|Its distribution is marked in red.|
The axolotl ( // ; from Classical Nahuatl : āxōlōtl [aːˈʃoːloːtɬ] ( listen )) (Ambystoma mexicanum) is a paedomorphic salamander closely related to the tiger salamander. It is unusual among amphibians in that it reaches adulthood without undergoing metamorphosis. Instead of taking to the land, adults remain aquatic and gilled. The species was originally found in several lakes underlying what is now Mexico City, such as Lake Xochimilco and Lake Chalco. These lakes were drained by Spanish settlers after the conquest of the Aztec Empire, leading to the destruction of much of the axolotl’s natural habitat.
Axolotls should not be confused with the larval stage of the closely related tiger salamander (A. tigrinum), which are widespread in much of North America and occasionally become paedomorphic. Neither should they be confused with mudpuppies (Necturus spp.), fully aquatic salamanders from a different family that are not closely related to the axolotl but bear a superficial resemblance.
As of 2020 [update] , the axolotl was near extinction due to urbanization in Mexico City and consequent water pollution, as well as the introduction of invasive species such as tilapia and perch. It is listed as critically endangered in the wild, with a decreasing population of around 50 to 1,000 adult individuals, by the International Union for Conservation of Nature and Natural Resources (IUCN) and is listed under Appendix II of the Convention on International Trade in Endangered Species (CITES). Axolotls are used extensively in scientific research due to their ability to regenerate limbs, gills and parts of their eyes and brains. Axolotls were also sold as food in Mexican markets and were a staple in the Aztec diet.
A sexually mature adult axolotl, at age 18–27 months, ranges in length from 15 to 45 cm (6 to 18 in), although a size close to 23 cm (9 in) is most common and greater than 30 cm (12 in) is rare. Axolotls possess features typical of salamander larvae, including external gills and a caudal fin extending from behind the head to the vent. External gills are usually lost when salamander species mature into adulthood, although the axolotl maintains this feature. This is due to their neoteny evolution, where axolotls are much more aquatic than other salamander species.
Their heads are wide, and their eyes are lidless. Their limbs are underdeveloped and possess long, thin digits. Males are identified by their swollen cloacae lined with papillae, while females are noticeable for their wider bodies full of eggs. Three pairs of external gill stalks (rami) originate behind their heads and are used to move oxygenated water. The external gill rami are lined with filaments (fimbriae) to increase surface area for gas exchange.Four-gill slits lined with gill rakers are hidden underneath the external gills, which prevent food from entering and allow particles to filter through.
Axolotls have barely visible vestigial teeth, which develop during metamorphosis. The primary method of feeding is by suction, during which their rakers interlock to close the gill slits. External gills are used for respiration, although buccal pumping (gulping air from the surface) may also be used to provide oxygen to their lungs.Buccal pumping can occur in a two-stroke manner that pumps air from the mouth to the lungs, and with four-stroke that reverses this pathway with compression forces.
Axolotls have four pigmentation genes; when mutated they create different color variants. The normal wild-type animal is brown/tan with gold speckles and an olive undertone. The five more common mutant colors are leucistic (pale pink with black eyes), golden albino (golden with gold eyes), xanthic (grey with black eyes), albino (pale pink/white with red eyes) which is more common in axolotls than some other creatures, and melanoid (all black/dark blue with no gold speckling or olive tone).In addition, there is wide individual variability in the size, frequency, and intensity of the gold speckling and at least one variant that develops a black and white piebald appearance on reaching maturity. Because pet breeders frequently cross the variant colors, double homozygous mutants are common in the pet trade, especially white/pink animals with pink eyes that are double homozygous mutants for both the albino and leucistic trait. Axolotls also have some limited ability to alter their color to provide better camouflage by changing the relative size and thickness of their melanophores.
The axolotl is native only to the freshwater of Lake Xochimilco and Lake Chalco in the Valley of Mexico. Lake Chalco no longer exists, having been drained as a flood control measure, and Lake Xochimilco remains a remnant of its former self, existing mainly as canals. The water temperature in Xochimilco rarely rises above 20 °C (68 °F), although it may fall to 6–7 °C (43–45 °F) in the winter, and perhaps lower.
Surveys in 1998, 2003, and 2008 found 6,000, 1,000, and 100 axolotls per square kilometer in its Lake Xochimilco habitat, respectively.A four-month-long search in 2013, however, turned up no surviving individuals in the wild. Just a month later, two wild ones were spotted in a network of canals leading from Xochimilco.
The wild population has been put under heavy pressure by the growth of Mexico City. The axolotl is currently on the International Union for Conservation of Nature's annual Red List of threatened species. Non-native fish, such as African tilapia and Asian carp, have also recently been introduced to the waters. These new fish have been eating the axolotls' young, as well as their primary source of food.
Axolotls are members of the tiger salamander, or Ambystoma tigrinum , species complex, along with all other Mexican species of Ambystoma. Their habitat is like that of most neotenic species—a high-altitude body of water surrounded by a risky terrestrial environment. These conditions are thought to favor neoteny. However, a terrestrial population of Mexican tiger salamanders occupies and breeds in the axolotl's habitat.[ citation needed ]
The axolotl is carnivorous, consuming small prey such as mollusks,worms, insects, other arthropods, and small fish in the wild. Axolotls locate food by smell, and will "snap" at any potential meal, sucking the food into their stomachs with vacuum force.
Today, the axolotl is still used in research as a model organism, and large numbers are bred in captivity. They are especially easy to breed compared to other salamanders in their family, which are rarely captive-bred due to the demands of terrestrial life. One attractive feature for research is the large and easily manipulated embryo, which allows viewing of the full development of a vertebrate. Axolotls are used in heart defect studies due to the presence of a mutant gene that causes heart failure in embryos. Since the embryos survive almost to hatching with no heart function, the defect is very observable. The axolotl is also considered an ideal animal model for the study of neural tube closure due to the similarities between human and axolotl neural plate and tube formation; the axolotl's neural tube, unlike the frog's, is not hidden under a layer of superficial epithelium.There are also mutations affecting other organ systems some of which are not well characterized and others that are. The genetics of the color variants of the axolotl have also been widely studied.
The feature of the axolotl that attracts most attention is its healing ability: the axolotl does not heal by scarring and is capable of the regeneration of entire lost appendages in a period of months, and, in certain cases, more vital structures, such as tail, limb, central nervous system, and tissues of the eye and heart.They can even restore less vital parts of their brains. They can also readily accept transplants from other individuals, including eyes and parts of the brain—restoring these alien organs to full functionality. In some cases, axolotls have been known to repair a damaged limb, as well as regenerating an additional one, ending up with an extra appendage that makes them attractive to pet owners as a novelty. In metamorphosed individuals, however, the ability to regenerate is greatly diminished. The axolotl is therefore used as a model for the development of limbs in vertebrates. There are three basic requirements for regeneration of the limb: the wound epithelium, nerve signaling, and the presence of cells from the different limb axes. A wound epidermis is quickly formed by the cells to cover up the site of the wound. In the following days, the cells of the wound epidermis divide and grow quickly forming a blastema, which means the wound is ready to heal and undergo patterning to form the new limb.
It is believed that during limb generation, axolotls have a different system to regulate their internal macrophage level and suppress inflammation, as scarring prevents proper healing and regeneration.However, this belief has been questioned by other studies. The axolotl’s regenerative properties leave the species as the perfect model to study the process of stem cells and its own neoteny feature. Current research can record specific examples of these regenerative properties through tracking cell fates and behaviors, lineage tracing skin triploid cell grafts, pigmentation imaging, electroporation, tissue clearing and lineage tracing from dye labeling. The newer technologies of germline modification and transgenesis are better suited for live imaging the regenerative processes that occur for axolotls.
The 32 billion base pair long sequence of the axolotl's genome was published in 2018 and was the largest animal genome completed at the time. It revealed species-specific genetic pathways that may be responsible for limb regeneration. bp) which are 13, 16 and 25 times that observed in human (1,750 bp), mouse (1,469 bp) and Tibetan frog (906 bp), respectively.Although the axolotl genome is about 10 times as large as the human genome, it encodes a similar number of proteins, namely 23,251 (the human genome encodes about 20,000 proteins). The size difference is mostly explained by a large fraction of repetitive sequences, but such repeated elements also contribute to increased median intron sizes (22,759
When most amphibians are young, they live in water, and they use gills that can breathe in the water. When they become adults, they go through a process called metamorphosis, in which they lose their gills and start living on land. However, the axolotl is unusual in that it has a lack of thyroid stimulating hormone, which is needed for the thyroid to produce thyroxine in order for the axolotl to go through metamorphosis; therefore, it keeps its gills and lives in water all its life, even after it becomes an adult and is able to reproduce. Its body has the capacity to go through metamorphosis if given the necessary hormone, but axolotls do not produce it, and must be exposed to it from an external source,after which an axolotl undergoes an artificially-induced metamorphosis and begins living on land. One method of artificial metamorphosis induction is through an injection of iodine, which is used in the production of thyroid hormones.
An axolotl undergoing metamorphosis experiences a number of physiological changes that help them adapt to life on land. These include increased muscle tone in limbs, the absorption of gills and fins into the body, the development of eyelids, and a reduction in the skin's permeability to water, allowing the axolotl to stay more easily hydrated when on land. The lungs of an axolotl, though present alongside gills after reaching non-metamorphosed adulthood, develop further during metamorphosis.
An axolotl that has gone through metamorphosis resembles an adult plateau tiger salamander, though the axolotl differs in its longer toes.[ citation needed ] The process of artificially inducing metamorphosis can often result in death during or even following a successful attempt, and so casual hobbyists are generally discouraged from attempting to induce metamorphosis in pet axolotls.
Neoteny is the term for reaching sexual maturity without undergoing metamorphosis.Many other species within the axolotl's genus are also either entirely neotenic or have neotenic populations. Sirens and Necturus are other neotenic salamanders, although unlike axolotls, they cannot be induced to metamorphose by an injection of iodine or thyroxine hormone.
The genes responsible for neoteny in laboratory animals may have been identified; however, they are not linked in wild populations, suggesting artificial selection is the cause of complete neoteny in laboratory and pet axolotls.
Six adult axolotls (including a leucistic specimen) were shipped from Mexico City to the Jardin des Plantes in Paris in 1863. Unaware of their neoteny, Auguste Duméril was surprised when, instead of the axolotl, he found in the vivarium a new species, similar to the salamander.[ verification needed ] This discovery was the starting point of research about neoteny. It is not certain that Ambystoma velasci specimens were not included in the original shipment.[ citation needed ] Vilem Laufberger in Prague used thyroid hormone injections to induce an axolotl to grow into a terrestrial adult salamander. The experiment was repeated by Englishman Julian Huxley, who was unaware the experiment had already been done, using ground thyroids. Since then, experiments have been done often with injections of iodine or various thyroid hormones used to induce metamorphosis.
Neoteny has been observed in all salamander families in which it seems to be a survival mechanism, in aquatic environments only of mountain and hill, with little food and, in particular, with little iodine. In this way, salamanders can reproduce and survive in the form of a smaller larval stage, which is aquatic and requires a lower quality and quantity of food compared to the big adult, which is terrestrial. If the salamander larvae ingest a sufficient amount of iodine, directly or indirectly through cannibalism, they quickly begin metamorphosis and transform into bigger terrestrial adults, with higher dietary requirements.In fact, in some high mountain lakes there live dwarf forms of salmonids that are caused by deficiencies in food and, in particular, iodine, which causes cretinism and dwarfism due to hypothyroidism, as it does in humans.
The axolotl is a popular exotic pet like its relative, the tiger salamander (Ambystoma tigrinum). As for all poikilothermic organisms, lower temperatures result in slower metabolism and a very unhealthily reduced appetite. Temperatures at approximately 16 °C (61 °F) to 18 °C (64 °F) are suggested for captive axolotls to ensure sufficient food intake; stress resulting from more than a day's exposure to lower temperatures may quickly lead to disease and death, and temperatures higher than 24 °C (75 °F) may lead to metabolic rate increase, also causing stress and eventually death. Chlorine, commonly added to tapwater, is harmful to axolotls. A single axolotl typically requires a 150-litre (40-US-gallon) tank. Axolotls spend the majority of the time at the bottom of the tank.
Salts, such as Holtfreter's solution, are often added to the water to prevent infection.
In captivity, axolotls eat a variety of readily available foods, including trout and salmon pellets, frozen or live bloodworms, earthworms, and waxworms. Axolotls can also eat feeder fish, but care should be taken as fish may contain parasites.
Substrates are another important consideration for captive axolotls, as axolotls (like other amphibians and reptiles) tend to ingest bedding material together with food cm (or approximately the size of the animal's head) should be available to the animal.and are commonly prone to gastrointestinal obstruction and foreign body ingestion. Some common substrates used for animal enclosures can be harmful for amphibians and reptiles. Gravel (common in aquarium use) should not be used, and is recommended that any sand consists of smooth particles with a grain size of under 1mm. One guide to axolotl care for laboratories notes that bowel obstructions are a common cause of death, and recommends that no items with a diameter below 3
There is some evidence that axolotls might seek out appropriately-sized gravel for use as gastrolithsbased on experiments conducted at the University of Manitoba axolotl colony, but these studies are outdated and not conclusive. As there is no conclusive evidence pointing to gastrolith use, gravel should be avoided due to the high risk of impaction.
The species is named after the Aztec deity Xolotl, who transformed himself into an axolotl. They continue to play an outsized cultural role in Mexico, and have appeared in cartoons and murals.
In 2020, it was announced that the axolotl will be featured on the new design for Mexico's 50-peso banknote, along with images of maize and chinampas.
Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arboreal or freshwater aquatic ecosystems. Thus amphibians typically start out as larvae living in water, but some species have developed behavioural adaptations to bypass this.
Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some insects, fish, amphibians, mollusks, crustaceans, cnidarians, echinoderms, and tunicates undergo metamorphosis, which is often accompanied by a change of nutrition source or behavior. Animals can be divided into species that undergo complete metamorphosis ("holometaboly"), incomplete metamorphosis ("hemimetaboly"), or no metamorphosis ("ametaboly").
Neoteny, also called juvenilization, is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared to other primates. In progenesis or paedogenesis, sexual development is accelerated.
Salamanders are a group of amphibians typically characterized by their lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All ten extant salamander families are grouped together under the order Urodela. Salamander diversity is highest in eastern North America, especially in the Appalachian Mountains; most species are found in the Holarctic realm, with some species present in the Neotropical realm.
The mole salamanders are a group of advanced salamanders endemic to North America. The group has become famous due to the presence of the axolotl, widely used in research due to its paedomorphosis, and the tiger salamander which is the official amphibian of many states, and often sold as a pet.
The tiger salamander is a species of mole salamander and one of the largest terrestrial salamanders in North America.
Ambystomatidae is a family of salamanders belonging to the order Caudata in the class Amphibia. It contains two genera, Ambystoma and Dicamptodon. Ambystoma contains 32 species and are distributed widely across North America, while Dicamptodon contains four species restricted to the Pacific Northwest. These salamanders are mostly terrestrial and eat invertebrates, although some species are known to eat smaller salamanders. They can be found throughout the US and some areas of Canada in damp forests or plains. This family contains some of the largest terrestrial salamanders in the world, the tiger salamander and the coastal giant salamander. Some species are toxic and can secrete poison from their bodies as protection against predators or infraspecific competition. Neoteny has been observed in several species in Ambystomatidae, and some of them like the axolotl live all of their lives under water in their larval stage.
In biology, regeneration is the process of renewal, restoration, and tissue growth that makes genomes, cells, organisms, and ecosystems resilient to natural fluctuations or events that cause disturbance or damage. Every species is capable of regeneration, from bacteria to humans. Regeneration can either be complete where the new tissue is the same as the lost tissue, or incomplete where after the necrotic tissue comes fibrosis.
In evolutionary developmental biology, heterochrony is any genetically controlled difference in the timing, rate, or duration of a developmental process in an organism compared to its ancestors or other organisms. This leads to changes in the size, shape, characteristics and even presence of certain organs and features. It is contrasted with heterotopy, a change in spatial positioning of some process in the embryo, which can also create morphological innovation. Heterochrony can be divided into intraspecific heterochrony, variation within a species, and interspecific heterochrony, phylogenetic variation, i.e. variation of a descendant species with respect to an ancestral species.
The common mudpuppy is a species of salamander in the genus Necturus. They live an entirely aquatic lifestyle in parts of North America in lakes, rivers, and ponds. They go through paedomorphosis and retain their external gills. Because skin and lung respiration alone is not sufficient for gas exchange, mudpuppies must rely on external gills as their primary means of gas exchange. They are usually a rusty brown color and can grow to an average length of 13 in (330 mm). Mudpuppies are nocturnal creatures, and come out during the day only if the water in which they live is murky. Their diet consists of almost anything they can get in their mouths, including insects, mollusks, and earthworms. Once a female mudpuppy reaches sexual maturity at six years of age, she can lay an average of 60 eggs. In the wild, the average lifespan of a mudpuppy is 11 years.
The ringed salamander is a species of mole salamander native to hardwood and mixed hardwood-pine forested areas in and around the Ozark Plateau and Ouachita Mountains of Arkansas, Oklahoma, and Missouri. This species of salamander has slander body, small head, and long tail. They are usually found to have various dorsal color from dark gray to dark brown. Various close relatives are found such as Ringed salamander, marbled salamander, and spotted salamander. This species of salamander has cannibal behavior espeically those in large body size.
The northwestern salamander is a species of mole salamander that inhabits the northwest Pacific coast of North America. These fairly large salamanders grow to 8.7 in (220 mm) in length. It is found from southeastern Alaska on May Island, through Washington and Oregon south to the mouth of the Gualala River, Sonoma County, California. It occurs from sea level to the timberline, but not east of the Cascade Divide. Its range includes Vancouver Island in British Columbia and The San Juan Islands, Cypress, Whidbey, Bainbridge, and Vashon Islands in Washington.
The long-toed salamander is a mole salamander in the family Ambystomatidae. This species, typically 4.1–8.9 cm (1.6–3.5 in) long when mature, is characterized by its mottled black, brown, and yellow pigmentation, and its long outer fourth toe on the hind limbs. Analysis of fossil records, genetics, and biogeography suggest A. macrodactylum and A. laterale are descended from a common ancestor that gained access to the western Cordillera with the loss of the mid-continental seaway toward the Paleocene.
Taylor's salamander is a species of salamander found only in Laguna Alchichica, a high-altitude crater lake to the southwest of Perote, Mexico. It was first described in 1982 but had been known to science prior to that. It is a neotenic salamander, breeding while still in the larval state and not undergoing metamorphosis. The lake in which it lives is becoming increasingly saline and less suitable for the salamander, which is declining in numbers. The International Union for Conservation of Nature (IUCN) has rated it as being "critically endangered".
Anderson's salamander is a neotenic salamander from Zacapu Lagoon in the Mexican state of Michoacán.
The Lake Patzcuaro salamander, locally known as achoque is a neotenic salamander species.
The Puerto Hondo stream salamander or Michoacan stream salamander is a mole salamander from the Trans-Mexican Volcanic Belt within the Mexican state of Michoacán.
The yellow-peppered salamander also known as the salamandra de Champala and the yellow-headed salamander, is a species of mole salamander native to areas at an elevation of 4900 ft around Santa Cruz, Rancho Malveste and Tapalpa in Jalisco, Mexico.
Ambystoma talpoideum, the mole salamander, is a species of salamander found in much of the eastern and central United States, from Florida to Texas, north to Illinois, east to Kentucky, with isolated populations in Virginia and Indiana. Older sources often refer to this species as the tadpole salamander because some individuals remain in a neotenic state. This salamander lives among the leaf litter on the forest floor, migrating to ponds to breed.
The mountain stream salamander or mountain stream siredon is a species of mole salamander that only lives in central México.