The Axolotl is unusual in nature because it retains its larval form into adulthood.

In fact, it becomes sexually mature in this state. This adaptation, known as neoteny, is often viewed as a backward step in evolution because it prevents the axolotl from living on land, and as a result, it can't colonise new habitats. However, it has led to the axolotl being quite successful in its native habitat, at least until the arrival of man. The Axolotl is carnivorous and has reasonably typical internal carnivore anatomy, with the main exception of the teeth. Its teeth are pedicalate (i.e. they're small stumps, like cones). With these it grips its food, manoeuvering it into position before swallowing it whole. It has a three-chambered amphibian heart (unlike the mammalian four-chambered heart), and, like all amphibians, it is poikilothermic (its body temperature is dependent upon its surroundings). One thing to note is that although they retain larval morphology, they do develop rudimentary lungs, and axolotls can be seen to occasionally rise to the surface, take a quick gulp of air to fill these lungs, and then quickly descend to the bottom once more. From my observations, I believe that the lungs develop shortly after the rear legs reach their full length. Axolotls are famous for their fabulous regeneration ability. Regeneration studies carried out around the world often involve the Axolotl. For example, a young axolotl that loses a foot to a sibling will usually grow it back over a period of a few weeks. This regeneration occurs via the formation of a "bud" at the end of the damaged appendage, followed by growth of the new foot. Entire limbs can be re ge ne ra te d an d ev en po rti on s of th e br ai n an d sp in e. T he diagram on the right is a summary of the development of the Axolotl, from egg to adult. The red line below each animal represents roughly 2 mm. The egg (stage 1) is typically amphibian. It comprises of the embryo, which measures about 2 mm in diameter, and also the surrounding layers of jelly. The jelly is the product of water and a substance that is secreted around the egg when it is laid. Stage 2 is the embryo prior to hatching. At this stage it is approximately 11mm in length. Stage 3 is the young larva, prior to the growth of limbs.

Larval axolotls tend to be transparent for their first few weeks of life, or until the skin has thickened and pigment cells have proliferated over the body, so the organs are quite visible, as well as the progress of any food in the digestive tract. After approximately two weeks, the larva reaches stage 4. Like all other caudates (newts and salamanders), and unlike anurans (frogs and toads), the front legs develop first, followed within a few weeks by the hind legs. At stage 5 the axolotl is, to all intents and purposes, a miniature adult. I once saw a friend's female axolotl that was about 43 cm (17 inches) in length. However, a large size for most axolotls would be 25-30 cm (10-12 inches). The average seems to be about 23-25 cm (9-10 inches). Adults tend to reach their full size after eighteen months to two years, the growth rate depending on how well they are fed and at what temperature they are kept. I've raised axolotls to 26 cm in 7 months, but that's exceptionally fast. The two pictures on the left and right are of a young adult female (hence the lack of eggs which would normally give the female a much more rounded appearance) . Notice that one of her gill branches is held in a different orientation to normal (the gill on the left actually). I have redrawn the diagram in Peter Scott's book (see Books and Links) of the Axolotl's digestive system on a real axolotl, in order to better illustrate the proportions and help us to visualise the internal arrangement of the organs. Imagine you're seeing through the animal. The cloaca/vent is on the underside. Sexual Maturity Male and female axolotls differ in a number of ways. Peter Scott gives a few pointers, but some of these seem to be

rather inaccurate and unreliable. There are only two reliable methods of distinguishing males from females. The first is that mature females tend to have very rounded bodies, due to the number of eggs present in their bodies. And the most reliable is that the sexually mature male's cloacal region is swollen, while that of the sexually mature female is considerably less so. See the photos at the bottom of this page. Males generally reach sexual maturity slightly earlier than females, and mature males tend to be more elongated, and longer tails than females. One point of note is that white, golden, and albino axolotls that have reached sexual maturity will have dark brown tips to their toes. The soles of their feet may even appear "dirty" (see the photo below). In wild type and melanoid animals the toe tips become slightly paler than the rest of the body, at maturity, but it is harder to see than in the lighter colour variants. Male axolotls go through a cycle where, although sexually mature, they may not have sperm available for mating. In the wild this seems to be determined to some extent by the seasons. However, in captivity it's less regular and may occur at any time of the year. It usually takes the male about 2-3 months to produce sperm and perhaps a further 2 months for the sperm to move into the vas deferens so that it is available for mating. This could mean that the overlap between having and not having sperm available could be a few weeks to a few months.

Dark toe tips on lightly coloured axolotls indicate sexual maturity.

The first photo below is of the cloacal region of a mature male and the second is of a mature female's cloacal region. The male is a golden albino, while the female is a melanoid albino. Their cloacas are circled in blue.

Male Albino

Golden

Female melanoid albino

Welcome! This web site is devoted to the Axolotl (pronounced Ax-oh-lot-ul), scientific name Ambystoma mexicanum. The site describes the Axolotl's background, biology, and care in captivity. You will also find information about the Tiger Salamander, Ambystoma tigrinum and Ambystoma mavortium spp., because these animals are closely related. Australians and New Zealanders frequently refer to the Axolotl as the Mexican Walking Fish, though the Axolotl is not a fish but an amphibian, a salamander, part of the order Caudata/Urodela. Because it's a salamander, it's part of one of the three branches of class Amphibia, which also includes the frogs and toads (the Anurans), and the mainly eellike order, Gymnophiona, which are also known as the Caecilians. Have a look at the Biology Page for a short guide to the Axolotl's body and characteristics. One common misconception is that axolotls and other salamanders are lizards or reptiles. In fact, amphibians are a completely separate group of animals. For example, did you know that reptiles and human beings have a four-chambered heart? Well amphibians have only three chambers. That's just one example of how appearance can be deceiving: salamanders might look like lizards, but they are very different indeed. This page is a brief introduction for those new to the Axolotl and salamanders. If you require specific information, you can search this site using the search facility at the top right of this page. Caudata.org also contains a wealth of axolotl information and it's a great place to buy axolotls or trade with other hobbyists. Caudata.org is the Internet's premier source of salamander and newt information and it places an emphasis on their maintenance in captivity. There is a very busy axolotl forum at Caudata.org, used by people just like you. I hope that you find this site useful, but most of all I hope you enjoy what you read and find here. If you're looking for information about metamorphosed axolotls, click here. Axolotls of various colours occur in captivity, including grey, shades of brown, leucistic (white with black eyes), golden albino, white albino,

as well as other varieties, such as the melanoid (a near-black animal). The normally coloured axolotl, the "wild type", can be near-black like the one in the group photo to the left, chocolate brown like the one in the site's logo, or even creamy in colour, and anywhere in between. There are even "piebald" axolotls in various colours, and a variety that is piebald in more than one colour, known as the "harlequin". You can learn more about how colour comes about and how it is passed on by taking a look at the Genetics Page. And why not take a look at the hundreds of photos of the weird and wonderful varieties of axolotls submitted by enthusiasts like yourself at the Axolotl Section of the Caudata.org User Photo Galleries? The name "Axolotl" comes from the Aztec language, "Nahuatl". One of the most popular translations of the name connects the Axolotl to the god of deformations and death, Xolotl, while the most commonly accepted translation is "water-dog" (from "atl" for water, and "xolotl", which can also mean dog). Prior to the growth of Mexico city in the basin of Mexico, the Axolotl was native to both Lake Xochimilco, and Lake Chalco. Of these two high altitude freshwater lakes, only the remnants of Xochimilco as canals can be seen today. Unfortunately many information sources mention these lakes as if they still exist (such as this illresearched article about a metamorphosed axolotl on the BBC News Web site). If only this were still the case: sadly it is rarely caught in the wild but at least the Axolotl is now on the CITES endangered species list. There have been efforts to breed and release the animal, in order to reestablish its numbers. However the location of the remaining waterways where the animal may live (located in the Mexico City metropolitan area) are likely to be very threatened by the city's continuing expansion and the days of the species surviving in the wild are surely quite limited. Fortunately, due to the importance of the Axolotl in scientific research, it is unheard of for them to be taken from the wild for that purpose because of the huge numbers bred in captivity each year. There are related Mexican Ambystoma species that also remain gilled as adults. These species are located in water bodies further from Mexico city and may have a slightly brighter future in the wild than the Axolotl. Despite its endangered status, the use of the Axolotl as a laboratory animal should ensure the species' survival, if only in captivity. It has long been known that the Axolotl is a worthy study due to its amazing healing and regeneration abilities. Normal wound healing in animals occurs through the growth of scar tissue, which is not the same as the original tissue, nor is it as robust. Normal wound healing also does not allow for most animals to re-grow a lost limb. However the axolotl is fully

capable of complete limb re-growth. The animal has the added scientific attraction of having especially large embryos, making it easier to deal with under laboratory conditions. Its embryo is also very robust, and can be spliced and combined with different parts of other axolotl embryos with a high degree of success. The Axolotl is a fascinating creature for a number of reasons, including its grotesque appearance, its ability to regenerate, and primarily the fact that it exhibits the phenomenon known as neoteny. Ordinarily, amphibians undergo metamorphosis from egg to larva (the tadpole of a frog is a larva), and finally to adult form. The Axolotl, along with a number of other amphibians, remains in its larval form throughout its life. This means that it retains its gills and fins, and it doesn't develop the protruding eyes, eyelids and characteristics of other adult salamanders. It grows much larger than a normal larval salamander, and it reaches sexual maturity in this larval stage. Another term to describe this state is "perennibranchiate". The animal is completely aquatic, and although it does possess rudimentary lungs, it breathes primarily through its gills and to a lesser extent, its skin. It is generally accepted that neoteny is a "backward " step in evolution, because the Axolotl is descended from what were once terrestrial salamande rs, like the closely related species, the Tiger Salamander, Ambystoma tigrinum and Ambystoma mavortium spp. (in fact, one likely theory suggests that the Axolotl is in fact a Tiger salamander off-shoot, as it can interbreed with that species with some success). Through some quirk of nature, a neotenous form developed and, probably due to environmental conditions, prospered. Neoteny is sometimes found in other amphibians, but tends to be caused by low levels of iodine (an essential element for animals to make thyroxine hormones, necessary for growth and development), or possibly by random genetic mutation. Research has also shown that very low

temperatures can suppress the production of these hormones, thus also inducing neoteny. In the Axolotl, neoteny is now totally genetic (click for more information on the Axolotl's genetics). When treated with hormones, the axolotl will usually begin to metamorphose, but in very rare cases it will metamorphose spontaneously, such as the metamorphosed wild type axolotl pictured here. The metamorphosed wild type axolotl bears a close resemblance to the Mexican race of the Tiger Salamander, Ambystoma velasci. There is a wonderful thread on the Caudata.org forum here about the metamorphosed axolotl in the photo.

Breeding Axolotls
Before reading this page, you may find it helpful to first read the Biology Page and the Genetics Page.

Basics
Axolotls can reach sexual maturity anywhere between 5 months and several years, depending on frequency and quality of food, and the water temperature and conditions in which the animals are kept. My personal record for a fully mature male is just under 6 months (at 25 cm or 10 inches). Axolotls generally begin to mature once they have reached about 18 cm (7 inches) in total length. Females tend to take a little longer to mature than males (usually a difference of a month or two). The Biology Page has a great deal of information about sexing axolotls and their sexual maturity. It is advisable that you don't attemt to breed axolotls until they reach at least 18 months of age. This gives them time to reach their full size (greater than 30 cm or 12 inches in many cases) and condition (a female ready to breed will be very round towards the end of the body when viewed from above). In my opinion it is safe to breed males at an earlier stage than females, because they have much less physical output during the mating process than females, and therefore there is less strain on their bodies. However, females should be prevented from breeding until they reach their full size. There is a very good reason for not breeding your female axolotl(s) too early. A female axolotl can lay in excess of 1000 eggs. Producing so many eggs is a strain on the animal's metabolism, and the body prioritises production of eggs over body growth while the animal is in conditions suitable for breeding. Since females may breed several times each year, as soon as the first batch of eggs are laid, the body attempts to produce new eggs to replace those that have been laid. Female axolotls may fall ill at this point unless due care is taken, and for a female that is still growing in length, the strain is increased. For the same reason, females that have recently bred should be kept away from males for at least a month, preferably two or three, in order for them to recover. From personal experience, I know that just because an axolotl breeds, it doesn't mean it is in good overall health.

Breeding methods
Most sources state that the breeding season for axolotls is from December to June. However, they can be bred at any time of the year, although most success is

reported in the early part of the year. The former Indiana University Axolotl Colony used changes in the length of light period to trigger spawning. Males and females are subjected to a decreasing "daylight" period over a few weeks, and then the duration of lighting is steadily increased. They then put a male and a female together and courtship behaviour usually follows. An alternate school (Peter W. Scott and some others) instead recommends a sudden change in temperature to trigger courtship behaviour. Keeping the pair separately for a few weeks at 20-22 C (68-71 F) and then transferring them both into a tank with a water temperature at least 5 C lower frequently triggers courtship behaviour. In fact, Scott recommends 12-14 C (54-57 F). My own experience, and that of some others, is that this thermal shock method usually just stimulates the male. In order for this procedure to be successful, the female most be receptive and ready to breed. In my experience, by keeping axolotls in a room that receives at least partial seasonal change in temperature and light period (if there is a window in the room), breeding will occur naturally, usually at least once before the peak of winter and once in the spring, if the animals are adequately fed. As mentioned briefly above, exposure to natural day length throughout the year by having the tank in a room that receives natural light is a good idea because light seems to have at least as an important role as temperature in simulating the seasons. A pair of axolotls kept in good conditions should breed at least once a year, albeit unpredictably. Axolotls may spawn for no obvious reason, at "odd" times of the year, as mine have done in the past. Axolotls may surprise you.

My Breeding Setup
The breeding setup that I use routinely is furnished with many plants (plastic plants are good because they don't rot, but I also use clumps of live Java Moss, Vesicularia dubyana). The plants are for the female axolotl to affix her eggs (pictured on the right is a female melanoid albino in the process of laying its eggs on plastic plant leaves). Slates or flat, rough pieces of stone should be placed on the bottom of the tank for the male to deposit its spermatophores. Spermatophores are packets of sperm - you can read more on these in the courtship and spawning section. Spermatophores will not readily stick to bare glass or plastic, so in order for mating to be successful the spermatophore must be stationary during courtship. It is usually a good idea to put the tank in a room where it will be left alone, so as not to disturb the pair.

Courtship and Spawning

Spawning is initiated by the male, who swims around, raising its tail and making vigorous writhing motions. The male nudges the female's vent occasionally and then leads her around the tank. The spermatophore is a common "device" in the salamander and newt world (an old one, about 12 hours old, is pictured beside this paragraph). It is a packet of sperm attached to the top of a cone of jelly. The male deposits between 5 and 25 of these around the tank and attempts to lead the female over them. She picks up the sperm cap (from one or more spermatophores) in her cloaca - fertilisation takes place internally. She may also nudge the male's vent, and this can lead to a prolonged "dance" around the tank. Between a few hours and two days later, she commences spawning, laying each egg individually. She will lay them on the leaves of plants, if available, but if not, she will place them about the tank, attaching them to rocks, pipes and any other object available. There may be between 100 and over a thousand eggs laid in one spawning, depending on the size of the female and if she is in optimal condition at spawning. After the female has finished laying, it's best to remove her and the male.

Hatching Eggs
Pictu red to the right are four 12hour old eggs. Note the lack of pigment this indicates that the mother was albino. Normal eggs are dark brown (there is a picture of two at the top of this page). An albino mother will lay white, pigmentless eggs. If the offspring are not albino, pigment will appear during embryo development. The eggs take about 2 to 3 weeks to hatch. Development seems to be optimal when the eggs are attached to plants. This is due to the circulation of water around the egg, aiding gaseous exchange. Assuming the eggs are fertile, the majority of the eggs should hatch if kept in well-aerated water. An air pump and air stone at one end of the tank will be

helpful, just be sure it doesn't cause vigorous water flow. At 20 C (68 F) the eggs should hatch after about 17 days. Have a look at the Embryo Series Photo Log for a daily photographic account of the development of some white eggs.

Requirements & Water Conditions

Overview
Water quality is an important factor in the health of the Axolotl. They are forgiving animals, but the correct care of axolotls in captivity is only possible under the right water conditions. Coupled with water conditions, we must also exercise caution when considering what to put in the water, be it ornaments or other tank mates. Topics covered on this page: 1. Other Tank Inmates, & Other Axolotls 2. Temperature & Cooling 3. Water Flow 4. pH: Acidity & Basicity/Alkalinity 5. Chlorine & Chloramines 6. Ammonia, Nitrite, & Nitrate 7. Water Hardness & Dissolved Salts 8. Water Changes & Final Words

Other Tank Inmates, & Other Axolotls

Here are the best three words of advice regarding other tank inmates: Just say no. Why? Well, let us suppose we would like to keep something with our axolotls, for instance, a fish, or another salamander or newt. The fish will invariably attempt to nibble on an axolotl's gills. After all, they're so attractive and feathery! The salamander or newt may try the same trick, particularly at feeding time. Let us suppose that our axolotl is bigger than the fish or salamander. It's quite likely we'll end up with one well fed axolotl! There is a simple rule that axolotls follow: if it moves and it's smaller than our axolotl, it'll end up in our axolotl's stomach. So, follow my three words of advice: Just say no! Young axolotls, less than 8 cm in length (3 inches), shouldn't be kept together in a confined space. If they are to be kept together, it would be wise to use an aquarium that allows them plenty of space. Young axolotls will nip each other's feet and gills, more so when very young. Even axolotls up to 15 or 16 cm (6 inches) may nip their tank mates, occasionally inflicting serious damage. This is particularly noticeable in wild type axolotls, since they are naturally more aggressive than homozygous colour mutants, and wild types will bite colour mutants in preference to other wild types. This behaviour is frequently observed by scientists and hobbyists alike and has yet to be fully explained.

Cannibalistic tendencies are much reduced in adult axolotls. However adults will predate upon considerably smaller axolotls, so keep different generations apart until they all reach adulthood. Lastly, consider keeping the sexes apart, unless you are prepared to deal with the occasional batch of eggs.

Temperature & Cooling

If you've been reading the pages in sequence, you will have read on the Housing Page that the optimum temperature for axolotls is between about 16 C and 18 C (60-64 F). Lower temperatures lead to sluggish behaviour, slower metabolism, and decreased appetite. Axolotls do not hibernate, so it is not helpful to cool them below 10 C, although they shouldn't suffer unduly if kept at these lower temperatures. Some hobbyists in temperate regions maintain axolotls in outdoor ponds. These ponds may even ice over during the winter. Provided the winter isn't particularly harsh or long, axolotls can do quite well under outdoor conditions. Obviously, a food source must be present during the rest of the year. Temperatures above 24 C (75 F) are very stressful to axolotls. Such temperatures cause metabolism to increase (the rate at which the body "works"), and consequently, an increase in appetite. However, the stress resulting from more than a day or two of exposure to these temperatures will quickly lead to disease and death. You can read more about axolotl diseases and their treatment on the Health Page. The first symptoms of heat stress in axolotls include refusal of food and/or the development of pale patches of mucus-like material on the skin (see the photo below).

This axolotl is exhibiting symptoms of heat stress: note the pale patch of mucus-like material on the head

If you are having difficulty maintaining the temperature of your axolotl's aquarium below 24 C (75 F), there are a few options to consider. The easiest short-term solution is to move the aquarium to a cooler part of the home. Remember, in every room, the temperature at ground level will be at least 1-2 C (3-5 F) cooler than high up on a shelf. In the summer, the same rule holds true for

a house as a whole: the lower rooms in the house (or the basement) will be cooler than rooms on higher floors. So, moving the aquarium may easily allow you to reduce the temperature in your axolotl tank from 26 C to 23 C. Such a temperature change could make the difference between your axolotl living and dying. Remember: if you plan to move an aquarium, make sure you remove most of the water first (just leave enough to cover the axolotl). Make sure there is a cushioned support at the destination (such as a polystyrene board), and ask someone to help you carry the aquarium. In more extreme cases, and in localities where high temperatures persist for more than a day or two at a time, you will need to take different measures. Some people use aquarium chillers (proprietary or home-made). I have no experience of chillers, but the people on the Axolotl Forum should be able to offer you appropriate advice. Another commonly used option, which should be safe to use for a week or two at a time is the "ice bottle". This is aplastic bottle of water, such as the 2 litre bottle (4 pints) used for carbonated soft drinks. The bottle is filled to between 80 and 90% capacity, and then frozen solid. Using a plastic bottle is important - a water-filled glass bottle will explode in the freezer. The bottle is then floated in the aquarium and the temperature monitored, and if necessary, the bottles are operated in relays. Larger aquariums would do better with larger bottles (2 L is a maximum though), smaller aquariums would do better with 500 mL bottles (about a pint). Using ice bottles safely is a skill: if the temperature changes too much, too rapidly, it can be more stressful to the axolotls than maintaining a stable, if high, temperature. Typically, ice bottles cause the temperature to crash into the teens Celsius (50s to early 60s F). After thawing, the temperature will begin to climb back into the mid-to-late 20s once again. Then, if we put another ice bottle back in to the aquarium, the process repeats itself. This is a fast and effective method to kill an axolotl. This is because the temperature changes so rapidly and does so repeatedly, stressing the axolotls to the extreme. So, if you intend to use ice bottles, do so carefully. Do not use them on very small tanks (under 60 cm/24 inches in the longest dimension). For a larger tank, using several smaller bottles in succession is probably safer than one large one and then another large one. And be prepared to switch bottles before the first one thaws out completely so that you can attempt to maintain a relatively constant temperature. To summarise tank cooling, the three things you need to worry about are: 1. How rapid is the temperature change? It shouldn't take place in less than 30-60 mins. 2. By how much does the temperature change? The larger the change, the more stress it will cause. 3. How stable is the new temperature? Fluctuations are quite stressful. 4. How often does the temperature change in 24 hours? If it's more than twice, you really should consider another option. There are no absolutes here. We just have to minimise the stress caused by these factors as best we can. I have a very clear opinion on temperature problems: I feel that if one can't maintain stress-free conditions for 360 of the 365 days of the year, one shouldn't keep axolotls. If your axolotl develops the pale mucus-like patches that the axolotl in the photo has developed, treat it as you would a fungal infection, but remember that

treating this condition is not going to be successful if you do not do something about the temperature too.

Water Flow
Water flow is usually caused by a filter or when you use an air pump on an aquarium. Output from a filter can cause significant flow and this is perhaps the most common cause of stress in axolotls. Excessive water flow will, sooner or later, lead to disease. The photos below depict an axolotl that has succumbed to the stress caused by excessive water flow.

This golden albino's forward-turned gills are typical of an axolotl stressed by flowing water.

A curled tail end is a sure sign of a stressed axolotl.

Here are several approaches to minimise concentrated water flow, such as that typical of a filter's outflow:

1. If the filter has a built-in facility to reduce the flow, use it. 2. Use a spray bar. Either make one from a piece of tubing that has had holes drilled in it, or buy a proprietary spray bar kit, the longer the better. Orientate it against the glass so that water force is lessened. An aquarium equipped with such a system is depicted and described on the Housing Page. 3. Angle the filter so that the water flow is aimed at a glass side of the tank. 4. Angle the filter so that the water flow is directed upwards towards the water's surface. 5. Partially obstruct the filter's output using a piece of filter wool, or a home-made device, such as a piece of filter tube. Be careful not to obstruct the output nozzle too much, as this may cause your filter's motor and impellor to wear rapidly. 6. Consider using a smaller or different filter.

pH: Acidity & Basicity/Alkalinity

pH stands for "power of hydronium". Water (H2O) exists in a constant equilibrium with itself: H2O OH- + H+. The hydrogen ion (H+) is very small and strongly hydrated (meaning it attaches itself to another water molecule), essentially existing as the ion hydronium (H30+). pH is a convenient way of expressing the hydrogen ion concentration. In actual fact, it's the negative logarithm of the hydrogen ion concentration. The pH scale ranges from 0 to 14. A solution (i.e. water containing something else) with a pH of less than 7, is said to be acidic. A pH greater than 7 is considered basic (also known as alkaline). And a pH of exactly 7 is considered neutral, neither acidic nor basic. The further the pH is from 7, the stronger the acidity or alkalinity/basicity of the solution. You can obtain simple-to-use pH test kits from your local aquarium retailer. Most municipalities treat their water so that it is within a few degrees of pH 7 (neutral). My local water is 7.2 after treatment. For axolotls, a pH of 6.5 to around 8.0 is acceptable, but 7.4 to 7.6 is probably ideal. pH can affect the toxicity of ammonia and this is discussed below. If you have particularly acidic or basic water, you can adjust the pH using the kits sold by your local aquarium retailer, or you can basicify the water by adding salts, as described in the Water Hardness & Dissolved Salts topic, below.

Chlorine & Chloramines

Chlorine is a nasty green gas, denser than air, that was used in the trenches in World War I. It was dangerous enough to kill people by reacting vigorously with lung tissue. This alarming property is used to kill bacteria in municipal water supplies around the world today, which makes the water safe for human consumption. Ammonia (some more info below) is another noxious gas, and many municipalities also add it to their water. Chloramines (NH2Cl, NHCl2, and NCl3) result from a combination of chlorine and ammonia. Unfortunately, aquatic animals are not top of the list of concerns of Municipalities. These substances make it necessary to use de-chlorinator in the water before it can be used in the aquarium. Alternatively, water can be left to stand for 24 hours or more to let the chlorine dissipate, but be aware that it takes a lot longer for chloramines to reach safe levels in water if just left to stand. De-chlorinators also remove chloramines. Most de-chlorinators also remove traces of metals such as iron, mercury, copper, lead, cadmium, and manganese. Metals may be found in water due to the pipes through which it must travel.

I use Aquarium Pharmaceuticals' Stress Coat, which also contains aloe vera, a plant extract that, although slightly toxic, replaces the natural slime coat that fish have. It seems to have a similar benefit for axolotls and it's not harmful. Hagen's Amquel also contains aloe vera. Aloe vera also has a slight anti-biotic effect. Every time you change the water using tap water, be sure to treat it first for chlorine and chloramines. If for some reason you're unsure of the presence of chlorine in the water, you can obtain a chlorine test kit from your local aquarium shop.

Ammonia, Nitrite, & Nitrate

These three substances, as discussed on the Housing Page, are part of the biological filtration cycle in your aquarium. Ammonia, NH3, the main waste product produced by your axolotls, is very toxic in its unionised form (NH3 as opposed to NH4+). A low pH means a higher concentration of H+ ions, which in turn results in a higher degree of ionisation of NH3 to NH4+. Conversely, a high pH means that unionised ammonia, toxic NH3, is the main form of ammonia. What this means in English is that the higher the pH, the more toxic the ammonia. Ammonia can kill, and at a pH of 8 or more, it kills even more effectively. Water temperature can also affect its toxicity, a higher temperature resulting in a higher toxicity. Unless you're extremely proficient with maintaining aquarium systems, a periodic ammonia test is advisable. Even the most experienced hobbyist should occasionally test for ammonia. Nitrite, NO2-, is produced from ammonia by the bacterium Nitrosomonas. It is not as toxic as ammonia, but should also be tested for regularly. Again, test kits are available and should be used about as regularly as ammonia tests. Nitrate, NO3-, is the least toxic of this family of nitrogenous compounds. It is produced from nitrite by the bacterium Nitrobacter. Although it should be tested for, regular water changes and plants in the aquarium will keep the levels of nitrate in check. Although not toxic at low levels, if let build up through lack of water changes, it too can be dangerous, and high levels usually lead to blooms of algae.

Water Hardness & Dissolved Salts

The degree of water hardness can be thought of as the amount of dissolved salts in the water. There are many kinds of salt apart from the one people use on their food. Some commonly encountered salts include epsom salt (MgSO4), baking/bread soda (NaHCO3), and "Low-Sodium" salt (KCl) used by people as an alternative to ordinary salt (NaCl). If you live in a hard water area, the chances are that you've seen limescale in your kettle or pipes. This is caused by the deposition of dissolved salts of calcium and magnesium on the heating element of the kettle and the inside surface of the pipes. Soft water contains little dissolved salts, while hard water contains significant amounts. There tends to be a correlation between soft water and acidic pH, and hard water and alkaline pH. This is because these minerals affect the chemical equilibrium in the water (discussed in a simple form in the pH section). Axolotls prefer somewhat hard water, and those that live in soft water will often suffer from temporary anaemia - the animal becomes pale and its gills lose their colouration for a few minutes or hours. This is not a dangerous condition but it can be prevented by supplementing the hardness of the water with added salts. Currently, I live in an area in which the water is a little on the soft side. I regularly supplement it with added salts. Laboratories use one of two types of salt mixture (each is known as a medium) in their water: Holtfreter's solution and Steinberg's solution. The ingredients and proportions used in each are detailed in

the table below, together with the "recipe" I personally use. This table includes a brief version of a table found in "Developmental Biology of the Axolotl". Medium Components to make 100% concentration (per litre of water) Holtrefeter's Solution NaCl KCl CaCl2 NaHCO3 NaCl KCl Ca(NO3)2.4H2O MgSO4.7H2O Tris (a sort of buffer) Add HCl to pH 7.4 NaCl MgSO4.7H2O NaHCO3 Weight in grams 3.46 0.05 0.1 0.2 3.4 0.05 0.08 0.205 0.56

Steinberg's Solution

John's Solution

1 0.1 0.1

Indiana University Axolotl Colony use a modified Holtfreter's solution which leaves out the NaHCO3 and uses MgSO4in the same weight. Typically, 40% or 50% is used for adults, and 20% for embryos. I have also used another modified Holtfreter's mix at 50% concentration. The modification is that I added MgSO4 in the same weight as NaHCO3 instead of replacing NaHCO3 as done by Indiana University Axolotl Colony. As a more practical straight-forward alternative to making up your own solutions, you can purchase kits for adjusting the hardness of water at your local aquarium outlet (they also directly adjust pH and are sold as such). If you want to get the salts for the solutions mentioned above and you have problems finding them, I have seen them at the Chemistry Store, though I have never purchased anything from that company.

Final Words
To sum up, water conditions are an important consideration when keeping axolotls. Remember to regularly replace 20% of the water each week. Depending on the size of the aquarium or container, and whether or not it is filtered, these changes may need to be more or less frequent. Do occasionally test the water for its different constituents and pollutants. Taking care of water conditions will help to prevent disease outbreaks. It will also keep your axolotls healthy and comfortable, and it will encourage breeding.

Rearing Axolotls
Introduction
I first bred axolotls successful ly in the late 1990s. Thankfully I was very successful . By following the advice and directions on this page, you should be too. Axolotl larvae are surprisingl y tough, as far as newt and salamande r larvae go. By the way, "larvae" is the plural of larva, the term used to describe axolotls (and other newts and salamanders) that have yet to develop all four legs. Due to their toughness, if we follow the guidelines on this page we shouldn't have any major problems rearing the larvae. We should bear in mind that there will always be a few casualities when rearing large numbers of larvae. Some larvae will have "unseen" genetic problems, some will succumb to stress that others will survive, and some will just be plain unlucky. For the beginner, it is most advisable to attempt to rear only a few larvae at a time. It's better to give away or, though it may seem unpleasant, cull most of the eggs, giving the remainder all of your attention and resources, than to try to rear many hundreds of larvae for the first time and see them all die due to stretched resources, bad water conditions, or other problems frequently encountered by beginners. If you have too many eggs, you will find plenty of people willing to accept them from you on the Caudata.org Forum. Likewise, if you are looking for axolotl eggs or young axolotls, you will find them there too. Unusual colours, like the melanoid axolotl larva we can see at the top of this page developing in its egg, are often available through the Forum.

Initial care of eggs

Finding axolotl eggs, or seeing a female spawning, often occurs when we least expect it, or at least not quite on the day we expect. It's always a good idea to keep an eye out for old spermatophores in the parents' tank so that we have some warning that there may soon be many new mouths to feed. On finding eggs, we must decide if they are to be removed, or the parents are to be removed and the eggs left behind. Many first time breeders opt for the latter choice, but axolotl eggs are quite tough, so the eggs can usually be removed from a tank and moved to another without issue. They can even be removed from rocks if you're careful: there is generally a point of attachment to the egg's outermost jelly layer, which can be sliced with a finger nail, thus freeing the egg. Each egg is surrounded by several layers of jelly, so don't be too afraid. Eggs attached to plants are great, because they will stick to the plants. The plants can be moved carefully to the new tank, thus avoiding all hand contact with the eggs themselves. Once you have made your choice (to move or not to move the eggs), it's time to make sure we have correct conditions for the eggs to hatch. Pictured to the left is a small aquarium (45x20x25 cm, 18x8x10 inch) solely for hatching eggs (I opted to hatch just 100 in this case). Remember to use dechlorinated water. Water shouldn't be soft: do you get limescale in your water pipes or kettle? If you get limescale, your water isn't soft. If you're not sure, you should read theRequirements Page for more information about dissolved salts and their importance. Our next concern is water temperature. Keeping the eggs at a warm temperature (to an absolute maximum of 25 C /77 F) will cause the eggs to hatch sooner (generally in less than 14 days), whilst a lower temperature (such as 18 C / 64 F) will result in them taking perhaps more than 20 days. Being able to manipulate the time taken to hatch can be very useful if you need a while to secure a food source for the newly hatched axolotls. By lowering the temperature you can give yourself an extra week or two in order to acquire some brineshrimp eggs, culture your Daphnia, or order some microworms from a dealer.

Food

Speak ing of food for little mouths, it's time we considere d our options. If you bred your axolotls deliberate ly, the chances are you are already setup as far as providing tiny live food items is concerned. More often than not though, we need to acquire these food items in a hurry. Newly hatched axolotl larvae can vary in size from 10-13 mm (roughly 0.5 inch). On hatching, they will still possess some egg yolk in their stomachs (the white substance that should be quite visible to the naked eye). Until this is used up they will be motionless and won't require food. Within 24-72 hours after hatching, they will require food. From this point until they reach approximately 20 mm in length (a little under an inch), their diet must consist solely of livefood items of a very small size. They will ignore dead food until they have grown significantly. This is because, instinctively, young larvae respond to prey movement alone. It is not until later that smell will play an important role in feeding. If you cannot meet this demand for tiny live food, your larvae will starve to death, unless they eat each other (very hard to do for newly hatched larvae). For newly hatched larvae, the live food choices are: newly hatched brineshrimp (Artemia), smallDaphnia (see the photo on the right) or Moina, and microworms (microworms are not ideal and won't be well received until the axolotl larvae develop their front legs). There is an excellent article about tiny live foods at the Caudata Culture web site, and I recommend you read that before proceeding. There is a good article about foods for newly hatched salamanders over at Caudata Culture. Here's a link to that article. My preference is to feed young Daphnia, with some microworms as back up. I culture both, myself. I wouldn't recommend feeding wildcaught Daphnia or Daphnia obtained from an unknown source when feeding axolotls, or even fish. They have been known to carry diseases if taken from nature or sources containing other animals. Culturing your own, however, removes most of this danger, and although not as small or quite as nutritious as newly-hatched brineshrimp (young Daphnia are approximately two and a half times the size of newly hatched brineshrimp), youngDaphnia are a good first food for axolotl larvae, and they can be free! I feed mine on crushed trout pellets (the same ones I feed the axolotls). The Daphnia don't actually eat the pellets but rather the bacteria that grow as a result of the leeching of nutrients from the pellets into the water.

Microworms aren't the most nutritious food - axolotls fed solely on these will grow very slowly, and won't be received well by larvae that have yet to develop their front legs. They are a good food when you are in need though.

After Hatching
If you'd like to see a photographic diary of my first albino embryos as they developed before hatching, then look at the Embryo Photo Series Page. On hatching, most axolotl larvae are about 11 mm (less than half an inch) in length. Just before hatching I normally lower the water level to about three or four centimetres (an inch and a half) so that any daphnia in the tank are concentrated near the larvae for easy access. Once most of the larvae have hatched, it is a good idea to tear the jelly coat of those eggs that haven't hatched yet in order to free the larva inside. This can be done with a sharp forceps or narrow scissors that is inserted into the egg and then the prongs/blades are pushed apart. As explained earlier, at this stage the larvae usually won't eat because they're still absorbing the yolk from their eggs. You should be ready with your chosen first food. I normally have some Daphnia in the tank a few days before hatching in order to have lots of little tiny young Daphnia present when needed. The larvae can get air bubbles in their stomachs if not fed early enough, but these will be expelled once they start to feed. Very young larvae can be kept together without much risk of cannibalism since this really only begins once the front legs develop. When very young they should be fed frequently (once or twice daily). If kept at about 20 C (68 F) they should reach about 1.5 cm in length within a week. The larva pictured on the right is four days old and 14 mm in length. It is D/D M/m a/a (see the Genetics Page for more information about colour). If feeding brineshrimp, at least some of the water should be replaced each day because newly hatched brineshrimp die quickly in fresh water and can foul the water in a matter of hours. As mentioned above, Daphnia will live in the tank with the larvae until eaten, so occasional water changes are fine (partial water changes a few times a week). Larvae never grow at the same rate, so it is advisable to divide them up according to size once they begin to reach about 2 cm in length. At this size the larvae become more cannibalistic, since they snap at anything that moves and at that size can damage their siblings (missing limbs and gills are an obvious sign). The larva below-left is a D/D m/m a/a sibling of the larva above-right but at 7 days old. It is 19 mm in length.

Cannibalism is a natural tendency, and studies have been carried out on the cannibalistic variants of salamander larvae. If left to cannibalise their siblings, they actually develop different morphology (shape and appearance). This is most noticeable in the shape of the head and the teeth. James Petranka's Book, "Salamanders of the United States and Canada" discusses cannibalism in detail, especially in the case of the Tiger Salamander, with some excellent photography, and William Duellman and Linda Trueb's book, "Biology of Amphibians", contains some excellent diagrams of the head morphology of the different morphs of the tiger salamander, a close relative of the Axolotl (see the Books and Links Page). In young larvae, particularly melanoid albino larvae like that on the left, you can actually see the liver, heart, stomach and intestines right through the skin. To minimise cannibalism, Peter Scott recommends that the tank is heavily planted and that light levels are lowered. A more reliable method is to reduce the numbers as much as possible in each container, but appetite does seem to decrease in low light. It's been my experience that front legs begin to develop once the larvae reach 20 mm (within about 9 days of hatching at about 22 C) and the hind legs begin to develop at the end of the third week. This all depends on temperature and feeding. The lungs first develop around the time that the rear-legs develop (yes, axolotls do possess lungs as well as gills). As the larvae grow they need to be thinned out, and any deformed or markedly inferior larvae (such as those that don't re-grow limbs and gills easily) should be euthanised/culled. There has been a lot of inbreeding in axolotls over the many years they have been kept in captivity, and this means that the likelihood of defects and oddities developing is greatly increased in this species. Using the conditions just described, at 7 days, I still feed young Daphnia to the larvae, and most of the larvae are about 18 mm in length. At 9 days, most larvae have very noticeable front limb buds and most are about 20 mm in length. This is a high growth rate and I attribute it to the daily feeding of young Daphnia and the temperature at which they are kept (22 C). The golden albino larva on the right is 10 days old and is 22 mm in length. The arrow is pointing at a limb bud. At this size they eat adult Daphnia (which are 2.5-3 mm in length). At 25 mm (1 inch) I try to maintain a maximum of 25 larvae per 45x20x25 cm aquarium. Remember, if you have too many larvae, either find new homes for them or cull them to prevent disasters. At 25 mm, hind leg buds should be quite apparent on most larvae and some should have well-developed front legs. At this point it is possible to begin feeding the larvae with thawed frozen bloodworm (see the Feeding Page). These can replace partially, or

fully, the small live foods you've been using up to this point. If you've been keeping up with the appetites of the larvae, you will probably be running low on live food any way. The photo on the left shows a 32 mm melanoid albino larva (D/D a/a m/m). At this stage its front legs are almost fully grown and hind leg buds are quite visible from the side (not visible in this photo though). Note the lack of iridophores (shiny pigment) in this larva, due to it being homozygous for the melanoid gene m. Xanthophores are almost absent in this larva, another effect of the melanoid gene. At 36 mm, the colour phenotype of each larva is generally very apparent - this means you can tell different colour types apart quite readily. The front legs are now fully grown and the hind legs are growing. At 40 mm (1.5 inches), the hind legs are about half grown. At this size, cannibalism can be a noticeable problem, so do try to keep the numbers in each tank at a low number (in the tank mentioned above, try to keep less than tens 40 mm larvae together). Feet and gills will still regrow rapidly on larvae at this size, however the older they get, the slower regeneration occurs. Another fact to be aware of is that wild type larvae tend to be more aggressive than non-wild types. Wild types will also show a tendency to attack non-wild types rather than other wild types. You can use biofoam filters to try to reduce the number of water changes. These aren't great filters for animals like axolotls, but for larvae they are quite safe (no risk of sucking up larvae). If you know what you're doing, you could use a canister/external filter on a large aquarium. You will need to make sure that the input is protected from sucking up larvae, and the output flow spread out so that it doesn't stress the larvae (excessive water flow is a common cause of stress in axolotls, leading to disease).

Juveniles
The larvae in the photo to the right are about 50 mm (2 inches) in length. The hind legs are not fully grown at this point but are quite visible. At this size they are more than capable of

taking the 3 mm axolotl pellets. Once the hind legs are fully developed, the larvae are now miniature versions of the adults, so they should now be called juveniles, or sub-adults. Growth should be steady, and juveniles are quite capable of taking small pieces of earthworm (or whole earthworms that are small in size). If you find frozen bloodworm expensive and messy, it's worth knowing that at about 2.5-4 cm (1-1.5 inches) most laboratories begin feeding with axolotl pellets. Be sure to gradually changes foods though, rather than doing so abruptly: it may take a few days before the animals begin to take the "new" food. If you would like to know more about the stages of larval growth, look at the Biology Page. There is a nice diagram of the stages of axolotl growth, from egg to juvenile. The photo below shows 15 cm (6 inches) juveniles of various colours.

Genetics and Colour

Overview
The Axolotl is studied the world over for several reasons. All of the traits which make it so suitable for study, as for all living things, are dependent upon its genes. This page will attempt to give a brief overview of axolotl genetics, mainly from the viewpoint of the hobbyist, who tends to be most interested in colour. Axolotls have 28 chromosomes per cell, in fourteen pairs. Humans have 46 chromosomes in 23 pairs. A chromosome is a thread-like structure composed of DNA and protein. The length of a chromosome is made up of many units of DNA called genes. Each gene has a special place on a chromosome and the position which it occupies is called the locus of that gene.

When an axolotl reproduces, the sperm from the male (which contains 14 chromosomes) and the egg from the female (which also contains 14 chromosomes) fuse to form the zygote, the first cell of the new axolotl. So each new cell of this new animal has 28 chromosomes. However, in the production of gametes (the sperm and egg cells) via the process known as meiosis, small exchanges of parts of the chromosomes take place (known as "crossing-over"), as well as a random allotment of chromosomes from the mother or father's own parents to each gamete (which means a gamete could have 2 maternal chromosomes and 12 paternal, or any other combination). So when axolotls reproduce, each new larva is a genetically distinct individual, different from its siblings and its parents. This is the essence of genetic variation. An animal's genotype is what its genes "say" it is, and its phenotype is the result of the gene, its expression. Mutant animals are those with genes differing from what is accepted as normal.

Colour
The colour of axolotls is dependent upon pigment cells called chromatophores. These cells are melanophores (containing eumelanin, a black-brown pigment), xanthophores (containing carotenoids and pteridines, yellow and reddish pigments) and iridophores (containing crystalised purines, which impart a shiny iridescence). Each cell in an axolotl, as stated above, contains 14 pairs of chromosomes. Every characteristic of the animals is coded for by genes on pairs of chromosomes. The genes for the pigment cells are inherited independently of one another, and there is no known linkage to any other genes. So, each pigment type is coded for by two different genes, one on each of a pair of chromosomes. These contrasting genes that code for the same characteristic are known as alleles. A pair of alleles is written like this: X/x. A capital letter means that gene is a dominant gene, as opposed to the small letter, which means that gene is recessive. For example, the allele that controls albinism could be found in an axolotl in one of the following combinations: A/a, A/A, or a/a. If the animal was A/a, because a is recessive and A is dominant, the animal's phenotype wouldn't be albino, but it would still carry the gene for albinism (since it has an "a"). Since it carries both "A" and "a", it is known as "heterozygous". If the animal had the A/A combination, its phenotype wouldn't be albino, and it wouldn't carry the gene for albinism (both genes being the same, it is called "homozygous" for "A"). If it were homozygous for "a" (i.e. a/a), the animal's phenotype would be albino. Since "a" is recessive, both alleles need to be "a" in order for albinism to be expressed in the phenotype. Albinism results in a lack of eumelanin (the dark pigment). In axolotls, it also results in an increased number of xanthophores (yellow pigment cells). In the same way that a/a results in a lack of eumelanin, m/m (melanoid) results in a lack of iridophores. Such animals are very dark, with no reflective pigment cells at all. M/m or M/M would result in normal iridophore development. Animals homozygous for "ax" (i.e. ax/ax) are axanthic, meaning they have no visible xanthophores or iridophores. Such animals are almost as dark as melanoids. Animals homozygous for both the albino gene and the axanthic gene appear to be slightly off-white (yellowish). The following table summarises the colour genes.

You may have noticed the "d" gene. This gene is a developmental mutant and not a pigment mutant like the others. Animals homozygous or heterozygous for "D" produce large numbers of yellow xanthophores. In combination with melanophores, we get the wild type colouration (dark brown/olive-green). However, in animals homozygous for "d", the normal pigment cells are produced, but they never migrate off the neural crest of the embryonic animal, resulting in the white phenotype. It is important to realise the this animal is not albino. This phenotype is white, but has dark eyes. It is known as leucistic. Simple albinism in axolotls leads to a yellow/golden animal, with red/pink eyes. In order to produce a white albino, the animal must have the d/d genotype in combination with the a/a genotype. Melanoid albinos (m/m with a/a) are also white animals with pink/red eyes. This can make initial identification of a white albino's phenotype difficult to determine for the novice.

Colour types
Here are photos and descriptions of some of the commonly available colour variants and an explanation of the genetics behind them. Wild Type Wild types vary somewhat in exact colour, but are generally a shade of dark brown with black, yellowish, and shiny patches/speckles. The phenotype of wild type animals is dark, non-melanoid, non-albino and non-axanthic. In the photo below, the animal on the right is a wild type female. This is the typical wild type appearance. You may also encounter tan or very lightly speckled variants. Wild types are not homozygous for any of the colour mutations.

Melanoid In the above photo, the animal on the left is a female melanoid. As described earlier, melanoids lack the "shiny" pigments (crystalised purines). The amount of yellow present is also much reduced. In contrast, the number of melanophores (dark pigment cells) is greatly increased, resulting in a black animal. The easiest way to determine if an animal is melanoid or a dark wild type is to look at the eyes: nonmelanoids have a shiny ring around the pupil of the eye, while melanoids (and melanoid albinos) do not. Melanoids are homozygous for "m" only (i.e. m/m), but they may be heterozygous for other colour mutations. White (also called Leucistic) Commonly known as leucistic, the phenotype is d/d, non-melanoid, non-albino and non-axanthic. Here's a picture of a large adult female. Notice the black eyes and small number of melanophores on the head and back which indicate that it is not an albino. White axolotls with black eyes are not albino. While d/d prevents the axolotl's pigment cells from migrating off the top of the animal, this does not

necessarily mean that all leucistics will possess colour cells on the the head and back - look at the eyes to be certain.

Albino Phenotype is albino a/a (lacking melanophores). There are many kinds of phenotypically-different albino axolotl. Here are some of them. The first is the golden albino (D/D a/a or D/d a/a). It has normal migration of pigment cells, but lacks melanophores, hence the yellow/gold appearance.

The second kind of albino I will mention is the axanthic albino. It has normal pigment cell migration but is homozygous for the albino gene and the axanthic gene (a/a and ax/ax), meaning it lacks melanophores, xanthophores and iridophores. It is almost white, but becomes yellow with age due to the accumulation of riboflavins from its diet. Here's a picture of one of the former Indiana University Axolotl Colony's albino axanthic specimens

The third kind of albino is the white albino. It is homozygous for "d" and "a" (d/d and a/a). The photo below is one of the former Indiana University Axolotl Colony's white albinos. Note the presence of iridophores (shiny pigment cells) in its gill branches.

The fourth kind of albino that I will discuss is the melanoid albino. It is homozygous for "m" and "a" (m/m a/a). This is a male axolotl. While a non-albino melanoid would be black, the combination of melanism and albinism "removes" all pigment except a tiny hint of yellow xanthophores on the head and back.

Conclusion
Axolotls come in many different colour variations. Axolotl breeders often produce an odd offspring whose phenotype defies what the breeder knows to be its genotype. Piebald axolotls (not just on the top of the body like a leucistic), yellow leucistics with black spots, and the harlequin (orange and black patches on a white axolotl) are just a few examples of what chance can present. You can see the huge variety of axolotl colour variations in Caudata.org's User Photo Gallery.