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Endolith lifeform found inside an Antarctic rock

An endolith or endolithic is an organism (archaeon, bacterium, fungus, lichen, algae or amoeba) that is able to acquire the necessary resources for growth in the inner part of a rock,^[1] mineral, coral, animal shells, or in the pores between mineral grains of a rock. Many are extremophiles, living in places long imagined 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.^[2]^[3]^[4]^[5] Normally, the endoliths colonize the areas within lithic substrates to withstand intense solar radiation, temperature fluctuations, wind, and desiccation.^[6] They are of particular interest to astrobiologists, who theorize that endolithic environments on Mars and other planets constitute potential refugia for extraterrestrial microbial communities.^[7]^[8]

Subdefinitions

The term "endolith", which defines an organism that colonizes the interior of any kind of rock, has been further classified into four subclasses:^[9]

Chasmoendolith: Colonizes fissures and cracks in the rock connected to the surface (chasm = cleft)
Cryptoendolith: Colonizes structural cavities within natural pore spaces within the rocks. These pores are usually indirectly connected to the rock surface; (crypto = hidden)
Euendolith: Penetrates actively into the interior of rocks forming channels and grooves that conform with the shape of its body, rock boring organism (eu = true)
Hypoendolith: Colonizes the pore spaces located on the underside of the rock and that make contact with the soil (hypo = under)
Autoendolith: Capable of rocks formation by mineral depositation (self = self)

Environment

Endolithic microorganisms have been reported in many areas around the globe, there are reports in warm hyper-arid and arid deserts such as Monjave and Sonora (E.U.), Atacama (Chile), Gobi (China, Mongolia), Negev (Israel), Namib (Namibia Angola), Al-Jafr basin (Jordania) and the Depression of Turpan (China),^[10] ^[11] ^[12] ^[13] ^[14] ^[15] ^[16](Bell, 1993), also in cold deserts as Arctic and Antarctic ^[17] ^[18] ^[19] ^[20] ^[21] ^[22] ^[23](Friedmann et al., 1987; Smith et al., 2000; Omelon et al., 2006; Makhalanyane et al., 2014), and deep subsoil and ocean trenches rocks (Inagaki et al., 2009). However, there are reports of endolithic microorganisms in inter-tropical zones ^[24], where humidity and solar radiation are significantly different from the above-mentioned biomes. Endoliths have been found in rock down to a depth of 3 kilometres (1.9 mi), though it is unknown if that is their limit (due to the cost involved in digging so deeply).^[25]^[26] The main threat to their survival seems not to result from the pressure at such depth, but from the increased temperature. Judging from hyperthermophile organisms, the temperature limit is at about 120 °C (Strain 121 can reproduce at 121 °C), which limits the possible depth to 4-4.5 km below the continental crust, and 7 or 7.5 km below the ocean floor. Endolithic organisms have also been found in surface rocks in regions of low humidity (hypolith) and low temperature (psychrophile), including the Dry Valleys and permafrost of Antarctica,^[27] the Alps,^[28] and the Rocky Mountains.^[29]^[30]

Metabolism and Survival

Endoliths can survive by feeding on traces of iron, potassium, or sulfur as well as some carbon. (See lithotroph.) Whether they metabolize these directly from the surrounding rock, or rather excrete an acid to dissolve them first is yet undetermined. The Ocean Drilling Program found microscopic trails in basalt from the Atlantic, Indian, and Pacific oceans that contain DNA.^[31]^[32] Photosynthetic endoliths have also been discovered.^[33]

As water and nutrients are rather sparse in the environment of the endolith, they have a very slow reproduction cycle. Early data suggest some only engage in cell division 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.^[34] Most of their energy is spent repairing cell damage caused by cosmic rays or racemization, and very little is available for reproduction or growth. It is thought that they weather long ice ages in this fashion, emerging when the temperature in the area warms.^[26]

Ecology

As most endoliths are autotrophs, 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-biotope where these different endolithic species live together has been called a Subsurface Lithoautotrophic Microbial Ecosystem (SLiME),^[35] or endolithic systems within the subterranean lithic biome.

Endolithic systems 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 growth and reproduction. In deeper rock layers microbes are exposed to high pressures and temperatures.^[36]

Endolithic fungi and algae in marine ecosystems

Only limited research has been done concerning the distribution of marine endolithic fungi and its diversity even though there is a probability that endolithic fungi could perhaps play an important role in the health of coral reefs.

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: Ostracoblabe implexis and Lithopythium gangliiforme. Discovery of endolithic fungi, such as Dodgella priscus and Conchyliastrum, 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.^[37]

In the wake of worldwide coral bleaching, 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 Coelenterate and endosymbiotic Symbiodinia.^[38]

According to a study done by Astrid Gunther endoliths were also found in the island of Cozumel (Mexico). The endoliths found there not only included algae and fungi but also included cyanobacteria, sponges as well as many other microborers.^[39]

Endolithic parasitism

Until the 1990s phototrophic endoliths were thought of as somewhat benign, but evidence has since surfaced that phototrophic endoliths (primarily cyanobacteria) have infested 50 to 80% of midshore populations of the mussel species Perna perna located in South Africa. 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.^[40]

Endolithic fungi and the mass extinction of Cretaceous dinosaurs

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."^[41]

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 fungi that formed on the eggshells would have resulted in the abnormal incubation of the eggs and may have contributed to the mass extinction of these dinosaurs. It may also have led to the preservation of dinosaur eggs, including some that contained embryos.^[41]

References

^ Omelon, CR (2016). "Endolithic Microorganisms and Their Habitats". In Hurst, C.J. (ed.). Advances in environmental microbiology. Their World: A Diversity of Microbial Environments. Cincinnati, USA: Springer.
^ 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". Geomicrobiology Journal. 26 (7): 491–507. doi:10.1080/01490450903061101.
^ Herrera, A.; Cockell, C. S.; Self, S.; Blaxter, M.; Reitner, J.; Thorsteinsson, T.; Tindle, A. G. (2009). "A cryptoendolithic community in volcanic glass". Astrobiology. 9 (4): 369–381. doi:10.1089/ast.2008.0278.
^ 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". Microbial ecology. 62 (1): 69–79. doi:10.1007/s00248-011-9864-1.
^ Omelon, C. R.; Pollard, W. H.; Ferris, F. G. (2007). "Inorganic species distribution and microbial diversity within high Arctic cryptoendolithic habitats". Microbial ecology. 54 (4): 740–752. doi:10.1007/s00248-007-9235-0.
^ Walker, J. J.; Pace, N. R. (2007). "Endolithic microbial ecosystems". Annu Rev Microbiol. 61: 331–347.
^ 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". Geobiology. 9 (1): 44–60. doi:10.1111/j.1472-4669.2010.00254.x. PMID 20726901.
^ Chang, Kenneth (12 September 2016). "Visions of Life on Mars in Earth's Depths". The New York Times. Retrieved 12 September 2016.
^ Golubic, Stjepko; Friedmann, E. Imre; Schneider, Jürgen (June 1981). "The lithobiotic ecological niche, with special reference to microorganisms". SEPM Journal of Sedimentary Research. 51 (2): 475–478. doi:10.1306/212F7CB6-2B24-11D7-8648000102C1865D. Archived from the original on 30 December 2010.
^ Ascaso, C (2002). "Ecología microbiana de sustratos líticos". Ciencia y medio ambiente: 90–103.
^ Bungartz, F; Garvie, L. A.; Nash, T. H. "Anatomy of the endolithic Sonoran Desert lichen Verrucaria rubrocincta Breuss: implications for biodeterioration and biomineralization". The Lichenologist. 36 (1): 55–73. doi:10.1017/S0024282904013854.
^ Dong, H; Rech, J. A.; Jiang, H; Sun, H; Buck, B. J. (2007). "Endolithic cyanobacteria in soil gypsum: Occurrences in Atacama (Chile), Mojave (United States), and Al-Jafr Basin (Jordan) Deserts". Journal of Geophysical Research: Biogeosciences. 112: G2. doi:10.1029/2006JG000385.
^ Lacap, D. C.; Warren-Rhodes, K. A.; McKay, C. P.; Pointing, S. B. (2011). "Cyanobacteria and chloroflexi-dominated hypolithic colonization of quartz at the hyper-arid core of the Atacama Desert, Chile". Extremophiles. 15 (1): 31–38. doi:10.1007/s00792-010-0334-3.
^ 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". Ecology. 84 (12): 3222–3231. doi:10.1890/02-0549.
^ 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". Extremophiles. 17 (2): 329–337. doi:10.1007/s00792-.
^ Vítek, 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". Analytical and bioanalytical chemistry. 408 (15): 4083–4092. doi:10.1007/s00216-016-9497-9.
^ Ascaso, C (2002). "Ecología microbiana de sustratos líticos". Ciencia y medio ambiente: 90–103.
^ Cockell, C. S.; Stokes, M. D. (2004). "Widespread colonization by polar hypoliths". Natuure. 431 (7007): 414–414. doi:10.1038/431414a.
^ Cowan, D. A.; Khan, N.; Pointing, S. B.; Cary, S. C. (2010). "Diverse hypolithic refuge communities in the McMurdo Dry Valleys". Antarctic Science. 22 (6): 714–720. doi:10.1017/S0954102010000507.
^ Friedmann, E. I. "Endolithic microbial life in hot and cold deserts". Orig. Life. 10: 223–235. doi:10.1007/978-94-009-9085-2_3.
^ Smith, M. C.; Bowman, J. P.; Scott, F. J.; Line, M. A. (2000). "Sublithic bacteria associated with Antarctic quartz stones". Antarctic Science. 12 (2): 177–184. doi:10.1017/S0954102000000237.
^ Omelon, C. R.; Pollard, W. H.; Ferris, F. G. (2006). "Environmental controls on microbial colonization of high Arctic cryptoendolithic habitats". Polar Biology. 30 (1): 19–29. doi:10.1007/s00300-006-0155-0.
^ Makhalanyane, T. P.; Pointing, S. B.; Cowan, D. A. (2014). "Lithobionts: cryptic and refuge niches". Antarctic Terrestrial Microbiology: 163–179. doi:10.1007/978-3-642-45213-0_9.
^ 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". Biofouling. 33 (2): 113–127. doi:10.1080/08927014.2016.1269893.
^ Schultz, Steven (13 December 1999). "Two miles underground". Princeton Weekly Bulletin. Archived from the original on 13 January 2016. — Gold mines present "ideal environment" for geologists studying subsurface microbes
^ ^a ^b Hively, Will (May 1997). "Looking for life in all the wrong places — research on cryptoendoliths". Discover. Retrieved 5 December 2019.
^ de la Torre, J. R.; Goebel, B. M.; Friedmann, E. I.; Pace, N. R. (2003). "Microbial Diversity of Cryptoendolithic Communities from the Mc Murdo Dry Valleys, Antarctica". Applied and Environmental Microbiology. 69 (7): 3858–3867. Bibcode:2003ApEnM..69.3858D. doi:10.1128/AEM.69.7.3858-3867.2003. PMC 165166. PMID 12839754.
^ Horath, Thomas; Bachofen, Reinhard (August 2009). "Molecular Characterization of an Endolithic Microbial Community in Dolomite Rock in the Central Alps (Switzerland)" (PDF). Microbial Ecology. 58 (2): 290–306. doi:10.1007/s00248-008-9483-7. PMID 19172216. S2CID 845383.
^ Walker, Jeffrey J.; Spear, John R.; Pace, Norman R. (2005). "Geobiology of a microbial endolithic community in the Yellowstone geothermal environment". Nature. 434 (7036): 1011–1014. Bibcode:2005Natur.434.1011W. doi:10.1038/nature03447. PMID 15846344. S2CID 4408407.
^ Walker, J. J.; Pace, N. R. (2007). "Phylogenetic Composition of Rocky Mountain Endolithic Microbial Ecosystems". Applied and Environmental Microbiology. 73 (11): 3497–3504. Bibcode:2007ApEnM..73.3497W. doi:10.1128/AEM.02656-06. PMC 1932665. PMID 17416689.
^ Mullen, Leslie. "Glass Munchers Under the Sea". NASA Astrobiology Institute. Archived from the original on 20 February 2013.
^ Lysnes, Kristine; Torsvik, Terje; Thorseth, Ingunn H.; Pedersen, Rolf B. (2004). "Microbial Populations in Ocean Floor Basalt: Results from ODP Leg 187" (PDF). Proc ODP Sci Results. Proceedings of the Ocean Drilling Program. 187: 1–27. doi:10.2973/odp.proc.sr.187.203.2004.
^ Wierzchos, Jacek; Ascaso, Carmen; McKay, Christopher P. (2006). "Endolithic Cyanobacteria in Halite Rocks from the Hyperarid Core of the Atacama Desert". Astrobiology. 6 (3): 415–422. Bibcode:2006AsBio...6..415W. doi:10.1089/ast.2006.6.415. PMID 16805697.
^ Yirka, Bob (29 August 2013). "Soil beneath ocean found to harbor long-lived bacteria, fungi and viruses". Phys.org. Archived from the original on 29 October 2015.
^ "Frequently Requested Information about the SLiME Hypothesis". Archived from the original on 30 September 2006.
^ Keith, DA; Iliffe, TM; Gerovasileiou, V; Gonzalez, B; Brankovits, D; Martínez García, A (2020). "S1.2 Endolithic systems". In Keith, D.A.; Ferrer-Paris, J.R.; Nicholson, E.; Kingsford, R.T. (eds.). The IUCN Global Ecosystem Typology 2.0: Descriptive profiles for biomes and ecosystem functional groups. Gland, Switzerland: IUCN. doi:10.2305/IUCN.CH.2020.13.en. ISBN 978-2-8317-2077-7.
^ Golubic, Stjepko; Radtke, Gudrun; Campion-Alsumard, Therese Le (2005). "Endolithic fungi in marine ecosystems". Trends in Microbiology. 13 (5): 229–235. doi:10.1016/j.tim.2005.03.007. PMID 15866040.
^ Fine, Maoz; Loya, Yossi (2002). "Endolithic algae: an alternative source of photoassimilates during coral bleaching". Proceedings of the Royal Society of London. Series B: Biological Sciences. 269 (1497): 1205–1210. doi:10.1098/rspb.2002.1983. PMC 1691023. PMID 12065035.
^ Günther, Astrid (1990). "Distribution and bathymetric zonation of shell-boring endoliths in recent reef and shelf environments: Cozumel, Yucatan (Mexico)". Facies. 22 (1): 233–261. doi:10.1007/bf02536953. S2CID 130403994.
^ Kaehler, S.; McQuaid, C. D. (1999). "Lethal and sub-lethal effects of phototrophic endoliths attacking the shell of the intertidal mussel Perna perna". Marine Biology. 135 (3): 497–503. doi:10.1007/s002270050650. S2CID 84103549.
^ ^a ^b Gong, YiMing; Xu, Ran; Hu, Bi (2008). "Endolithic fungi: A possible killer for the mass extinction of Cretaceous dinosaurs". Science in China Series D: Earth Sciences. 51 (6): 801–807. Bibcode:2008ScChD..51..801G. doi:10.1007/s11430-008-0052-1. S2CID 126670640.

External links

Endoliths General Collection — This collection of online resources such as news articles, web sites, and reference pages provides a comprehensive array of information about endoliths.
Endolith Advanced Collection — Compiled for professionals and advanced learners, this endolith collection includes online resources such as journal articles, academic reviews, and surveys.

[1] Omelon, CR (2016). "Endolithic Microorganisms and Their Habitats". In Hurst, C.J. (ed.). Advances in environmental microbiology. Their World: A Diversity of Microbial Environments. Cincinnati, USA: Springer.

[2] 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". Geomicrobiology Journal. 26 (7): 491–507. doi:10.1080/01490450903061101.

[3] Herrera, A.; Cockell, C. S.; Self, S.; Blaxter, M.; Reitner, J.; Thorsteinsson, T.; Tindle, A. G. (2009). "A cryptoendolithic community in volcanic glass". Astrobiology. 9 (4): 369–381. doi:10.1089/ast.2008.0278.

[4] 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". Microbial ecology. 62 (1): 69–79. doi:10.1007/s00248-011-9864-1.

[5] Omelon, C. R.; Pollard, W. H.; Ferris, F. G. (2007). "Inorganic species distribution and microbial diversity within high Arctic cryptoendolithic habitats". Microbial ecology. 54 (4): 740–752. doi:10.1007/s00248-007-9235-0.

[6] Walker, J. J.; Pace, N. R. (2007). "Endolithic microbial ecosystems". Annu Rev Microbiol. 61: 331–347.

[Wierzchos_et_al._2011-7] 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". Geobiology. 9 (1): 44–60. doi:10.1111/j.1472-4669.2010.00254.x. PMID 20726901.

[NYT-20160912-8] Chang, Kenneth (12 September 2016). "Visions of Life on Mars in Earth's Depths". The New York Times. Retrieved 12 September 2016.

[9] Golubic, Stjepko; Friedmann, E. Imre; Schneider, Jürgen (June 1981). "The lithobiotic ecological niche, with special reference to microorganisms". SEPM Journal of Sedimentary Research. 51 (2): 475–478. doi:10.1306/212F7CB6-2B24-11D7-8648000102C1865D. Archived from the original on 30 December 2010.

[10] Ascaso, C (2002). "Ecología microbiana de sustratos líticos". Ciencia y medio ambiente: 90–103.

[11] Bungartz, F; Garvie, L. A.; Nash, T. H. "Anatomy of the endolithic Sonoran Desert lichen Verrucaria rubrocincta Breuss: implications for biodeterioration and biomineralization". The Lichenologist. 36 (1): 55–73. doi:10.1017/S0024282904013854.

[12] Dong, H; Rech, J. A.; Jiang, H; Sun, H; Buck, B. J. (2007). "Endolithic cyanobacteria in soil gypsum: Occurrences in Atacama (Chile), Mojave (United States), and Al-Jafr Basin (Jordan) Deserts". Journal of Geophysical Research: Biogeosciences. 112: G2. doi:10.1029/2006JG000385.

[13] Lacap, D. C.; Warren-Rhodes, K. A.; McKay, C. P.; Pointing, S. B. (2011). "Cyanobacteria and chloroflexi-dominated hypolithic colonization of quartz at the hyper-arid core of the Atacama Desert, Chile". Extremophiles. 15 (1): 31–38. doi:10.1007/s00792-010-0334-3.

[14] 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". Ecology. 84 (12): 3222–3231. doi:10.1890/02-0549.

[15] 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". Extremophiles. 17 (2): 329–337. doi:10.1007/s00792-.

[16] Vítek, 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". Analytical and bioanalytical chemistry. 408 (15): 4083–4092. doi:10.1007/s00216-016-9497-9.

[17] Ascaso, C (2002). "Ecología microbiana de sustratos líticos". Ciencia y medio ambiente: 90–103.

[18] Cockell, C. S.; Stokes, M. D. (2004). "Widespread colonization by polar hypoliths". Natuure. 431 (7007): 414–414. doi:10.1038/431414a.

[19] Cowan, D. A.; Khan, N.; Pointing, S. B.; Cary, S. C. (2010). "Diverse hypolithic refuge communities in the McMurdo Dry Valleys". Antarctic Science. 22 (6): 714–720. doi:10.1017/S0954102010000507.

[20] Friedmann, E. I. "Endolithic microbial life in hot and cold deserts". Orig. Life. 10: 223–235. doi:10.1007/978-94-009-9085-2_3.

[21] Smith, M. C.; Bowman, J. P.; Scott, F. J.; Line, M. A. (2000). "Sublithic bacteria associated with Antarctic quartz stones". Antarctic Science. 12 (2): 177–184. doi:10.1017/S0954102000000237.

[22] Omelon, C. R.; Pollard, W. H.; Ferris, F. G. (2006). "Environmental controls on microbial colonization of high Arctic cryptoendolithic habitats". Polar Biology. 30 (1): 19–29. doi:10.1007/s00300-006-0155-0.

[23] Makhalanyane, T. P.; Pointing, S. B.; Cowan, D. A. (2014). "Lithobionts: cryptic and refuge niches". Antarctic Terrestrial Microbiology: 163–179. doi:10.1007/978-3-642-45213-0_9.

[24] 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". Biofouling. 33 (2): 113–127. doi:10.1080/08927014.2016.1269893.

[two_miles_underground-25] Schultz, Steven (13 December 1999). "Two miles underground". Princeton Weekly Bulletin. Archived from the original on 13 January 2016. — Gold mines present "ideal environment" for geologists studying subsurface microbes

[hively-26] Hively, Will (May 1997). "Looking for life in all the wrong places — research on cryptoendoliths". Discover. Retrieved 5 December 2019.

[dry_valleys-27] Torre, J. R.; Goebel, B. M.; Friedmann, E. I.; Pace, N. R. (2003). "Microbial Diversity of Cryptoendolithic Communities from the Mc Murdo Dry Valleys, Antarctica". Applied and Environmental Microbiology. 69 (7): 3858–3867. Bibcode:2003ApEnM..69.3858D. doi:10.1128/AEM.69.7.3858-3867.2003. PMC 165166. PMID 12839754.

[28] Horath, Thomas; Bachofen, Reinhard (August 2009). "Molecular Characterization of an Endolithic Microbial Community in Dolomite Rock in the Central Alps (Switzerland)" (PDF). Microbial Ecology. 58 (2): 290–306. doi:10.1007/s00248-008-9483-7. PMID 19172216. S2CID 845383.

[29] Walker, Jeffrey J.; Spear, John R.; Pace, Norman R. (2005). "Geobiology of a microbial endolithic community in the Yellowstone geothermal environment". Nature. 434 (7036): 1011–1014. Bibcode:2005Natur.434.1011W. doi:10.1038/nature03447. PMID 15846344. S2CID 4408407.

[30] Walker, J. J.; Pace, N. R. (2007). "Phylogenetic Composition of Rocky Mountain Endolithic Microbial Ecosystems". Applied and Environmental Microbiology. 73 (11): 3497–3504. Bibcode:2007ApEnM..73.3497W. doi:10.1128/AEM.02656-06. PMC 1932665. PMID 17416689.

[glass_munchers-31] Mullen, Leslie. "Glass Munchers Under the Sea". NASA Astrobiology Institute. Archived from the original on 20 February 2013.

[basalt-32] Lysnes, Kristine; Torsvik, Terje; Thorseth, Ingunn H.; Pedersen, Rolf B. (2004). "Microbial Populations in Ocean Floor Basalt: Results from ODP Leg 187" (PDF). Proc ODP Sci Results. Proceedings of the Ocean Drilling Program. 187: 1–27. doi:10.2973/odp.proc.sr.187.203.2004.

[photendolith-33] Wierzchos, Jacek; Ascaso, Carmen; McKay, Christopher P. (2006). "Endolithic Cyanobacteria in Halite Rocks from the Hyperarid Core of the Atacama Desert". Astrobiology. 6 (3): 415–422. Bibcode:2006AsBio...6..415W. doi:10.1089/ast.2006.6.415. PMID 16805697.

[34] Yirka, Bob (29 August 2013). "Soil beneath ocean found to harbor long-lived bacteria, fungi and viruses". Phys.org. Archived from the original on 29 October 2015.

[slime-35] "Frequently Requested Information about the SLiME Hypothesis". Archived from the original on 30 September 2006.

[36] Keith, DA; Iliffe, TM; Gerovasileiou, V; Gonzalez, B; Brankovits, D; Martínez García, A (2020). "S1.2 Endolithic systems". In Keith, D.A.; Ferrer-Paris, J.R.; Nicholson, E.; Kingsford, R.T. (eds.). The IUCN Global Ecosystem Typology 2.0: Descriptive profiles for biomes and ecosystem functional groups. Gland, Switzerland: IUCN. doi:10.2305/IUCN.CH.2020.13.en. ISBN 978-2-8317-2077-7.

[37] Golubic, Stjepko; Radtke, Gudrun; Campion-Alsumard, Therese Le (2005). "Endolithic fungi in marine ecosystems". Trends in Microbiology. 13 (5): 229–235. doi:10.1016/j.tim.2005.03.007. PMID 15866040.

[38] Fine, Maoz; Loya, Yossi (2002). "Endolithic algae: an alternative source of photoassimilates during coral bleaching". Proceedings of the Royal Society of London. Series B: Biological Sciences. 269 (1497): 1205–1210. doi:10.1098/rspb.2002.1983. PMC 1691023. PMID 12065035.

[39] Günther, Astrid (1990). "Distribution and bathymetric zonation of shell-boring endoliths in recent reef and shelf environments: Cozumel, Yucatan (Mexico)". Facies. 22 (1): 233–261. doi:10.1007/bf02536953. S2CID 130403994.

[40] Kaehler, S.; McQuaid, C. D. (1999). "Lethal and sub-lethal effects of phototrophic endoliths attacking the shell of the intertidal mussel Perna perna". Marine Biology. 135 (3): 497–503. doi:10.1007/s002270050650. S2CID 84103549.

[Gong-41] Gong, YiMing; Xu, Ran; Hu, Bi (2008). "Endolithic fungi: A possible killer for the mass extinction of Cretaceous dinosaurs". Science in China Series D: Earth Sciences. 51 (6): 801–807. Bibcode:2008ScChD..51..801G. doi:10.1007/s11430-008-0052-1. S2CID 126670640.

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 ==Environment==
-Endolithic microorganisms have been reported in many areas around the globe, there are reports in warm hyper-arid and arid deserts such as Monjave and Sonora (E.U.), Atacama (Chile), Gobi (China, Mongolia), Negev (Israel), Namib (Namibia Angola), Al-Jafr basin (Jordania) and the Depression of Turpan (China),<ref>{{cite journal |last1=Ascaso |first1=C |title=Ecología microbiana de sustratos líticos |journal=Ciencia y medio ambiente |date=2002 |pages=90-103 |url=http://hdl.handle.net/10261/111133}}</ref> <ref>{{cite journal |last1=Bungartz |first1=F |last2=Garvie |first2=L. A. |last3=Nash |first3=T. H. |title=Anatomy of the endolithic Sonoran Desert lichen Verrucaria rubrocincta Breuss: implications for biodeterioration and biomineralization |journal=The Lichenologist |volume=36 |issue=1 |pages=55-73 |doi=10.1017/S0024282904013854}}</ref> <ref>{{cite journal |last1=Dong |first1=H |last2=Rech |first2=J. A. |last3=Jiang |first3=H |last4=Sun |first4=H |last5=Buck |first5=B. J. |title=Endolithic cyanobacteria in soil gypsum: Occurrences in Atacama (Chile), Mojave (United States), and Al-Jafr Basin (Jordan) Deserts |journal=Journal of Geophysical Research: Biogeosciences |date=2007 |volume=112 |pages=G2 |doi=10.1029/2006JG000385}}</ref> <ref>{{cite journal |last1=Lacap |first1=D. C. |last2=Warren-Rhodes |first2=K. A. |last3=McKay |first3=C. P. |last4=Pointing |first4=S. B. |title=Cyanobacteria and chloroflexi-dominated hypolithic colonization of quartz at the hyper-arid core of the Atacama Desert, Chile. |journal=Extremophiles |date=2011 |volume=15 |issue=1 |pages=31-38 |doi=10.1007/s00792-010-0334-3}}</ref> <ref>{{cite journal |last1=Schlesinger |first1=W. H |last2=Pippen |first2=J. S. |last3=Wallenstein |first3=M. D. |last4=Hofmockel |first4=K. S. |last5=Klepeis |first5=D. M. |last6=Mahall |first6=B. E. |title=Community composition and photosynthesis by photoautotrophs under quartz pebbles, southern Mojave Desert. |journal=Ecology |date=2003 |volume=84 |issue=12 |pages=3222-3231 |doi=10.1890/02-0549}}</ref> <ref>{{cite journal |last1=Stomeo |first1=F |last2=Valverde |first2=A |last3=Pointing |first3=S. B. |last4=McKay |first4=C. P. |last5=Warren-Rhodes |first5=K. A. |last6=Tuffin |first6=M. I. |last7=Cowan |first7=D. A. |title=Hypolithic and soil microbial community assembly along an aridity gradient in the Namib Desert |journal=Extremophiles |date=2013 |volume=17 |issue=2 |pages=329-337 |doi=10.1007/s00792-}}</ref> <ref>{{cite journal |last1=Vítek |first1=P. |last2=Ascaso |first2=C |last3=Artieda |first3=O |last4=Wierzchos |first4=J |title=Raman imaging in geomicrobiology: endolithic phototrophic microorganisms in gypsum from the extreme sun irradiation area in the Atacama Desert |journal=Analytical and bioanalytical chemistry |date=2016 |volume=408 |issue=15 |pages=4083-4092 |doi=10.1007/s00216-016-9497-9}}</ref>(Ascaso, 2002; Bell, 1993; Bungartz et al., 2004; Dong et al., 2007; Lacap et al., 2011; Schlesinger et al., 2003; Stomeo et al., 2013; Vítek et al., 2016), also in cold deserts as Arctic and Antarctic <ref>{{cite journal |last1=Ascaso |first1=C |title=Ecología microbiana de sustratos líticos |journal=Ciencia y medio ambiente |date=2002 |pages=90-103 |url=http://hdl.handle.net/10261/111133}}</ref> <ref>{{cite journal |last1=Cockell |first1=C. S. |last2=Stokes |first2=M. D. |title=Widespread colonization by polar hypoliths |journal=Natuure |date=2004 |volume=431 |issue=7007 |pages=414-414 |doi=10.1038/431414a}}</ref>(Ascaso 2002; Cockell & Stokes 2004; Cowan et al., 2010; Friedmann 1980; Friedmann et al., 1987; Smith et al., 2000; Omelon et al., 2006; Makhalanyane et al., 2014), and deep subsoil and ocean trenches rocks (Inagaki et al., 2009). However, there are reports of endolithic microorganisms in inter-tropical zones (Gaylarde et al., 2017), where humidity and solar radiation are significantly different from the above-mentioned biomes.
+Endolithic microorganisms have been reported in many areas around the globe, there are reports in warm hyper-arid and arid deserts such as Monjave and Sonora (E.U.), Atacama (Chile), Gobi (China, Mongolia), Negev (Israel), Namib (Namibia Angola), Al-Jafr basin (Jordania) and the Depression of Turpan (China),<ref>{{cite journal |last1=Ascaso |first1=C |title=Ecología microbiana de sustratos líticos |journal=Ciencia y medio ambiente |date=2002 |pages=90-103 |url=http://hdl.handle.net/10261/111133}}</ref> <ref>{{cite journal |last1=Bungartz |first1=F |last2=Garvie |first2=L. A. |last3=Nash |first3=T. H. |title=Anatomy of the endolithic Sonoran Desert lichen Verrucaria rubrocincta Breuss: implications for biodeterioration and biomineralization |journal=The Lichenologist |volume=36 |issue=1 |pages=55-73 |doi=10.1017/S0024282904013854}}</ref> <ref>{{cite journal |last1=Dong |first1=H |last2=Rech |first2=J. A. |last3=Jiang |first3=H |last4=Sun |first4=H |last5=Buck |first5=B. J. |title=Endolithic cyanobacteria in soil gypsum: Occurrences in Atacama (Chile), Mojave (United States), and Al-Jafr Basin (Jordan) Deserts |journal=Journal of Geophysical Research: Biogeosciences |date=2007 |volume=112 |pages=G2 |doi=10.1029/2006JG000385}}</ref> <ref>{{cite journal |last1=Lacap |first1=D. C. |last2=Warren-Rhodes |first2=K. A. |last3=McKay |first3=C. P. |last4=Pointing |first4=S. B. |title=Cyanobacteria and chloroflexi-dominated hypolithic colonization of quartz at the hyper-arid core of the Atacama Desert, Chile. |journal=Extremophiles |date=2011 |volume=15 |issue=1 |pages=31-38 |doi=10.1007/s00792-010-0334-3}}</ref> <ref>{{cite journal |last1=Schlesinger |first1=W. H |last2=Pippen |first2=J. S. |last3=Wallenstein |first3=M. D. |last4=Hofmockel |first4=K. S. |last5=Klepeis |first5=D. M. |last6=Mahall |first6=B. E. |title=Community composition and photosynthesis by photoautotrophs under quartz pebbles, southern Mojave Desert. |journal=Ecology |date=2003 |volume=84 |issue=12 |pages=3222-3231 |doi=10.1890/02-0549}}</ref> <ref>{{cite journal |last1=Stomeo |first1=F |last2=Valverde |first2=A |last3=Pointing |first3=S. B. |last4=McKay |first4=C. P. |last5=Warren-Rhodes |first5=K. A. |last6=Tuffin |first6=M. I. |last7=Cowan |first7=D. A. |title=Hypolithic and soil microbial community assembly along an aridity gradient in the Namib Desert |journal=Extremophiles |date=2013 |volume=17 |issue=2 |pages=329-337 |doi=10.1007/s00792-}}</ref> <ref>{{cite journal |last1=Vítek |first1=P. |last2=Ascaso |first2=C |last3=Artieda |first3=O |last4=Wierzchos |first4=J |title=Raman imaging in geomicrobiology: endolithic phototrophic microorganisms in gypsum from the extreme sun irradiation area in the Atacama Desert |journal=Analytical and bioanalytical chemistry |date=2016 |volume=408 |issue=15 |pages=4083-4092 |doi=10.1007/s00216-016-9497-9}}</ref>(Bell, 1993), also in cold deserts as Arctic and Antarctic <ref>{{cite journal |last1=Ascaso |first1=C |title=Ecología microbiana de sustratos líticos |journal=Ciencia y medio ambiente |date=2002 |pages=90-103 |url=http://hdl.handle.net/10261/111133}}</ref> <ref>{{cite journal |last1=Cockell |first1=C. S. |last2=Stokes |first2=M. D. |title=Widespread colonization by polar hypoliths |journal=Natuure |date=2004 |volume=431 |issue=7007 |pages=414-414 |doi=10.1038/431414a}}</ref> <ref>{{cite journal |last1=Cowan |first1=D. A. |last2=Khan |first2=N. |last3=Pointing |first3=S. B. |last4=Cary |first4=S. C. |title=Diverse hypolithic refuge communities in the McMurdo Dry Valleys |journal=Antarctic Science |date=2010 |volume=22 |issue=6 |pages=714-720 |doi=10.1017/S0954102010000507}}</ref> <ref>{{cite journal |last1=Friedmann |first1=E. I. |title=Endolithic microbial life in hot and cold deserts |journal=Orig. Life |volume=10 |pages=223–235 |doi=10.1007/978-94-009-9085-2_3}}</ref> <ref>{{cite journal |last1=Smith |first1=M. C. |last2=Bowman |first2=J. P. |last3=Scott |first3=F. J. |last4=Line |first4=M. A. |title=Sublithic bacteria associated with Antarctic quartz stones |journal=Antarctic Science |date=2000 |volume=12 |issue=2 |pages=177-184 |doi=10.1017/S0954102000000237}}</ref> <ref>{{cite journal |last1=Omelon |first1=C. R. |last2=Pollard |first2=W. H. |last3=Ferris |first3=F. G. |title=Environmental controls on microbial colonization of high Arctic cryptoendolithic habitats |journal=Polar Biology |date=2006 |volume=30 |issue=1 |pages=19-29 |doi=10.1007/s00300-006-0155-0}}</ref> <ref>{{cite journal |last1=Makhalanyane |first1=T. P. |last2=Pointing |first2=S. B. |last3=Cowan |first3=D. A. |title=Lithobionts: cryptic and refuge niches |journal=Antarctic Terrestrial Microbiology |date=2014 |pages=163-179 |doi=10.1007/978-3-642-45213-0_9}}</ref>(Friedmann et al., 1987; Smith et al., 2000; Omelon et al., 2006; Makhalanyane et al., 2014), and deep subsoil and ocean trenches rocks (Inagaki et al., 2009). However, there are reports of endolithic microorganisms in inter-tropical zones <ref>{{cite journal |last1=Gaylarde |first1=C. |last2=Baptista-Neto |first2=J. A. |last3=Ogawa |first3=A. |last4=Kowalski |first4=M. |last5=Celikkol-Aydin |first5=S. |last6=Beech |first6=I. |title=Epilithic and endolithic microorganisms and deterioration on stone church facades subject to urban pollution in a sub-tropical climate |journal=Biofouling |date=2017 |volume=33 |issue=2 |pages=113-127 |doi=10.1080/08927014.2016.1269893}}</ref>, where humidity and solar radiation are significantly different from the above-mentioned biomes.
 Endoliths have been found in rock down to a depth of {{convert|3|km}}, though it is unknown if that is their limit (due to the cost involved in digging so deeply).<ref name="two miles underground">{{cite web|last1=Schultz|first1=Steven|title=Two miles underground|url=http://www.princeton.edu/pr/pwb/99/1213/microbe.shtml|publisher=Princeton Weekly Bulletin|archive-url=https://web.archive.org/web/20160113130655/http://www.princeton.edu/pr/pwb/99/1213/microbe.shtml|archive-date=13 January 2016|date=13 December 1999}} — Gold mines present "ideal environment" for geologists studying subsurface microbes</ref><ref name="hively">{{cite news|url=http://discovermagazine.com/1997/may/lookingforlifein1124|title= Looking for life in all the wrong places — research on cryptoendoliths|work=[[Discover (magazine)|Discover]]|date= May 1997 |first1= Will|last1= Hively|access-date=December 5, 2019}}</ref> The main threat to their survival seems not to result from the pressure at such depth, but from the increased temperature. Judging from [[hyperthermophile]] organisms, the temperature limit is at about 120&nbsp;°C ([[Strain 121]] can reproduce at 121&nbsp;°C), which limits the possible depth to 4-4.5&nbsp;km below the [[continent]]al crust, and 7 or 7.5&nbsp;km below the [[ocean]] floor. Endolithic organisms have also been found in surface rocks in regions of  low humidity ([[hypolith]]) and low temperature ([[psychrophile]]), including the [[Dry Valleys]] and [[permafrost]] of [[Antarctica]],<ref name="dry valleys">{{Cite journal |doi = 10.1128/AEM.69.7.3858-3867.2003|pmid = 12839754|pmc = 165166|title = Microbial Diversity of Cryptoendolithic Communities from the Mc ''Murdo'' Dry Valleys, Antarctica|journal = Applied and Environmental Microbiology|volume = 69|issue = 7|pages = 3858–3867|year = 2003|last1 = de la Torre|first1 = J. R.|last2 = Goebel|first2 = B. M.|last3 = Friedmann|first3 = E. I.|last4 = Pace|first4 = N. R.|bibcode = 2003ApEnM..69.3858D}}</ref> the [[Alps]],<ref>{{cite journal|last1=Horath|first1=Thomas|last2=Bachofen|first2=Reinhard|title=Molecular Characterization of an Endolithic Microbial Community in Dolomite Rock in the Central Alps (Switzerland)|journal=Microbial Ecology|date=August 2009|volume=58|issue=2|pages=290–306|doi=10.1007/s00248-008-9483-7|pmid=19172216|s2cid=845383|url=https://www.zora.uzh.ch/id/eprint/18300/34/ZORA_NL_18300.pdf}}</ref> and the [[Rocky Mountains]].<ref>{{Cite journal | doi=10.1038/nature03447| pmid=15846344| title=Geobiology of a microbial endolithic community in the Yellowstone geothermal environment| journal=Nature| volume=434| issue=7036| pages=1011–1014| year=2005| last1=Walker| first1=Jeffrey J.| last2=Spear| first2=John R.| last3=Pace| first3=Norman R.| bibcode=2005Natur.434.1011W| s2cid=4408407}}</ref><ref>{{Cite journal |doi = 10.1128/AEM.02656-06|pmid = 17416689|pmc = 1932665|title = Phylogenetic Composition of Rocky Mountain Endolithic Microbial Ecosystems|journal = Applied and Environmental Microbiology|volume = 73|issue = 11|pages = 3497–3504|year = 2007|last1 = Walker|first1 = J. J.|last2 = Pace|first2 = N. R.|bibcode = 2007ApEnM..73.3497W}}</ref>