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The bright colors of Grand Prismatic Spring, Yellowstone National Park, are produced by Thermophiles, a type of extremophile. Grand prismatic spring.jpg
The bright colors of Grand Prismatic Spring, Yellowstone National Park, are produced by Thermophiles, a type of extremophile.

An extremophile (from Latin extremus meaning "extreme" and Greek philiā (φιλία) meaning "love") is an organism with optimal growth in environmental conditions considered extreme in comparison to the environmental conditions that are comfortable to humans. [1] [2] [3] In contrast, organisms that live in more moderate environmental conditions, according to an anthropocentric view, may be termed mesophiles or neutrophiles.

Organism Any individual living physical entity

In biology, an organism is any individual entity that propagates the properties of life. It is a synonym for "life form".

An extreme environment is a habitat that is considered very hard to survive in due to its considerably extreme conditions such as temperature, accessibility to different energy sources or under high pressure. For an area to be considered an extreme environment, it must contain certain conditions and aspects that are considered very hard for other life forms to survive. Pressure conditions may be extremely high or low; high or low content of oxygen or carbon dioxide in the atmosphere; high levels of radiation, acidity, or alkalinity; absence of water; water containing a high concentration of salt or sugar; the presence of sulphur, petroleum, and other toxic substances.

Human Hominin events for the last 10 million years

Humans are the only extant members of the subtribe Hominina. Together with chimpanzees, gorillas, and orangutans, they are part of the family Hominidae. A terrestrial animal, humans are characterized by their erect posture and bipedal locomotion; high manual dexterity and heavy tool use compared to other animals; open-ended and complex language use compared to other animal communications; larger, more complex brains than other animals; and highly advanced and organized societies.



In the 1980s and 1990s, biologists found that microbial life has great flexibility for surviving in extreme environments—niches that are acidic or extraordinarily hot, for example—that would be completely inhospitable to complex organisms. Some scientists even concluded that life may have begun on Earth in hydrothermal vents far under the ocean's surface. [4]

Hydrothermal vent A fissure in a planets surface from which geothermally heated water issues

A hydrothermal vent is a fissure on the seafloor from which geothermally heated water issues. Hydrothermal vents are commonly found near volcanically active places, areas where tectonic plates are moving apart at spreading centers, ocean basins, and hotspots. Hydrothermal deposits are rocks and mineral ore deposits formed by the action of hydrothermal vents.

According to astrophysicist Steinn Sigurdsson, "There are viable bacterial spores that have been found that are 40 million years old on Earth—and we know they're very hardened to radiation." [5] Some bacteria were found living in the cold and dark in a lake buried a half-mile deep under the ice in Antarctica, [6] and in the Marianas Trench, the deepest place in Earth's oceans. [7] [8] Some microorganisms have been found thriving inside rocks up to 1,900 feet (580 m) below the sea floor under 8,500 feet (2,600 m) of ocean off the coast of the northwestern United States. [7] [9] According to one of the researchers, "You can find microbes everywhere—they're extremely adaptable to conditions, and survive wherever they are." [7] A key to extremophile adaptation is their amino acid composition, affecting their protein folding ability under particular conditions. [10]

Endospore protective structure formed by bacteria

An endospore is a dormant, tough, and non-reproductive structure produced by some bacteria in the phylum Firmicutes. The name "endospore" is suggestive of a spore or seed-like form, but it is not a true spore. It is a stripped-down, dormant form to which the bacterium can reduce itself. Endospore formation is usually triggered by a lack of nutrients, and usually occurs in gram-positive bacteria. In endospore formation, the bacterium divides within its cell wall, and one side then engulfs the other. Endospores enable bacteria to lie dormant for extended periods, even centuries. There are many reports of spores remaining viable over 10,000 years, and revival of spores millions of years old has been claimed. There is one report of viable spores of Bacillus marismortui in salt crystals approximately 250 million years old. When the environment becomes more favorable, the endospore can reactivate itself to the vegetative state. Most types of bacteria cannot change to the endospore form. Examples of bacteria that can form endospores include Bacillus and Clostridium.

Radiation Waves or particles propagating through space or through a medium, carrying energy

In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or through a material medium. This includes:

Bacteria A domain of prokaryotes – single celled organisms without a nucleus

Bacteria are a type of biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a number of shapes, ranging from spheres to rods and spirals. Bacteria were among the first life forms to appear on Earth, and are present in most of its habitats. Bacteria inhabit soil, water, acidic hot springs, radioactive waste, and the deep biosphere of the earth's crust. Bacteria also live in symbiotic and parasitic relationships with plants and animals. Most bacteria have not been characterised, and only about 27 percent of the bacterial phyla have species that can be grown in the laboratory . The study of bacteria is known as bacteriology, a branch of microbiology.

Tom Gheysens from Ghent University in Belgium and some of his colleagues have presented research findings that show spores from a species of Bacillus bacteria survived and were still viable after being heated to temperatures of 420 °C (788 °F). [11]

The limits of known life on Earth. [12]
FactorEnvironment / sourceLimitsExamples
High temperature Submarine hydrothermal vents 110 °C to 121 °C Pyrolobus fumarii , Pyrococcus furiosus
Low temperatureIce-17 °C to -20 °C Synechococcus lividus
Alkaline systems Soda lakes pH > 11 Psychrobacter , Vibrio , Arthrobacter , Natronobacterium
Acidic systemsVolcanic springs, acid mine drainagepH -0.06 to 1.0 Bacillus , Clostridium paradoxum
Ionizing radiation Cosmic rays, X-rays, radioactive decay 1,500 to 6,000 Gy Deinococcus radiodurans , Rubrobacter , Thermococcus gammatolerans
UV radiation Sunlight5,000 J/m2Deinococcus radiodurans, Rubrobacter, Thermococcus gammatolerans
High pressure Mariana Trench 1,100 bar Pyrococcus sp.
SalinityHigh salt concentration aw ~ 0.6 Halobacteriaceae , Dunaliella salina
Desiccation Atacama Desert (Chile), McMurdo Dry Valleys (Antarctica)~60% relative humidity Chroococcidiopsis
Deep crustaccessed at some gold mines Halicephalobus mephisto , Mylonchulus brachyurus , unidentified arthropods


There are many classes of extremophiles that range all around the globe; each corresponding to the way its environmental niche differs from mesophilic conditions. These classifications are not exclusive. Many extremophiles fall under multiple categories and are classified as polyextremophiles. For example, organisms living inside hot rocks deep under Earth's surface are thermophilic and barophilic such as Thermococcus barophilus. [13] A polyextremophile living at the summit of a mountain in the Atacama Desert might be a radioresistant xerophile, a psychrophile, and an oligotroph. Polyextremophiles are well known for their ability to tolerate both high and low pH levels. [14]

Atacama Desert desert in South America

The Atacama Desert is a desert plateau in South America covering a 1,000-km (600-mi) strip of land on the Pacific coast, west of the Andes mountains. The Atacama desert is one of the driest places in the world, as well as the only true desert to receive less precipitation than the polar deserts. According to estimates, the Atacama Desert occupies 105,000 km2 (41,000 sq mi), or 128,000 km2 (49,000 sq mi) if the barren lower slopes of the Andes are included. Most of the desert is composed of stony terrain, salt lakes (salares), sand, and felsic lava that flows towards the Andes.

A xerophile is an extremophilic organism that can grow and reproduce in conditions with a low availability of water, also known as water activity. Water activity (aw) is measured as the humidity above a substance relative to the humidity above pure water. Xerophiles are "xerotolerant", meaning tolerant of dry conditions. They often can survive in environments with water activity below 0.8; above which is typical for most life on Earth. Typically xerotolerance is used with respect to matric drying, where a substance has a low water concentration. These environments include arid desert soils. The term osmotolerance is typically applied to organisms that can grow in solutions with high solute concentrations, such as halophiles.

Psychrophiles or cryophiles are extremophilic organisms that are capable of growth and reproduction in low temperatures, ranging from −20 °C to +10 °C. They are found in places that are permanently cold, such as the polar regions and the deep sea. They can be contrasted with thermophiles, which are organisms that thrive at unusually high temperatures. Psychrophile is Greek for 'cold-loving'.


An organism with optimal growth at pH levels of 3.0 or below
An organism with optimal growth at pH levels of 9.0 or above
An organism with optimal growth in the absence of molecular oxygen. Two sub-types exist: facultative anaerobe and obligate anaerobe. A facultative anaerobe can tolerate anoxic and oxic conditions; however, an obligate anaerobe dies in the presence of even trace levels of molecular oxygen.
An organism that lives in microscopic spaces within rocks, such as pores between aggregate grains. These may also be called endolith, a term that also includes organisms populating fissures, aquifers, and faults filled with groundwater in the deep subsurface.
An organism with optimal growth at a concentration of dissolved salts of 50 g/L (= 5% m/v) or above.
An organism with optimal growth at hydrostatic pressures above 50 MPa (= 493 atm = 7,252 psi).
An organism with optimal growth at temperatures above 80 °C (176 °F).
An organism that lives underneath rocks in cold deserts.
Capable of tolerating high levels of dissolved heavy metals in solution, such as copper, cadmium, arsenic, and zinc. Examples include Ferroplasma sp., Cupriavidus metallidurans and GFAJ-1. [15] [16] [17]
An organism with optimal growth in nutritionally limited environments.
An organism with optimal growth in environments with a high sugar concentration.
An organism with optimal growth in hydrostatic pressures above 10 MPa (= 99 atm = 1,450 psi). Also referred to as barophile.
A polyextremophile (faux Ancient Latin/Greek for 'affection for many extremes') is an organism that qualifies as an extremophile under more than one category.
An organism with optimal growth at temperatures of 15 °C (59 °F) or lower.
Organisms resistant to high levels of ionizing radiation, most commonly ultraviolet radiation. This category also includes organisms capable of resisting nuclear radiation.
An organism with optimal growth at temperatures above 45 °C (113 °F).
An organism with optimal growth at water activity below 0.8.

In astrobiology

Astrobiology is the study of the origin, evolution, distribution, and future of life in the universe: extraterrestrial life and life on Earth. Astrobiology makes use of physics, chemistry, astronomy, solar physics, biology, molecular biology, ecology, planetary science, geography, and geology to investigate the possibility of life on other worlds and help recognize biospheres that might be different from that on Earth. [18] Astrobiologists are particularly interested in studying extremophiles, [19] as their habitats may be analogous to conditions on other planets. For example, analogous deserts of Antarctica are exposed to harmful UV radiation, low temperature, high salt concentration and low mineral concentration. These conditions are similar to those on Mars. Therefore, finding viable microbes in the subsurface of Antarctica suggests that there may be microbes surviving in endolithic communities and living under the Martian surface. Research indicates it is unlikely that Martian microbes exist on the surface or at shallow depths, but may be found at subsurface depths of around 100 meters. [20]

Astrobiology Science concerned with life in the universe

Astrobiology, formerly known as exobiology, is an interdisciplinary scientific field concerned with the origins, early evolution, distribution, and future of life in the universe. Astrobiology considers the question of whether extraterrestrial life exists, and if it does, how humans can detect it.

Abiogenesis The natural process by which life arises from non-living matter

Abiogenesis, or informally the origin of life, is the natural process by which life has arisen from non-living matter, such as simple organic compounds. While the details of this process are still unknown, the prevailing scientific hypothesis is that the transition from non-living to living entities was not a single event, but an evolutionary process of increasing complexity that involved molecular self-replication, self-assembly, autocatalysis, and the emergence of cell membranes. Although the occurrence of abiogenesis is not controversial among scientists, its possible mechanisms are poorly understood. This article presents several principles and hypotheses for how abiogenesis could have occurred.

Evolution change in the heritable characteristics of biological populations over successive generations

Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes that are passed on from parent to offspring during reproduction. Different characteristics tend to exist within any given population as a result of mutation, genetic recombination and other sources of genetic variation. Evolution occurs when evolutionary processes such as natural selection and genetic drift act on this variation, resulting in certain characteristics becoming more common or rare within a population. It is this process of evolution that has given rise to biodiversity at every level of biological organisation, including the levels of species, individual organisms and molecules.

Recent research carried out on extremophiles in Japan involved a variety of bacteria including Escherichia coli and Paracoccus denitrificans being subject to conditions of extreme gravity. The bacteria were cultivated while being rotated in an ultracentrifuge at high speeds corresponding to 403,627 g (i.e. 403,627 times the gravity experienced on Earth). Paracoccus denitrificans was one of the bacteria which displayed not only survival but also robust cellular growth under these conditions of hyperacceleration which are usually found only in cosmic environments, such as on very massive stars or in the shock waves of supernovas. Analysis showed that the small size of prokaryotic cells is essential for successful growth under hypergravity. The research has implications on the feasibility of panspermia. [21] [22] [23]

On 26 April 2012, scientists reported that lichen survived and showed remarkable results on the adaptation capacity of photosynthetic activity within the simulation time of 34 days under Martian conditions in the Mars Simulation Laboratory (MSL) maintained by the German Aerospace Center (DLR). [24] [25]

On 29 April 2013, scientists at Rensselaer Polytechnic Institute, funded by NASA, reported that, during spaceflight on the International Space Station, microbes seem to adapt to the space environment in ways "not observed on Earth" and in ways that "can lead to increases in growth and virulence". [26]

On 19 May 2014, scientists announced that numerous microbes, like Tersicoccus phoenicis , may be resistant to methods usually used in spacecraft assembly clean rooms. It's not currently known if such resistant microbes could have withstood space travel and are present on the Curiosity rover now on the planet Mars. [27]

On 20 August 2014, scientists confirmed the existence of microorganisms living half a mile below the ice of Antarctica. [28] [29]

On September 2015, scientists from CNR-National Research Council of Italy reported that S.soflataricus was able to survive under Martian radiation at a wavelength that was considered extremely lethal to most bacteria. This discovery is significant because it indicates that not only bacterial spores, but also growing cells can be remarkably resistant to strong UV radiation. [30]

On June 2016, scientists from Brigham Young University conclusively reported that endospores of Bacillus subtilis were able to survive high speed impacts up to 299±28 m/s, extreme shock, and extreme deceleration. They pointed out that this feature might allow endospores to survive and to be transferred between planets by traveling within meteorites or by experiencing atmosphere disruption. Moreover, they suggested that the landing of spacecraft may also result in interplanetary spore transfer, given that spores can survive high-velocity impact while ejected from the spacecraft onto the planet surface. This is the first study which reported that bacteria can survive in such high-velocity impact. However, the lethal impact speed is unknown, and further experiments should be done by introducing higher-velocity impact to bacterial endospores. [31]

Examples and recent findings

New sub-types of -philes are identified frequently and the sub-category list for extremophiles is always growing. For example, microbial life lives in the liquid asphalt lake, Pitch Lake. Research indicates that extremophiles inhabit the asphalt lake in populations ranging between 106 to 107 cells/gram. [32] [33] Likewise, until recently boron tolerance was unknown but a strong borophile was discovered in bacteria. With the recent isolation of Bacillus boroniphilus, borophiles came into discussion. [34] Studying these borophiles may help illuminate the mechanisms of both boron toxicity and boron deficiency.

In July 2019, a scientific study of Kidd Mine in Canada discovered sulfur-breathing organisms which live 7900 feet below the surface, and which breathe sulfur in order to survive. these organisms are also remarkable due to eating rocks such as pyrite as their regular food source. [35] [36] [37]


The thermoalkaliphilic catalase, which initiates the breakdown of hydrogen peroxide into oxygen and water, was isolated from an organism, Thermus brockianus , found in Yellowstone National Park by Idaho National Laboratory researchers. The catalase operates over a temperature range from 30 °C to over 94 °C and a pH range from 6–10. This catalase is extremely stable compared to other catalases at high temperatures and pH. In a comparative study, the T. brockianus catalase exhibited a half life of 15 days at 80 °C and pH 10 while a catalase derived from Aspergillus niger had a half life of 15 seconds under the same conditions. The catalase will have applications for removal of hydrogen peroxide in industrial processes such as pulp and paper bleaching, textile bleaching, food pasteurization, and surface decontamination of food packaging. [38]

DNA modifying enzymes such as Taq DNA polymerase and some Bacillus enzymes used in clinical diagnostics and starch liquefaction are produced commercially by several biotechnology companies. [39]

DNA transfer

Over 65 prokaryotic species are known to be naturally competent for genetic transformation, the ability to transfer DNA from one cell to another cell followed by integration of the donor DNA into the recipient cell's chromosome. [40] Several extremophiles are able to carry out species-specific DNA transfer, as described below. However, it is not yet clear how common such a capability is among extremophiles.

The bacterium Deinococcus radiodurans is one of the most radioresistant organisms known. This bacterium can also survive cold, dehydration, vacuum and acid and is thus known as a polyextremophile. D. radiodurans is competent to perform genetic transformation. [41] Recipient cells are able to repair DNA damage in donor transforming DNA that had been UV irradiated as efficiently as they repair cellular DNA when the cells themselves are irradiated. The extreme thermophilic bacterium Thermus thermophilus and other related Thermus species are also capable of genetic transformation. [42]

Halobacterium volcanii , an extreme halophilic (saline tolerant) archaeon, is capable of natural genetic transformation. Cytoplasmic bridges are formed between cells that appear to be used for DNA transfer from one cell to another in either direction. [43]

Sulfolobus solfataricus and Sulfolobus acidocaldarius are hyperthermophilic archaea. Exposure of these organisms to the DNA damaging agents UV irradiation, bleomycin or mitomycin C induces species-specific cellular aggregation. [44] [45] UV-induced cellular aggregation of S. acidocaldarius mediates chromosomal marker exchange with high frequency. [45] Recombination rates exceed those of uninduced cultures by up to three orders of magnitude. Frols et al. [44] and Ajon et al. [45] hypothesized that cellular aggregation enhances species-specific DNA transfer between Sulfolobus cells in order to repair damaged DNA by means of homologous recombination. Van Wolferen et al. [46] noted that this DNA exchange process may be crucial under DNA damaging conditions such as high temperatures. It has also been suggested that DNA transfer in Sulfolobus may be an early form of sexual interaction similar to the more well-studied bacterial transformation systems that involve species-specific DNA transfer leading to homologous recombinational repair of DNA damage [47] (and see Transformation (genetics)).

Extracellular membrane vesicles (MVs) might be involved in DNA transfer between different hyperthermophilic archaeal species. [48] It has been shown that both plasmids [49] and viral genomes [48] can be transferred via MVs. Notably, a horizontal plasmid transfer has been documented between hyperthermophilic Thermococcus and Methanocaldococcus species, respectively belonging to the orders Thermococcales and Methanococcales. [50]

See also

Specific types of organisms

Related Research Articles

Microorganism Microscopic living organism

A microorganism, or microbe, is a microscopic organism, which may exist in its single-celled form or in a colony of cells.

Panspermia Hypothesis that life exists throughout the Universe, distributed by space dust, meteoroids, asteroids, comets, planetoids, and also by spacecraft carrying unintended contamination by microorganisms

Panspermia is the hypothesis that life exists throughout the Universe, distributed by space dust, meteoroids, asteroids, comets, planetoids, and also by spacecraft carrying unintended contamination by microorganisms. Distribution may have occurred spanning galaxies, and so may not be restricted to the limited scale of solar systems.

Thermophile organism that thrives are relatively high temperatures

A thermophile is an organism—a type of extremophile—that thrives at relatively high temperatures, between 41 and 122 °C . Many thermophiles are archaea. Thermophilic eubacteria are suggested to have been among the earliest bacteria.

Bacterial growth Growth of bacterial colonies

Bacterial growth is the asexual reproduction, or cell division, of a bacterium into two daughter cells, in a process called binary fission. Providing no event occurs, the resulting daughter cells are genetically identical to the original cell. Hence, bacterial growth occurs. Both daughter cells from the division do not necessarily survive. However, if the number surviving exceeds unity on average, the bacterial population undergoes exponential growth. The measurement of an exponential bacterial growth curve in batch culture was traditionally a part of the training of all microbiologists; the basic means requires bacterial enumeration by direct and individual, direct and bulk (biomass), indirect and individual, or indirect and bulk methods. Models reconcile theory with the measurements.

Unicellular organism Organism that consists of only one cell

A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of Multiple cells. Unicellular organisms fall into two general categories: prokaryotic organisms and eukaryotic organisms. Prokaryotes include bacteria and archaea. Many eukaryotes are multicellular, but the group includes the protozoa, unicellular algae, and unicellular fungi. Unicellular organisms are thought to be the oldest form of life, with early protocells possibly emerging 3.8–4 billion years ago.

A mesophile is an organism that grows best in moderate temperature, neither too hot nor too cold, typically between 14 and 21 °C. The term is mainly applied to microorganisms. Organisms that prefer extreme environments are known as extremophiles. Mesophiles have diverse classifications, belonging to two domains: Bacteria, Archaea, and to kingdom Fungi of domain Eucarya. Mesophiles belonging to the domain Bacteria can either be gram-positive or gram-negative. Gram-positive bacteria have a cell layer made of peptidoglycan and stains purple. Gram-negative bacteria also contains peptidoglycan, yet the layer is extremely thin and stains red or pink. Oxygen requirements for mesophiles are not just confined to aerobic or anaerobic. There are three basic shapes of mesophiles: coccus, bacillus, and spiral.

A hyperthermophile is an organism that thrives in extremely hot environments—from 60 °C upwards. An optimal temperature for the existence of hyperthermophiles is above 80 °C. Hyperthermophiles are often within the domain Archaea, although some bacteria are able to tolerate temperatures of around 100 °C, as well. Some bacteria can live at temperatures higher than 100 °C at large depths in sea where water does not boil because of high pressure. Many hyperthermophiles are also able to withstand other environmental extremes such as high acidity or high radiation levels. Hyperthermophiles are a subset of extremophiles.

Karl Otto Stetter is a German microbiologist and authority on astrobiology. He is an expert on microbial life at high temperatures.

Microbial ecology Study of the relationship of microorganisms with their environment

Microbial ecology is the ecology of microorganisms: their relationship with one another and with their environment. It concerns the three major domains of life—Eukaryota, Archaea, and Bacteria—as well as viruses.

Microbial genetics is a subject area within microbiology and genetic engineering. Microbial genetics studies microorganisms for different purposes. The microorganisms that are observed are bacteria, and archaea. Some fungi and protozoa are also subjects used to study in this field. The studies of microorganisms involve studies of genotype and expression system. Genotypes are the inherited compositions of an organism. Genetic Engineering is a field of work and study within microbial genetics. The usage of recombinant DNA technology is a process of this work. The process involves creating recombinant DNA molecules through manipulating a DNA sequence. That DNA created is then in contact with a host organism. Cloning is also an example of genetic engineering.

In taxonomy, Thermococcus is a genus of extreme thermophiles in the family the Thermococcaceae.

Prokaryote Group of organisms whose cells lack a cell nucleus

A prokaryote is a unicellular organism that lacks a membrane-bound nucleus, mitochondria, or any other membrane-bound organelle. The word prokaryote comes from the Greek πρό (pro) "before" and κάρυον (karyon) "nut" or "kernel". Prokaryotes are divided into two domains, Archaea and Bacteria. Species with nuclei and organelles are placed in the third domain, Eukaryota. Prokaryotes reproduce without fusion of gametes. The first living organisms are thought to have been prokaryotes.

Archaea A domain of single-celled prokaryotic microorganisms

Archaea constitute a domain of single-celled organisms. These microorganisms are prokaryotes, and have no cell nucleus. Archaea were initially classified as bacteria, receiving the name archaebacteria, but this classification is outmoded.

Microbes can be damaged or killed by elements of their physical environment such as temperature, radiation, or exposure to chemicals; these effects can be exploited in efforts to control pathogens, often for the purpose of food safety.

Thermoplasma volcanium is a moderate thermoacidophilic archaea isolated from acidic hydrothermal vents and solfatara fields. It contains no cell wall and is motile. It is a facultative anaerobic chemoorganoheterotroph. No previous phylogenetic classifications have been made for this organism. Thermoplasma volcanium reproduces asexually via binary fission and is nonpathogenic.

Microbial biogeography is a subset of biogeography, a field that concerns the distribution of organisms across space and time. Although biogeography traditionally focused on plants and larger animals, recent studies have broadened this field to include distribution patterns of microorganisms. This extension of biogeography to smaller scales—known as "microbial biogeography"—is enabled by ongoing advances in genetic technologies.

Saccharolobus solfataricus is a species of thermophilic archaeon. It was transferred from the genus Sulfolobus to the new genus Saccharolobus with the description of Saccharolobus caldissimus in 2018.

Metallosphaera hakonensis is a gram-negative, thermoacidophilic archaea discovered in the hot springs of Hakone National Park, Kanagawa, Japan.


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Further reading