Proteus anguinus

The olm or proteus (Proteus anguinus) is an aquatic salamander in the family Proteidae, the only exclusively cave-dwelling chordate species found in Europe. In contrast to most amphibians, it is entirely aquatic; it eats, sleeps, and breeds underwater. Living in caves found in the Dinaric Alps, it is endemic to the waters that flow underground through the extensive limestone bedrock of the karst of Central and Southeastern Europe, specifically southern Slovenia, the basin of the Soča River (Italian: Isonzo) near Trieste, Italy, southwestern Croatia, and Bosnia and Herzegovina. Introduced populations are found near Vicenza, Italy, and Kranj, Slovenia. It is also called the 'human fish' by locals because of its fleshy skin color (translated literally from Slovene: človeška ribica, Macedonian: човечка рипка, Serbo-Croatian: čovječja ribica/човечја рибица), as well as 'cave salamander' or 'white salamander'. In Slovenia, it is also known by the name močeril (from *močerъ 'earthworm, damp creepy-crawly' < moča 'dampness'). It was first mentioned in 1689 by the local naturalist Valvasor in his Glory of the Duchy of Carniola, who reported that, after heavy rains, the olms were washed up from the underground waters and were believed by local people to be a cave dragon's offspring. This cave salamander is most notable for its adaptations to a life of complete darkness in its underground habitat. The olm's eyes are undeveloped, leaving it blind, while its other senses, particularly those of smell and hearing, are acutely developed. It also lacks any pigmentation in its skin. It has three toes on its forelimbs, but only two toes on its hind feet. It also exhibits neoteny, retaining larval characteristics like external gills into adulthood, like some American amphibians, the axolotl and the mudpuppies (Necturus). The olm is the only species in the genus Proteus and the only European species of the family Proteidae, whose other extant genus is Necturus. The word olm is a German loanword that was incorporated into English in the late 19th century. The origin of the German original, Olm or Grottenolm 'cave olm', is unclear. It may be a variant of the word Molch 'salamander'. The olm's body is snakelike, 20–30 cm (8–12 in) long, with some specimens reaching up to 40 centimetres (16 in). Females grow larger than males, but otherwise the primary external difference between the sexes are in the cloaca region (shape and size) when breeding. The trunk is cylindrical, uniformly thick, and segmented with regularly spaced furrows at the myomere borders. The tail is relatively short, laterally flattened, and surrounded by a thin fin. The limbs are small and thin, with a reduced number of digits compared to other amphibians: the front legs have three digits instead of the normal four, and the rear have two digits instead of five. Its body is covered by a thin layer of skin, which contains very little of the pigment riboflavin, making it yellowish-white or pink in color. The internal organs can be seen shining through on the abdominal part of the body. The resemblance of the color of its skin to that of white people is why the olm is called human fish in some languages. However, its skin retains the ability to produce melanin, and will gradually turn dark when exposed to light; in some cases the larvae are also colored. Its pear-shaped head ends with a short, dorsoventrally flattened snout. The mouth opening is small, with tiny teeth forming a sieve to keep larger particles inside the mouth. The nostrils are so small as to be imperceptible, but are placed somewhat laterally near the end of the snout. The regressed eyes are covered by a layer of skin. The olm breathes with external gills that form two branched tufts at the back of the head. They are red in color because the oxygen-rich blood shows through the non-pigmented skin. The olm also has rudimentary lungs, but their role in respiration is only accessory, except during hypoxic conditions. Cave-dwelling animals have been prompted, among other adaptations, to develop and improve non-visual sensory systems in order to orient in and adapt to permanently dark habitats. The olm's sensory system is also adapted to life in the subterranean aquatic environment. Unable to use vision for orientation, the olm compensates with other senses, which are better developed than in amphibians living on the surface. It retains larval proportions, like a long, slender body and a large, flattened head, and is thus able to carry a larger number of sensory receptors. Although blind, the olm swims away from light. The eyes are regressed, but retain sensitivity. They lie deep below the dermis of the skin and are rarely visible except in some younger adults. Larvae have normal eyes, but development soon stops and they start regressing, finally atrophying after four months of development. The pineal body also has photoreceptive cells which, though regressed, retain visual pigment like the photoreceptive cells of the regressed eye. The pineal gland in Proteus probably possesses some control over the physiological processes. Behavioral experiments revealed that the skin itself is also sensitive to light. Photosensitivity of the integument is due to the pigment melanopsin inside specialized cells called melanophores. Preliminary immunocytochemical analyses support the existence of photosensitive pigment also in the animal's integument.