Temporal range: Cambrian–Recent
Tardigrades (//; also known colloquially as water bears, or moss piglets) are a phylum of water-dwelling, eight-legged, segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who gave them the name of "little water bears". The name Tardigrada (meaning "slow steppers") was given three years later by the Italian biologist Lazzaro Spallanzani. They have been found everywhere: from mountaintops to the deep sea and mud volcanoes; from tropical rain forests to the Antarctic. Tardigrades are among the most resilient known animals, with individual species able to survive extreme conditions that would be rapidly fatal to nearly all other known life forms, such as exposure to extreme temperatures, extreme pressures (both high and low), air deprivation, radiation, dehydration, and starvation. About 1,150 known species form the phylum Tardigrada, a part of the superphylum Ecdysozoa. The group includes fossils dating from 530 million years ago, in the Cambrian period.
Usually, tardigrades are about 0.5 mm (0.02 in) long when they are fully grown. They are short and plump, with four pairs of legs, each ending in claws (usually four to eight) and/or sucking disks. Tardigrades are prevalent in mosses and lichens and feed on plant cells, algae, and small invertebrates. When collected, they may be viewed under a very low-power microscope, making them accessible to students and amateur scientists.
Johann August Ephraim Goeze originally named the tardigrade kleiner Wasserbär (Bärtierchen today), meaning "little water bear" in German. The name Tardigradum means "slow walker" and was given by Lazzaro Spallanzani in 1776. The name "water bear" comes from the way they walk, reminiscent of a bear's gait. The biggest adults may reach a body length of 1.5 mm (0.059 in), the smallest below 0.1 mm. Newly hatched tardigrades may be smaller than 0.05 mm.
The most convenient place to find tardigrades is on lichens and mosses. Other environments are dunes, beaches, soil, and marine or freshwater sediments, where they may occur quite frequently (up to 25,000 animals per litre). Tardigrades, in the case of Echiniscoides wyethi, may be found on barnacles. Often, tardigrades can be found by soaking a piece of moss in water.
Anatomy and morphology
Tardigrades have barrel-shaped bodies with four pairs of stubby legs. Most range from 0.3 to 0.5 mm (0.012 to 0.020 in) in length, although the largest species may reach 1.2 mm (0.047 in). The body consists of a head, three body segments each with a pair of legs, and a caudal segment with a fourth pair of legs. The legs are without joints, while the feet have four to eight claws each. The cuticle contains chitin and protein and is moulted periodically. The first three pairs of legs are directed downward along the sides, and are the primary means of locomotion, while the fourth pair is directed backward on the last segment of the trunk and is used primarily for grasping the substrate.
Tardigrades lack several Hox genes and a large intermediate region of the body axis. In insects, this corresponds to the entire thorax and the abdomen. Practically the whole body, except for the last pair of legs, is made up of just the segments that are homologous to the head region in arthropods.
The body cavity consists of a haemocoel, but the only place where a true coelom can be found is around the gonad. No respiratory organs are found, with gas exchange able to occur across the entirety of the body. Some tardigrades have three tubular glands associated with the rectum; these may be excretory organs similar to the Malpighian tubules of arthropods, although the details remain unclear. Also nephridia are absent.
The tubular mouth is armed with stylets, which are used to pierce the plant cells, algae, or small invertebrates on which the tardigrades feed, releasing the body fluids or cell contents. The mouth opens into a triradiate, muscular, sucking pharynx. The stylets are lost when the animal molts, and a new pair is secreted from a pair of glands that lie on either side of the mouth. The pharynx connects to a short esophagus, and then to an intestine that occupies much of the length of the body, which is the main site of digestion. The intestine opens, via a short rectum, to an anus located at the terminal end of the body. Some species only defecate when they molt, leaving the feces behind with the shed cuticle.
The brain develops in a bilaterally symmetric pattern. The brain includes multiple lobes, mostly consisting of three bilaterally paired clusters of neurons. The brain is attached to a large ganglion below the esophagus, from which a double ventral nerve cord runs the length of the body. The cord possesses one ganglion per segment, each of which produces lateral nerve fibres that run into the limbs. Many species possess a pair of rhabdomeric pigment-cup eyes, and numerous sensory bristles are on the head and body.
Tardigrades all possess a buccopharyngeal apparatus (swallowing device made of muscles and spines that activates an inner jaw and begins digestion and movement along the throat and intestine) which, along with the claws, is used to differentiate among species.
Although some species are parthenogenic, both males and females are usually present, each with a single gonad located above the intestine. Two ducts run from the testes in males, opening through a single pore in front of the anus. In contrast, females have a single duct opening either just above the anus or directly into the rectum, which thus forms a cloaca.
Tardigrades are oviparous, and fertilization is usually external. Mating occurs during the molt with the eggs being laid inside the shed cuticle of the female and then covered with sperm. A few species have internal fertilization, with mating occurring before the female fully sheds her cuticle. In most cases, the eggs are left inside the shed cuticle to develop, but some species attach them to nearby substrate.
The eggs hatch after no more than 14 days, with the young already possessing their full complement of adult cells. Growth to the adult size therefore occurs by enlargement of the individual cells (hypertrophy), rather than by cell division. Tardigrades may molt up to 12 times.
Ecology and life history
This section needs expansion. You can help by adding to it. (November 2014)
Most tardigrades are phytophagous (plant eaters) or bacteriophagous (bacteria eaters), but some are carnivorous to the extent of eating other smaller species of tardigrades (e.g., Milnesium tardigradum).
Tardigrades share morphological characteristics with many species that differ largely by class. Biologists have a difficult time finding verification among tardigrade species because of this relationship. These animals are most closely related to the early evolution of arthropods. Tardigrade fossils go as far back as the Cretaceous period in North America. This specific species is considered cosmopolitan and can be located in regions all over the world. The eggs and cysts of tardigrades are so resistant to other dangers that they are carried great distances, on the feet of other animals, to a different location.
The lifespan of tardigrades range from 3–4 months for some species, up to 2 years for other species, not counting the time they spend in dormant states.
Scientists have reported tardigrades in hot springs, on top of the Himalayas (6,000 m; 20,000 ft, above sea level) to the deep sea (−4,000 m; −13,000 ft) and from the polar regions to the equator, under layers of solid ice, and in ocean sediments. Many species can be found in milder environments such as lakes, ponds, and meadows, while others can be found in stone walls and roofs. Tardigrades are most common in moist environments, but can stay active wherever they can retain at least some moisture.
Tardigrades are considered to be able to survive even complete global mass extinction events due to astrophysical events, such as gamma-ray bursts, or large meteorite impacts. Some of them can withstand extremely cold temperatures down to 1 K (−458 °F; −272 °C) (close to absolute zero), while others can withstand extremely hot temperatures up to 420 K (300 °F; 150 °C) for several minutes, pressures about six times greater than those found in the deepest ocean trenches, ionizing radiation at doses hundreds of times higher than the lethal dose for a human, and the vacuum of outer space. They can go without food or water for more than 30 years, drying out to the point where they are 3% or less water, only to rehydrate, forage, and reproduce. Tardigrades that live in harsh conditions undergo an annual process of cyclomorphosis, allowing for survival in sub-zero temperatures.
They are not considered extremophilic because they are not adapted to exploit these conditions, only to endure them. This means that their chances of dying increase the longer they are exposed to the extreme environments, whereas true extremophiles thrive in a physically or geochemically extreme environment that would harm most other organisms.
Tardigrades are one of the few groups of species that are capable of suspending their metabolism (see cryptobiosis). Many species of tardigrade can survive in a dehydrated state up to five years, or in exceptional cases longer. Depending on the environment, they may enter this state via anhydrobiosis, cryobiosis, osmobiosis, or anoxybiosis. While in this state, their metabolism lowers to less than 0.01% of normal and their water content can drop to 1% of normal. Their ability to remain desiccated for such long periods was thought to be largely dependent on the high levels of the nonreducing sugar trehalose, which protects their membranes, although recent research suggests that tardigrades have a unique type of disordered protein that serves a similar purpose: It replaces water in the cells and adopts a glassy, vitrified state when the animals dry out. Their DNA is further protected from radiation by a protein called "dsup" (short for damage suppressor). In this cryptobiotic state, the tardigrade is known as a tun.
Tardigrades are able to survive in extreme environments that would kill almost any other animal. Extremes at which tardigrades can survive include those of:
- Temperature – tardigrades can survive:
- Pressure – they can withstand the extremely low pressure of a vacuum and also very high pressures, more than 1,200 times atmospheric pressure. Tardigrades can survive the vacuum of open space and solar radiation combined for at least 10 days. Some species can also withstand pressure of 6,000 atmospheres, which is nearly six times the pressure of water in the deepest ocean trench, the Mariana trench.
- Dehydration – the longest that living tardigrades have been shown to survive in a dry state is nearly 10 years, although there is one report of leg movement, not generally considered "survival", in a 120-year-old specimen from dried moss. When exposed to extremely low temperatures, their body composition goes from 85% water to only 3%. As water expands upon freezing, dehydration ensures the tardigrades do not get ripped apart by the freezing ice.
- Radiation – tardigrades can withstand 1,000 times more radiation than other animals, median lethal doses of 5,000 Gy (of gamma rays) and 6,200 Gy (of heavy ions) in hydrated animals (5 to 10 Gy could be fatal to a human). The only explanation found in earlier experiments for this ability was that their lowered water state provides fewer reactants for ionizing radiation. However, subsequent research found that tardigrades, when hydrated, still remain highly resistant to shortwave UV radiation in comparison to other animals, and that one factor for this is their ability to efficiently repair damage to their DNA resulting from that exposure.
- Irradiation of tardigrade eggs collected directly from a natural substrate (moss) showed a clear dose-related response, with a steep decline in hatchability at doses up to 4 kGy, above which no eggs hatched. The eggs were more tolerant to radiation late in development. No eggs irradiated at the early developmental stage hatched, and only one egg at middle stage hatched, while eggs irradiated in the late stage hatched at a rate indistinguishable from controls.
- Environmental toxins – tardigrades are reported to undergo chemobiosis, a cryptobiotic response to high levels of environmental toxins. However, as of 2001, these laboratory results have yet to be verified.
- Outer space – tardigrades are the first known animal to survive in outer space. In September 2007, dehydrated tardigrades were taken into low Earth orbit on the FOTON-M3 mission carrying the BIOPAN astrobiology payload. For 10 days, groups of tardigrades were exposed to the hard vacuum of outer space, or vacuum and solar UV radiation. After being rehydrated back on Earth, over 68% of the subjects protected from high-energy UV radiation revived within 30 minutes following rehydration, but subsequent mortality was high; many of these produced viable embryos. In contrast, hydrated samples exposed to the combined effect of vacuum and full solar UV radiation had significantly reduced survival, with only three subjects of Milnesium tardigradum surviving. In May 2011, Italian scientists sent tardigrades on board the International Space Station along with extremophiles on STS-134, the final flight of Space Shuttle Endeavour. Their conclusion was that microgravity and cosmic radiation "did not significantly affect survival of tardigrades in flight, confirming that tardigrades represent a useful animal for space research." In November 2011, they were among the organisms to be sent by the U.S.-based Planetary Society on the Russian Fobos-Grunt mission's Living Interplanetary Flight Experiment to Phobos; however, the launch failed. Tardigrades are one of the few groups to have survived Earth's five mass extinctions.
Scientists have conducted morphological and molecular studies to understand how tardigrades relate to other lineages of ecdysozoan animals. Two plausible placements have been proposed: tardigrades are either most closely related to Arthropoda ± Onychophora, or to nematodes. Evidence for the former is a common result of morphological studies; evidence of the latter is found in some molecular analyses.
The latter hypothesis has been rejected by recent microRNA and expressed sequence tag analyses. Apparently, the grouping of tardigrades with nematodes found in a number of molecular studies is a long branch attraction artifact. Within the arthropod group (called panarthropoda and comprising onychophora, tardigrades and euarthropoda), three patterns of relationship are possible: tardigrades sister to onychophora plus arthropods (the lobopodia hypothesis); onychophora sister to tardigrades plus arthropods (the tactopoda hypothesis); and onychophora sister to tardigrades. Recent analyses indicate that the panarthropoda group is monophyletic, and that tardigrades are a sister group of Lobopodia, the lineage consisting of arthropods and Onychophora.
The minute sizes of tardigrades and their membranous integuments make their fossilization both difficult to detect and highly unusual. The only known fossil specimens are those from mid-Cambrian deposits in Siberia and a few rare specimens from Cretaceous amber.
The Siberian tardigrade fossils differ from living tardigrades in several ways. They have three pairs of legs rather than four, they have a simplified head morphology, and they have no posterior head appendages, but they share with modern tardigrades their columnar cuticle construction. Scientists think they represent a stem group of living tardigrades.
Rare specimens in Cretaceous amber have been found in two North American locations. Milnesium swolenskyi, from New Jersey, is the older of the two; its claws and mouthparts are indistinguishable from the living M. tardigradum. The other specimens from amber are from western Canada, some 15–20 million years earlier than M. swolenskyi. One of the two specimens from Canada has been given its own genus and family, Beorn leggi, but it bears a strong resemblance to many living specimens in the family Hypsibiidae.
Aysheaia from the middle Cambrian Burgess shale has been proposed as a sister taxon to an arthropod-tardigrade clade. Tardigrades have been proposed to be among the closest living relatives of the Burgess shale oddity Opabinia.
Genomes and genome sequencing
Tardigrade genomes vary in size, from about 75 to 800 megabase pairs of DNA.Hypsibius dujardini has a compact genome of 100 megabase pairs  and a generation time of about two weeks; it can be cultured indefinitely and cryopreserved.
The genome of Ramazzottius varieornatus, one of the most stress-tolerant species of Tardigrades, was sequenced by a team of researchers from the University of Tokyo in 2015. Analysis revealed less than 1.2% of its genes were the result of horizontal gene transfer. They also found evidence of a loss of gene pathways that are known to promote damage due to stress. This study also found a high expression of novel Tardigrade-unique proteins, including Damage suppressor (Dsup), which was shown to protect against DNA damage from X-ray radiation. The same team applied the Dsup protein to human cultured cells and found that it suppressed X-ray damage to the human cells by ~40%.
Many organisms that live in aquatic environments feed on species such as nematodes, tardigrades, bacteria, algae, mites, and collembolans. Tardigrades work as pioneer species by inhabiting new developing environments. This movement attracts other invertebrates to populate that space, while also attracting predators.
In popular culture
- List of microorganisms tested in outer space
- Living Interplanetary Flight Experiment – study of selected microorganisms in outer space
- Mopsechiniscus franciscae – tardigrade found in Victoria Land Antarctica
- Budd, Graham E (2001). "Tardigrades as 'Stem-Group Arthropods': The Evidence from the Cambrian Fauna". Zoologischer Anzeiger. 240 (3–4): 265–79. doi:10.1078/0044-5231-00034.
- Miller, William. "Tardigrades". American Scientist. Retrieved 2018-04-13.
- Simon, Matt (21 March 2014). "Absurd Creature of the Week: The Incredible Critter That's Tough Enough to Survive in the vacuum of Space". Wired. Retrieved 2014-03-21.
- Copley, Jon (23 October 1999). "Indestructible". New Scientist (2209). Retrieved 2010-02-06.
- "Stanford Tardigrade Project". Foldscope. Retrieved 2017-03-23.
-  The Hindu SEPTEMBER 09, 2015
- Bordenstein, Sarah. "Tardigrades (Water Bears)". Microbial Life Educational Resources. National Science Digital Library. Retrieved 2014-01-24.
- "BBC — Earth — The strange worms that live on erupting mud volcanoes". BBC. Retrieved 2017-04-15.
- "Tardigrades". Tardigrade. Retrieved 2015-09-21.
- Guarino, Ben (14 July 2017). "These animals can survive until the end of the Earth, astrophysicists say". Washington Post. Retrieved 14 July 2017.
- Sloan, David; Alves Batista, Rafael; Loeb, Abraham (2017). "The Resilience of Life to Astrophysical Events". Scientific Reports. 7 (1): 5419. arXiv:1707.04253. Bibcode:2017NatSR...7.5419S. doi:10.1038/s41598-017-05796-x. PMC 5511186. PMID 28710420.
- Zhang, Zhi-Qiang (2011). "Animal biodiversity: An introduction to higher-level classification and taxonomic richness" (PDF). Zootaxa. 3148: 7–12.
- Degma, Peter; Bertolani, Roberto; Guidetti, Roberto. "Actual checklist of Tardigrada species (2009-2015, Ver. 28: 31-03-2015)" (PDF). Archived from the original (PDF) on 8 May 2010.
- "Tardigrada (water bears, tardigrades)". biodiversity explorer. Retrieved 2013-05-31.
- Nelson, Diane (1 July 2002). "Current status of Tardigrada:Evolution and Ecology". Integrative and Comparative Biology. pp. 652–659. doi:10.1093/icb/42.3.652. Archived from the original on 2017-05-02. Retrieved 3 August 2018.
- Shaw, Michael W. "How to Find Tardigrades". tardigrades.us. Archived from the original on 10 February 2014. Retrieved 2013-01-14.
- Bordenstein, Sarah (17 December 2008). "Tardigrades (Water Bears)". Carleton College. Retrieved 2012-09-16.
- Staff (29 September 2015). "Researchers discover new tiny organism, name it for Wyeths". AP News. Retrieved 2015-09-29.
- Perry, Emma S; Miller, William R (2015). "Echiniscoides wyethi, a new marine tardigrade from Maine, U.S.A. (Heterotardigrada: Echiniscoidea: Echiniscoididae)". Proceedings of the Biological Society of Washington. 128 (1): 103–10. doi:10.2988/0006-324X-128.1.103.
- Goldstein, Bob; Blaxter, Mark (2002). "Tardigrades". Current Biology. 12 (14): R475. doi:10.1016/S0960-9822(02)00959-4. PMID 12176341.
- Romano, Frank A. (2003). "On water bears". Florida Entomologist. 86 (2): 134–137. doi:10.1653/0015-4040(2003)086[0134:OWB]2.0.CO;2.
- Smith, Frank W.; Boothby, Thomas C.; Giovannini, Ilaria; Rebecchi, Lorena; Jockusch, Elizabeth L.; Goldstein, Bob (1 January 2016). "The Compact Body Plan of Tardigrades Evolved by the Loss of a Large Body Region". Current Biology. 26 (2): 224–229. doi:10.1016/j.cub.2015.11.059. Retrieved 29 July 2018.
- Seki, Kunihiro; Toyoshima, Masato (1998). "Preserving tardigrades under pressure". Nature. 395 (6705): 853–854. Bibcode:1998Natur.395..853S. doi:10.1038/27576.
- Kinchin, Ian M. (1994) The Biology of Tardigrades, Ashgate Publishing
- Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 877–80. ISBN 0-03-056747-5.
- Segmentation in Tardigrada and diversification of segmental patterns in Panarthropoda
- Gross, Vladimir; Mayer, Georg (2015). "Neural development in the tardigrade Hypsibius dujardini based on anti-acetylated α-tubulin immunolabeling". EvoDevo. 6: 12. doi:10.1186/s13227-015-0008-4. PMC 4458024. PMID 26052416.
- Zantke, Juliane; Wolff, Carsten; Scholtz, Gerhard (2007). "Three-dimensional reconstruction of the central nervous system of Macrobiotus hufelandi (Eutardigrada, Parachela): Implications for the phylogenetic position of Tardigrada". Zoomorphology. 127 (1): 21–36. doi:10.1007/s00435-007-0045-1.
- Greven, Hartmut (2007). "Comments on the eyes of tardigrades". Arthropod Structure & Development. 36 (4): 401–7. doi:10.1016/j.asd.2007.06.003. PMID 18089118.
- Elzinga, Richard J (1998). "Microspines in the alimentary canal of arthropoda, onychophora, annelida". International Journal of Insect Morphology and Embryology. 27 (4): 341–9. doi:10.1016/S0020-7322(98)00027-0.
- Morgan, Clive I. (1977). "Population dynamics of two species of Tardigrada, Macrobiotus hufelandii (Schultze) and Echiniscus (Echiniscus) testudo (Doyere), in roof moss from Swansea". Journal of Animal Ecology. British Ecological Society. 46 (1): 263–279. doi:10.2307/3960. JSTOR 3960.
- Lindahl, K. (2008-03-15). "Tardigrade Facts".
- Brent Nichols, Phillip (2005). Tardigrade Evolution and Ecology (PhD). Tampa, FL: University of South Florida.
- Glime, Janice (2010). "Tardigrades". Bryophyte Ecology: Volume 2, Bryogological Interaction.
- Hogan, C. Michael (2010). E. Monosson; C. Cleveland., eds. "Extremophile". Encyclopedia of Earth. Washington, DC: National Council for Science and the Environment.
- Simon, Matt. "Absurd Creature of the Week: The Incredible Critter That's Tough Enough to Survive in Space".
- Dean, Cornelia (7 September 2015). "The Tardigrade: Practically Invisible, Indestructible 'Water Bears'". New York Times. Retrieved 7 September 2015.
- Brennand, Emma (17 May 2011). "Tardigrades: Water bears in space". BBC. Retrieved 2013-05-31.
- Crowe, John H.; Carpenter, John F.; Crowe, Lois M. (October 1998). "The role of vitrification in anhydrobiosis". Annual Review of Physiology. 60. pp. 73–103. doi:10.1146/annurev.physiol.60.1.73. PMID 9558455.
- Guidetti, Roberto; Jönsson, K. Ingemar (2002). "Long-term anhydrobiotic survival in semi-terrestrial micrometazoans". Journal of Zoology. 257 (2): 181–187. doi:10.1017/S095283690200078X.
- Chimileski, Scott; Kolter, Roberto (2017). Life at the Edge of Sight: A Photographic Exploration of the Microbial World. Cambridge, MA: Belknap Press: An Imprint of Harvard University Press. ISBN 067497591X.
- Halberg, Kenneth Agerlin; Persson, Dennis; Ramløv, Hans; Westh, Peter; Kristensen, Reinhardt Møbjerg; Møbjerg, Nadja (1 September 2009). "Cyclomorphosis in Tardigrada: adaptation to environmental constraints". Journal of Experimental Biology. 212 (17): 2803–2811. doi:10.1242/jeb.029413. PMID 19684214. Retrieved 11 December 2017 – via jeb.biologists.org.
- Rampelotto, Pabulo Henrique (2010). "Resistance of Microorganisms to Extreme Environmental Conditions and Its Contribution to Astrobiology". Sustainability. 2 (6): 1602–23. Bibcode:2010Sust....2.1602R. doi:10.3390/su2061602.
- Rothschild, Lynn J; Mancinelli, Rocco L (2001). "Life in extreme environments". Nature. 409 (6823): 1092–101. Bibcode:2001Natur.409.1092R. doi:10.1038/35059215. PMID 11234023.
- Bell, Graham (2016). "Experimental macroevolution". Proceedings of the Royal Society B: Biological Sciences. 283 (1822): 20152547. doi:10.1098/rspb.2015.2547. PMC 4721102.
- Anderson, David. "Humans are just starting to understand this nearly invincible creature — and it's fascinating". BusinessInsider.com. Business Insider Inc. Retrieved 26 October 2017.
- Boothby, Thomas C; Tapia, Hugo; Brozena, Alexandra H; Piszkiewicz, Samantha; Smith, Austin E; Giovannini, Ilaria; Rebecchi, Lorena; Pielak, Gary J; Koshland, Doug; Goldstein, Bob (2017). "Tardigrades Use Intrinsically Disordered Proteins to Survive Desiccation". Molecular Cell. 65 (6): 975–984.e5. doi:10.1016/j.molcel.2017.02.018. PMID 28306513.
- Tauger, Nathan; Gill, Victoria (20 September 2016). "Survival secret of 'Earth's hardiest animal' revealed". BBC News. Retrieved 2016-09-21.
- Hashimoto, Takuma; Horikawa, Daiki D; Saito, Yuki; Kuwahara, Hirokazu; Kozuka-Hata, Hiroko; Shin-i, Tadasu; Minakuchi, Yohei; Ohishi, Kazuko; Motoyama, Ayuko; Aizu, Tomoyuki; Enomoto, Atsushi; Kondo, Koyuki; Tanaka, Sae; Hara, Yuichiro; Koshikawa, Shigeyuki; Sagara, Hiroshi; Miura, Toru; Yokobori, Shin-Ichi; Miyagawa, Kiyoshi; Suzuki, Yutaka; Kubo, Takeo; Oyama, Masaaki; Kohara, Yuji; Fujiyama, Asao; Arakawa, Kazuharu; Katayama, Toshiaki; Toyoda, Atsushi; Kunieda, Takekazu (2016). "Extremotolerant tardigrade genome and improved radiotolerance of human cultured cells by tardigrade-unique protein". Nature Communications. 7: 12808. Bibcode:2016NatCo...712808H. doi:10.1038/ncomms12808. PMC 5034306. PMID 27649274.
- Piper, Ross (2007), Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals, Greenwood Press. p. 277. ISBN 978-0-313-33922-6.
- Horikawa, Daiki D (2012). "Survival of Tardigrades in Extreme Environments: A Model Animal for Astrobiology". In Altenbach, Alexander V.; Bernhard, Joan M.; Seckbach, Joseph. Anoxia. Cellular Origin, Life in Extreme Habitats and Astrobiology. 21. pp. 205–17. doi:10.1007/978-94-007-1896-8_12. ISBN 978-94-007-1895-1.
- Tsujimoto, Megumu; Imura, Satoshi; Kanda, Hiroshi (February 2015). "Recovery and reproduction of an Antarctic tardigrade retrieved from a moss sample frozen for over 30 years". Cryobiology. 72 (1): 78–81. doi:10.1016/j.cryobiol.2015.12.003. PMID 26724522.
- Becquerel, Paul (1950). "La suspension de la vie au dessous de 1⁄20 K absolu par demagnetization adiabatique de l'alun de fer dans le vide les plus eléve" [The suspension of life below 1⁄20 K absolute by adiabatic demagnetization of iron alum in the highest vacuum]. Comptes Rendus des Séances de l'Académie des Sciences (in French). 231 (4): 261–3.
- Jönsson, K. Ingemar; Rabbow, Elke; Schill, Ralph O; Harms-Ringdahl, Mats; Rettberg, Petra (2008). "Tardigrades survive exposure to space in low Earth orbit". Current Biology. 18 (17): R729–R731. doi:10.1016/j.cub.2008.06.048. PMID 18786368.
- Jönsson, K. Ingemar; Bertolani, Roberto (2001). "Facts and fiction about long-term survival in tardigrades". Journal of Zoology. 255 (1): 121–3. doi:10.1017/S0952836901001169.
- Franceschi, T. (1948). "Anabiosi nei tardigradi" [Anabiosis in Tardigrades]. Bollettino dei Musei e degli Istituti Biologici dell'Università di Genova (in Italian). 22: 47–9.
- Kent, Michael (2000), Advanced Biology, Oxford University Press
- "Radiation tolerance in the tardigrade Milnesium tardigradum" (PDF).[permanent dead link]
- Horikawa, Daiki D; Sakashita, Tetsuya; Katagiri, Chihiro; Watanabe, Masahiko; Kikawada, Takahiro; Nakahara, Yuichi; Hamada, Nobuyuki; Wada, Seiichi; Funayama, Tomoo; Higashi, Seigo; Kobayashi, Yasuhiko; Okuda, Takashi; Kuwabara, Mikinori (2009). "Radiation tolerance in the tardigrade Milnesium tardigradum". International Journal of Radiation Biology. 82 (12): 843–8. doi:10.1080/09553000600972956. PMID 17178624.
- Horikawa, Daiki D. "UV Radiation Tolerance of Tardigrades". NASA.com. Retrieved 2013-01-15.
- Jönsson, Ingemar; Beltran-Pardo, Eliana; Haghdoost, Siamak; Wojcik, Andrzej; Bermúdez-Cruz, Rosa María; Bernal Villegas, Jaime E; Harms-Ringdahl, Mats (2013). "Tolerance to gamma-irradiation in eggs of the tardigrade Richtersius coronifer depends on stage of development". Journal of Limnology. 72 (1): 9. doi:10.4081/jlimnol.2013.s1.e9.
- "Creature Survives Naked in Space". Space.com. 8 September 2008. Retrieved 2011-12-22.
- Mustain, Andrea (22 December 2011). "Weird wildlife: The real land animals of Antarctica". MSNBC. Retrieved 2011-12-22.
- Courtland, Rachel (8 September 2008). "'Water bears' are first animal to survive space vacuum". New Scientist. Retrieved 2011-05-22.
- NASA Staff (17 May 2011). "BIOKon In Space (BIOKIS)". NASA. Retrieved 2011-05-24.
- Brennard, Emma (17 May 2011). "Tardigrades: Water bears in space". BBC. Retrieved 2011-05-24.
- "Tardigrades: Water bears in space". BBC Nature. 17 May 2011.
- Rebecchi, L.; Altiero, T.; Rizzo, A. M.; Cesari, M.; Montorfano, G.; Marchioro, T.; Bertolani, R.; Guidetti, R. (July 2012). "Two tardigrade species on board of the STS-134 space flight". 12th International Symposium on Tardigrada (PDF). p. 89. hdl:2434/239127. ISBN 978-989-96860-7-6.
- Cosmos: A Spacetime Odyssey[full citation needed]
- Campbell, Lahcen I.; Rota-Stabelli, Omar; Edgecombe, Gregory D.; Marchioro, Trevor; Longhorn, Stuart J.; Telford, Maximilian J.; Philippe, Hervé; Rebecchi, Lorena; Peterson, Kevin J.; Pisani, Davide (2011). "MicroRNAs and phylogenomics resolve the relationships of Tardigrada and suggest that velvet worms are the sister group of Arthropoda". Proceedings of the National Academy of Sciences. 108 (38): 15920–4. Bibcode:2011PNAS..10815920C. doi:10.1073/pnas.1105499108. PMC 3179045. PMID 21896763.
- Telford, Maximilian J.; Bourlat, Sarah J.; Economou, Andrew; Papillon, Daniel; Rota-Stabelli, Omar (2008). "The evolution of the Ecdysozoa". Philosophical Transactions of the Royal Society B: Biological Sciences. 363 (1496): 1529–37. doi:10.1098/rstb.2007.2243. JSTOR 20208544. PMC 2614232. PMID 18192181.
- Blaxter, Mark; Elsworth, Ben; Daub, Jennifer (2004). "DNA taxonomy of a neglected animal phylum: An unexpected diversity of tardigrades". Proceedings of the Royal Society B: Biological Sciences. 271: S189–92. doi:10.1098/rsbl.2003.0130. JSTOR 4142714. PMC 1810026. PMID 15252980.
- Grimaldi, David A.; Engel, Michael S. (2005). Evolution of the Insects. Cambridge University Press. pp. 96–97. ISBN 0-521-82149-5.
- Cooper, Kenneth W. (1964). "The first fossil tardigrade: Beorn leggi, from Cretaceous Amber". Psyche: A Journal of Entomology. 71 (2): 41–48. doi:10.1155/1964/48418.
- Fortey, Richard A.; Thomas, Richard H. (2001). Arthropod Relationships. Chapman & Hall. p. 383. ISBN 0-412-75420-7.
- Budd, Graham E (1996). "The morphology of Opabinia regalis and the reconstruction of the arthropod stem-group". Lethaia. 29 (1): 1–14. doi:10.1111/j.1502-3931.1996.tb01831.x.
- "Genome Size of Tardigrades".
- Yoshida, Yuki; Koutsovoulos, Georgios; Laetsch, Dominik R.; Stevens, Lewis; Kumar, Sujai; Horikawa, Daiki D.; Ishino, Kyoko; Komine, Shiori; Kunieda, Takekazu; Tomita, Masaru; Blaxter, Mark; Arakawa, Kazuharu; Tyler-Smith, Chris (27 July 2017). "Comparative genomics of the tardigrades Hypsibius dujardini and Ramazzottius varieornatus". PLOS Biology. 15 (7): e2002266. doi:10.1371/journal.pbio.2002266.
- Gabriel, Willow N; McNuff, Robert; Patel, Sapna K; Gregory, T. Ryan; Jeck, William R; Jones, Corbin D; Goldstein, Bob (2007). "The tardigrade Hypsibius dujardini, a new model for studying the evolution of development". Developmental Biology. 312 (2): 545–59. doi:10.1016/j.ydbio.2007.09.055. PMID 17996863.
- Kinchin, IM (1987). "The moss fauna 1; Tardigrades". Journal of Biological Education. 21(4): 288–290.
- "13 ways Neil deGrasse Tyson's "Cosmos" sent the religious right off the deep end". 14 June 2014.
- "Steam Greenlight page for Tardigrades game". 21 November 2014.
- "CBS' Star Trek: Discovery is being sued for 'plagiarising' a 2014 indie game". 30 August 2018. Retrieved 7 September 2018.
- "'Harbinger Down': New trailer for creature feature". Retrieved 3 October 2018.
- "'Harbinger Down' Review: A Bleak & Vanilla Creature Feature". bloody-disgusting.com. Retrieved 3 October 2018.
- "How the Quantum Realm could play into future Marvel films". 10 July 2018. Retrieved 29 July 2018.
- "The Science (and the Scientists) Behind 'Ant-Man'". Retrieved 29 July 2018.
- "Ant-Man and the Wasp needs a little help". 4 July 2018. Retrieved 29 July 2018.
- "The Scientific Truth About Ripper the 'Star Trek' Tardigrade Is a Huge Relief". Retrieved 5 September 2018.
- Salzberg, Steven. "New 'Star Trek' Series Makes Massive Science Blunder". Retrieved 5 September 2018.
- Placido, Dani Di. "'South Park' Review: Cartman Creates A Monster In 'Moss Piglets'". Retrieved 29 July 2018.
- "'South Park' season 21 episode 8 live stream: 'Moss Piglets'". 15 November 2017. Retrieved 29 July 2018.
|Wikispecies has information related to Tardigrada|
|Wikimedia Commons has media related to Tardigrada.|
- Tardigrade water bear NY Times, 2015
- Tardigrada Newsletter
- Tardigrades – Pictures and Movies
- The Edinburgh Tardigrade project
- Instructions for finding tardigrades
- The incredible water bear!
- Tardigrade Reference Center
- Tardigrades in space
- Tardigrade data and analysis
- A short film about tardigrade research from NPR's Science Friday
- Tardigrada at the Tree of Life Web Project
- Swiss Center of Tardigrade Research – Ecology, Physiology and Evolutionary Biology of Tardigrades
- NASA Astronomy Picture of the Day: Tardigrade in Moss (6 March 2013)
- Video (07:54) – First Animal to Survive in Space
- Video (00:38) – Tardigrade Movement in Water
- Tardigrades are so tough, they can survive outer space (March 2015). BBC
- The International Society of Tardigrade Hunters
- Tardigrades discussed on RNZ Critter of the Week, 14 July 2017