Axolotl

The axolotl (pronounced ACK-suh-LAH-tuhl) also known as a **Mexican salamander** (*Ambystoma mexicanum*) or a **Mexican walking fish**, is a neotenic salamander, closely related to the tiger salamander. Axolotis have the rare trait of retaining its larval features throughout its adult life. This condition, called neoteny, means it keeps its tadpole-like dorsal fin, which runs almost the length of its body, and its feathery external gills, which protrude from the back of its wide head.

As of 2010, wild axolotls were 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. They are currently listed by CITES as an endangered species and by IUCN as critically endangered in the wild, with a decreasing population. Axolotls are used extensively in scientific research due to their ability to regenerate limbs. Axolotls were also sold as food in Mexican markets and were a staple in the Aztec diet.

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 city is currently working on conserving axolotls by building "axolotl shelters" and conserving remaining and potential habitats for the salamanders.

## Description

A sexually mature adult axolotl, at age 18–24 months, ranges in length from 15–45 cm (6–18 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.

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.

Axolotls have barely visible vestigial teeth, which would have developed 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.

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 four mutant colors are leucistic (pale pink with black eyes), albino (golden with gold eyes), axanthic (grey with black eyes) and melanoid (all black 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, animals that are double recessive 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.

**Differences From Other Salamanders**

Found exclusively in the lake complex of Xochimilco (pronounced SO-chee-MILL-koh) near Mexico City, axolotls differ from most other salamanders in that they live permanently in water. In extremely rare cases, an axolotl will progress to maturity and emerge from the water, but by and large, they are content to stay on the bottom of Xochimilco’s lakes and canals.

Close relatives of the tiger salamander, axolotls can be quite large, reaching up to a foot in length, although the average size is closer to half that. They are typically black or mottled brown, but albino and white varieties are somewhat common, particularly among captive specimens.

## Habitat and ecology

The axolotl is only native to Lake Xochimilco and Lake Chalco in central Mexico. Lake Chalco no longer exists, as it was artificially drained to avoid periodic flooding, 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), though it may fall to 6 to 7 °C in the winter, and perhaps lower.

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 its 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. The axolotl is carnivorous, consuming small prey such as worms, insects, 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.

## Neoteny

Axolotls exhibit neoteny, meaning that they reach sexual maturity without undergoing metamorphosis. Many species within the axolotl's genus are either entirely neotenic or have neotenic populations. In the axolotl, metamorphic failure is caused by a lack of thyroid stimulating hormone, which is used to induce the thyroid to produce thyroxine in transforming salamanders. 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.

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 deficiencies in food and, in particular, iodine, which causes cretinism and dwarfism due to hypothyroidism, as it does in humans.

Unlike some other neotenic salamanders (sirens and *Necturus*), axolotls can be induced to metamorphose by an injection of iodine (used in the production of thyroid hormones) or by shots of thyroxine hormone. The adult form resembles a terrestrial plateau tiger salamander, but has several differences, such as longer toes, which support its status as a separate species.

## Population Decline

Axolotls are long-lived, surviving up to 15 years on a diet of mollusks, worms, insect larvae, crustaceans, and some fish. Accustomed to being a top predator in its habitat, this species has begun to suffer from the introduction of large fish into its lake habitat. Natural threats include predatory birds such as herons.

Populations are in decline as the demands of nearby Mexico City have led to the draining and contamination of much of the waters of the Xochimilco Lake complex. They are also popular in the aquarium trade, and roasted axolotl is considered a delicacy in Mexico, further shrinking their numbers. They are considered a critically endangered species.