Axolotl

## Description

The axolotl (; from Classical Nahuatl: āxōlōtl (listen)), Ambystoma mexicanum, is a paedomorphic salamander related to the tiger salamander. The species was originally found in several lakes, such as Lake Xochimilco underlying Mexico City. Axolotls are unusual among amphibians in that they reach adulthood without undergoing metamorphosis. Instead of taking to the land, adults remain aquatic and gilled.

Axolotls should not be confused with the larval stage of closely related tiger salamander species (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, 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 listed as critically endangered in the wild, with a decreasing population of around 50 to 1000 adult individuals, by the International Union for Conservation of Nature and Natural Resources (IUCN) and are 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.

## Appearance

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.

## Distribution

The axolotl is only native 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.

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.

## Behavior

Axolotls may be active at any time of the day and lead a solitary lifestyle. They locate food by smell, and will "snap" at any potential meal, sucking the food into their stomachs with vacuum force.

## Diet

Axolotls are carnivores (piscivores, molluscivores, vermivores, insectivores). They eat small prey such as mollusks, worms, insects, other arthropods, and small fish.

## References

Axolotls are polygunandrpus (promiscuous) meaning both the males and the females of this species mate with multiple partners during the breeding season. Axolotls usually breed from March to June. The female lays 100-300 eggs in the water attached to substrates. Eggs hatch after 10-14 days and the young (larvae) swim away on their own. They become reproductively mature at 18-27 months of age.

## Population

### Population threats

The main threat to this species is the loss of its native habitat. Axolotls occur only in the Mexican Central Valley and their native habitat is now limited to Lake Xochimilco as a result of the expansion of Mexico City. Lake Xochimilco is not a large body of water, but rather a small series of artificial channels, small lakes, and temporary wetlands. Lake Xochimilco has poor water quality, caused by the region’s aquaculture and agriculture demands. It is also maintained by inputs of only partially treated wastewater. In addition, the intensive use of pesticides from agriculture around Lake Xochimilco causes runoff into the lake and a reduction of habitat quality for axolotls. The pesticides used contain chemical compounds that studies show to sharply increase mortality in axolotl embryos and larvae. Of the surviving embryo and larvae, there is also an increase in morphological, behavioral, and activity abnormalities. Another factor that threatens the native axolotl population is the introduction of invasive species such as the Nile tilapia and Common carp. These invasive fish species threaten axolotl populations by eating their eggs or young and by out-competing them for natural resources. The presence of these species has also been shown to change the behavior of axolotls, causing them to be less active to avoid predation. This reduction in activity greatly impacts the axolotls foraging and mating opportunities.

### Population number

According to IUCN Red List, the total population size of the axolotl is between 500 and 1,000 mature individuals. Currently, this species is classified as Critically Endangered (CR) on the IUCN Red List, and its numbers today are decreasing.