanobacteria
283:
281:
277:
273:
269:
265:
261:
257:
253:
249:
245:
237:
235:
233:
229:
225:
221:
217:
213:
209:
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197:
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179:
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169:
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159:
155:
152:
149:
145:
142:
139:
135:
132:
131:
130:
124:
122:
120:
116:
112:
108:
107:extremophiles
104:
100:
96:
95:animal shells
92:
88:
84:
80:
76:
72:
68:
64:
60:
56:
52:
48:
39:
33:
19:
3515:
3436:Subdivisions
3326:Australasian
3316:Afrotropical
3284:
3278:Other biomes
3254:Benthic zone
3249:Pelagic zone
3244:Neritic zone
3234:Kelp forests
3106:Tropical and
2977:Thermotogota
2964:
2938:Methanopyrus
2936:
2924:
2917:
2913:Halobacteria
2893:Berkeley Pit
2848:Pompeii worm
2840:
2833:
2826:
2819:
2799:
2792:
2780:
2755:
2748:
2741:
2732:Deinococcota
2724:
2717:
2679: /
2667:Psychrophile
2601:
2512:
2503:
2466:
2462:
2452:
2419:
2416:Astrobiology
2415:
2409:
2376:
2372:
2365:
2332:
2328:
2294:
2290:
2284:
2259:
2255:
2249:
2214:
2210:
2200:
2175:
2171:
2165:
2130:
2120:
2112:the original
2102:
2089:
2079:
2038:
2034:
2027:
1984:
1981:Astrobiology
1980:
1974:
1957:
1953:
1940:
1932:the original
1927:
1917:
1898:
1844:
1840:
1830:
1789:
1785:
1779:
1746:
1742:
1729:
1686:
1682:
1676:
1668:
1656:. Retrieved
1650:
1628:the original
1617:
1584:
1580:
1573:
1548:
1544:
1538:
1513:
1509:
1503:
1476:
1470:
1448:(1): 19–29.
1445:
1441:
1435:
1402:
1398:
1392:
1337:
1331:
1288:
1284:
1278:
1251:
1247:
1237:
1219:10261/111133
1210:
1200:
1175:
1171:
1129:
1125:
1119:
1076:
1072:
1065:
1048:
1044:
1037:
1005:(1): 31–38.
1002:
998:
991:Lacap, D. C.
985:
958:
954:
944:
922:(1): 55–73.
919:
915:
909:
891:10261/111133
882:
872:
864:the original
851:
847:
837:
827:12 September
825:. Retrieved
819:
809:
779:(1): 44–60.
776:
770:
735:
731:
725:
692:
688:
682:
652:(1): 69–79.
649:
645:
639:
606:
603:Astrobiology
602:
596:
571:
567:
560:
551:
545:
506:
483:
474:
471:
462:South Africa
455:
448:phototrophic
445:
425:
414:Coelenterate
407:
402:
398:
392:
388:
386:
375:
364:
363:
354:
350:
344:
334:
321:racemization
304:reproduction
284:
241:
212:psychrophile
188:
177:
173:Autoendolith
167:
163:Hypoendolith
157:
147:
137:
128:
97:, or in the
50:
46:
44:
3528:Lithophiles
3346:Neotropical
3336:Indomalayan
3309:Terrestrial
3239:Coral reefs
3108:subtropical
3035:Terrestrial
2944:Movile Cave
2898:Blood Falls
2677:Thermophile
2662:Psammophile
2592:Alkaliphile
2516:References:
2507:References:
2469:: 151–159.
2041:: 262–271.
2010:10261/19099
738:: 331–347.
532:Volyn biota
460:located in
457:Perna perna
421:Symbiodinia
382:coral reefs
317:cosmic rays
313:cell damage
300:chlorophyll
294:, such as;
216:Dry Valleys
200:continental
185:Environment
3453:Ecoregions
3448:Bioregions
3356:Palearctic
3264:Cold seeps
3224:Intertidal
2967:polymerase
2959:Tardigrade
2788:Strain 121
2652:Piezophile
2642:Oligotroph
2632:Methanogen
2627:Lithophile
2597:Capnophile
2587:Acidophile
1658:5 December
1581:Biofouling
1314:10289/5090
1213:: 90–103.
1094:10566/3555
961:(G2): G2.
885:: 90–103.
772:Geobiology
538:References
527:Lithotroph
522:Lithophile
337:autotrophs
315:caused by
230:, and the
224:Antarctica
220:permafrost
196:Strain 121
153:Euendolith
51:endolithic
32:Pulp stone
3331:Holarctic
3321:Antarctic
3229:Mangroves
3075:Temperate
2954:Rio Tinto
2812:Eukaryota
2691:Xerophile
2647:Osmophile
2617:Lipophile
2607:Halophile
2357:126670640
2276:130403994
2157:241360441
1382:ignored (
1372:cite book
588:131694781
290:and some
266:from the
150:= hidden)
63:bacterium
3522:Category
3478:See also
3351:Oceanian
3341:Nearctic
3219:Littoral
3192:Mangrove
3182:Riparian
2737:Snottite
2710:Bacteria
2612:Hypolith
2602:Endolith
2444:12804371
2393:16865610
2311:84103549
2241:12065035
2192:15866040
2094:Archived
2090:Phys.org
2071:58604046
2063:30658280
2019:16805697
1960:: 1–27.
1879:17416689
1814:15846344
1763:19172216
1721:12839754
1652:Discover
1601:28054493
1565:84636295
1530:11536571
1462:22633158
1427:84337509
1323:53558610
1270:15386002
1192:85033484
1146:27055886
1111:11175962
1103:23397517
1029:21069402
936:86211017
793:20726901
752:17506683
717:19843927
709:17457639
674:23356098
666:21584756
631:19519213
516:See also
490:minerals
325:ice ages
268:Atlantic
208:hypolith
170:= under)
140:= cleft)
101:between
59:archaeon
55:organism
47:endolith
3205:Aquatic
3187:Wetland
2773:Archaea
2700:Notable
2471:Bibcode
2424:Bibcode
2401:1914122
2337:Bibcode
2232:1691023
2043:Bibcode
1989:Bibcode
1870:1932665
1849:Bibcode
1822:4408407
1794:Bibcode
1691:Bibcode
1609:3295932
1510:Science
1407:Bibcode
1364:6774304
1293:Bibcode
1154:8132118
1045:Ecology
1020:3017302
963:Bibcode
801:9458330
611:Bibcode
436:sponges
428:Cozumel
341:biotope
331:Ecology
276:Pacific
180:= self)
160:= true)
119:refugia
103:mineral
87:mineral
3377:Arctic
3365:Marine
3301:realms
3207:biomes
3161:Steppe
3051:Tundra
3037:biomes
3027:Biomes
2763:GFAJ-1
2442:
2399:
2391:
2355:
2309:
2274:
2256:Facies
2239:
2229:
2190:
2155:
2145:
2069:
2061:
2017:
1905:
1877:
1867:
1820:
1812:
1786:Nature
1771:845383
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1761:
1719:
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1709:
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1491:
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1362:
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1101:
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353:), or
274:, and
272:Indian
264:basalt
258:. The
226:, the
148:crypto
79:amoeba
71:lichen
67:fungus
53:is an
3056:Taiga
2580:Types
2495:Notes
2397:S2CID
2353:S2CID
2307:S2CID
2272:S2CID
2153:S2CID
2067:S2CID
1950:(PDF)
1818:S2CID
1767:S2CID
1739:(PDF)
1677:Murdo
1605:S2CID
1561:S2CID
1458:S2CID
1423:S2CID
1319:S2CID
1188:S2CID
1150:S2CID
1107:S2CID
932:S2CID
797:S2CID
713:S2CID
670:S2CID
584:S2CID
486:Earth
359:biome
351:SLiME
292:algae
204:ocean
138:chasm
99:pores
91:coral
75:algae
3214:Pond
2440:PMID
2389:PMID
2237:PMID
2188:PMID
2143:ISBN
2059:PMID
2015:PMID
1903:ISBN
1875:PMID
1810:PMID
1759:PMID
1717:PMID
1660:2019
1597:PMID
1526:PMID
1489:ISBN
1384:help
1360:PMID
1350:ISBN
1266:PMID
1223:ISBN
1142:PMID
1099:PMID
1025:PMID
895:ISBN
829:2016
789:PMID
748:PMID
705:PMID
662:PMID
627:PMID
494:Mars
416:and
401:and
391:and
298:and
250:and
228:Alps
218:and
178:auto
168:hypo
115:Mars
83:rock
3170:Wet
3144:Dry
2965:Taq
2479:doi
2467:128
2432:doi
2381:doi
2345:doi
2299:doi
2295:135
2264:doi
2227:PMC
2219:doi
2215:269
2180:doi
2135:doi
2051:doi
2039:212
2005:hdl
1997:doi
1962:doi
1958:187
1865:PMC
1857:doi
1802:doi
1790:434
1751:doi
1707:PMC
1699:doi
1589:doi
1553:doi
1518:doi
1514:236
1481:doi
1450:doi
1415:doi
1342:doi
1309:hdl
1301:doi
1256:doi
1252:431
1215:hdl
1180:doi
1134:doi
1130:408
1089:hdl
1081:doi
1053:doi
1015:PMC
1007:doi
971:doi
959:112
924:doi
887:hdl
856:doi
781:doi
740:doi
697:doi
654:doi
619:doi
576:doi
319:or
280:DNA
246:,
222:of
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