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Salamander life cycles

Continue LIFE CYCLE This is a blue spotted salamander. Its life cycle is the same as most . Click the image to follow the lifecycle. When you return to this page, you can see the lifecycle again or go elsewhere. Order of (Urodela) For the mythical lizard, Salamanders see in folklore. For other uses, see Salamander (disambiguation). SalamandersTemporal range: Late – Present Day,[1] 160-0 Ma PreꞒ Ꞓ O S D C P T J K Pg N Salamander, Ambystoma maculatum Scientific Classification Domain: Eukaryota Kingdom: Animalia Phylum: Chordata Class: Clade: Order: UrodelaDuméril, 1806 Suborders Cryptobranchoidea Salamandroidea Native distribution of salamanders (in green) Salamanders is a group of amphibians typically characterized by a lizard-like appearance , with slender bodies, blunt muzzers, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults. All modern salamander families are grouped together under the order Urodela. Salamander diversity is highest in the northern hemisphere and most species are found in the Holarctic realm, with some species found in the neotropical sphere. Salamanders rarely have more than four toes on the forelimbs and five on their hind legs, but some species have fewer numbers and others lack hind legs. Their permeable skin usually makes them dependent on habitats in or near water or other cool, moist places. Some salamander species are completely aquatic throughout life, some make their way to the water periodically, and others are completely terrestrial as adults. They can recreate lost limbs, as well as other damaged parts of their bodies. The researchers hope to reverse engineer the remarkable regenerative processes for potential human medical applications, such as brain and spinal cord injury treatment or prevent harmful scarring during cardiac surgery recovery. [2] Members of the family are best known as newts and lack the costal traces along the sides of their bodies that are typical of other groups. The skin of some species contains the powerful poison tetrodotoxin; these salamanders tend to be slow and have bright warning staining to advertise their toxicity. Salamanders usually lay eggs in water and have aquatic larvae, but great variation occurs in their life cycles. Some species in harsh environments reproduce while they are still in larval state. Description X-ray of salamander Skin lacks scales and is moist and smooth to the touch, except in newts of Salamandridae, which may have velvety or variesome skin, wet to the touch. The skin can be gray or colorful, exhibiting different patterns of stripes, bars, spots, spots, or dots. Male newts become dramatically colored during the breeding season. Cave species living in the dark lack pigmentation and have a pink or pearlescent appearance. [3] Salamanders range in size from minute salamanders, with a total length of 2.7 cm (1.1 in), including tail, to the Chinese reaching 1.8 m (5.9 feet) and weighing up to 65 kg (143 lb). However, most are between 10 and 20 cm (3.9 and 7.9 in) in length. [4] Trunk, limbs and tail An adult salamander generally resembles a small lizard, which has a basal body shape with a cylindrical trunk, four limbs, and a long tail. Except in the family Salamandridae, the head, body and tail have a number of vertical depressions in the surface that run from the middle of the dorsal region to the ventral area and are called costal grooves. Their function seems to be to help keep the skin moist by channeling water across the surface of the body. [5] Sirens have an eel- like appearance. Some aquatic species, such as sirens and , have decreased or absent hind legs, giving them an eel-like appearance, but in most species, the anterior and posterior limbs are about the same length and project sideways, barely raising the trunk from the ground. The feet are wide with short numbers, usually four on the front feet and five on the back. Salamanders have no claws, and the shape of the foot varies depending on the 's habitat. Climbing species have elongated, square toes, while mountain-dwellers have larger feet with short, blunt toes. The tree-climbing salamander (Bolitoglossa sp.) has flat-like webbed feet that follow smooth surfaces by suction, while rock climbing Hydromantes species from California have feet with fleshy scaffolding and short numbers and use their tails as an extra limb. Upon ascent, the tail props up the back of the body, while one hind foot moves forward and then swings to the other side to provide support as the other backfoot advances. [6] In larvae and aquatic salamanders, the tail is flattened sideways, its dorsal and ventral fins and undulates have side to side to push the animal through the water. In the and Salamandridae families, the male's tail, which is larger than the female's, is used under amplexus embrace to drive the mating pair to a secluded place. In terrestrial species, the tail goes to counterbalance the animal as it runs, while in arboreal salamander and other tree climbing species, it's prehensile. The tail is also used by some plethodontid salamanders that can jump, to help launch themselves into the air. [6] The tail is used in courtship and as a storage organ for proteins and lipids. It also serves as a defense against predation, when it can be lashed at the attacker or autotomised when seized. Unlike frogs, an adult salamander can regenerate limbs and its tail when these are lost. [6] Skin Rough-skinned newt Skin of salamanders, in common with other amphibians, is thin, permeable to water, services a respiratory membrane, and is well supplied with glands. It has highly cornified outer layers, renewed periodically by a skin shedding process controlled by hormones from the pituitary gland and thyroid gland. During moulting, the skin initially breaks around the mouth, and the animal moves forward through the gap to discard the skin. When the front limbs have worked clearly, a series of body ripples push the skin towards the back. The hind legs are extracted and push the skin further back, before it is eventually released by friction as the salamander moves forward with its tail pressed to the ground. [7] The animal often eats the resulting sloughed skin. [3] Glands in the skin discharge mucus that keeps the skin moist, an important factor in skin breathing and thermoregulation. The sticky layer helps protect against bacterial infections and molds, reduces friction during swimming, and makes the animal slippery and harder for predators to catch. Granular glands scattered on the upper surface, especially the head, back, and tail, produce repellent or toxic secretions. [7] Some salamander toxins are particularly potent. The rough-skinned newt (Taricha granulosa) produces neurotoxin tetrodotoxin, the most toxic nonprotein substance known. Handling newts does no harm, but ingestion of even one minute fragments of the skin is fatal. In feeding trials, fish, frogs, reptiles, birds and mammals were all found to be susceptible. [8] Mature adults of some salamander species have wedding glandular tissue in their kloacae, at the base of the tail, on the head or under the chin. Some females release chemical substances, possibly from the ventral sewage gland, to attract males, but males do not seem to use pheromones for this purpose. [9] In some plethodonts, males have conspicuous mental glands on their chins pressed against the nostrils of females during the courtship ritual. They can work to speed up the mating process, reducing the risk of it being disturbed by a predator or a rival male. [10] The gland at the tail base in Plethodon cinereus is used to mark fecal pellets to proclaim territorial ownership. [9] Senses The anterior part of olm's head carries sensitive chemo, mechano, and electroreceptors. Olfaction in salamanders plays a role in territory maintenance, recognition of predators, and courtship rituals, but is probably secondary to sight during prey selection and feeding. Salamanders have two types of sensory areas that respond to the chemistry of the environment. Olfactory epithelium in the nasal cavity picks up airborne and aquatic odors, while adjacent vomeronasal bodies detect nonvolatile chemical clues, such as flavors in the mouth. In plethodonts, the sensory epithelium of vomeronasal organs extends to the nasolabial grooves, ranging from the nostrils to the tongues of the mouth. Extended areas appear to be associated with identification of prey objects, recognition of conspecifics, and identification of individuals. [11] Most salamanders' eyes are primarily adapted to the sight of the night. In some permanent aquatic species, they are reduced in size and have a simplified retinal structure, and in cave inhabitants like Georgia blind salamander, they are absent or covered with a layer of skin. In amphibian species, the eyes are a compromise and are nearsighted in air and farsighted in water. Completely terrestrial species like the fire salamander have a flatter lens that can focus over a much wider distance. [12] To find their prey, salamanders use trichromatic color vision that extends into ultraviolet ranges, based on three photoreceptor types that are maximally sensitive around 450, 500, and 570 nm. [13] The larvae, and the adults of some very aquatic species, also have a lateral line organ, similar to that of fish, which can detect changes in water pressure. [3] All salamanders lack the middle ear cavity, eardrum and eustachian tube, but have an opercularis system like frogs, and can still detect airborne sound. [14] [15] The opercularis system consists of two ossicles: the columellan (corresponding to the stapes of higher vertebrates) fixed to the skull and the operculum. An opercularis muscle connects the latter to the chest girdle, and is kept under tension when the animal is alert. [16] The system appears to be able to detect low-frequency vibrations (500-600 Hz), which may be picked up from the ground by the forelimbs and transferred to the inner ear. These can serve to warn the animal of an approaching predator. [17] Salamanders are usually considered to have no voice and do not use audio for communication in the way frogs do; however, in mating systems they communicate through pheromone signaling; some species can make silent ticking or popping sounds, perhaps through the opening and closing of valves in the nose. The Californian giant salamander can produce a bark or rattle, and some species can squeak by pulling muscles in the throat. The arboreal salamander can squeak using another mechanism; it pulls its eyes into the head and forces air out of its mouth. Ensatina salamander sometimes makes a hissing sound, while the sirens sometimes produce silent clicks, and can resort to faint screams if attacked. Similar clicking behavior was observed in two European newts Lissotriton vulgaris and Ichthyosaura alpestris in their aqueous phase. [18] Vocalization in salamanders has been studied little and the purpose of these sounds is assumed to be startling by predators. [19] Salamanders need moist environments to respire through their skin. Respiration Respiration differs between the different species of salamanders, and may involve gills, lungs, skin, and membranes in the mouth and Larvsalamandras breathe mainly with the help of gills, which are usually outer and feather-light in appearance. Water is drawn in through the mouth and flows out through the gill slots. Some neotenic species such as mudpuppy (Necturus maculosus) retain their gills throughout their lives, but most species lose them in metamorphosis. The embryos of some terrestrial lungless salamanders, such as Ensatina, which undergo direct development, have large gills located near the egg surface. [20] When found in adult salamanders, the lungs vary greatly between species in size and structure. In aquatic, cold water species such as the southern (Rhyacotriton variegatus), the lungs are very small with smooth walls, while species living in warm water with little dissolved oxygen, such as the smaller siren (Siren intermedia), have large lungs with intricate surfaces. In the terrestrial lungless salamanders (family ), no lungs or gills are present, and gas exchange occurs mostly through the skin, supplemented with the tissues lining the mouth. To facilitate this, these salamanders have a dense network of blood vessels just below the skin and in the mouth. [20] [21] In the Amphiumas, metamorphosis is incomplete, and they retain a pair of gill slits as adults, with fully functioning internal lungs. [22] Some species lack ing lungs respire through gills. In most cases, these external gills are visible as tassels on either side of the head. Some terrestrial salamanders have lungs used in breathing, although these are simple and sac-like, unlike the more complex organs found in mammals. Many species, such as olm, have both lungs and gills as adults. [3] A dissected view of the levators arcuum muscles in a Necturus maculosus specimen. These (shown in the purple circles) move the outer gills, as a means of breathing. In Necturus, external gills begin to form as a means of combating hypoxia in the egg as egg yolks are converted into metabolically active tissue. [23] However, molecular changes in the mudpuppy during post-embryonic development primarily due to thyroid gland prevent the internalization of the external gills seen in most salamanders undergoing metamorphosis. [24] The outer gills seen in salamanders are very different from those of amphibians with internalized gills. Unlike amphibians with internalized gills that usually rely on the pressure to change within the buccala and pharyngeal cavities to ensure diffusion of oxygen on the gill curtain, neotenic salamanders such as Necturus use specified musculature, such as the levators arcuum, to move external gills to keep breathing surfaces constantly in contact with new oxygen water. [25] [26] Feeding and dieting Salamanders are opportunistic predators. They are generally not limited to specific foods, but feed on almost all organisms of reasonable size. [27] Large species such as the Japanese giant salamander (Andrias japonicus) eat crabs, fish, small mammals, amphibians, and aquatic insects. [28] In a study of smaller dusky salamanders (Desmogenthus) in the Appalachian Mountains, their diet includes earthworms, flies, beetles, beetles, leaf hoppers, springtails, moths, spiders, grasshoppers and mites. [27] Cannibalism sometimes occurs, especially when resources are short or time is limited. Tiger salamander tadpoles in ephemeral pools sometimes resort to eating each other, and are seemingly able to target unrelated individuals. [29] Adult blackberry salamanders (Desmogenthus quadramaculatus) hunt adults and young of other species of salamanders, while their larvae sometimes cannibalise smaller larvae. [30] The head of a tiger alammander Most species of salamander have small teeth in both the upper and lower jaws. Unlike frogs, even the larvae of salamanders possess these teeth. [3] Although caterpillar teeth are shaped like pointed cones, the teeth of adults are adapted to enable them to easily take prey. The crown, which has two cusps (bicuspid), is attached to a pedicel of collagen fibers. The joint formed between the bicuspid and the pedicel is partly flexible, as it can bend inwards, but not outwards. As struggling prey advances into the salamander's mouth, the tip of the teeth relaxes and bends in the same direction, encouraging movement towards the throat and resisting the prey's escape. [31] Many salamanders have spots of teeth attached to the dog and palatine bones in the mouthroof, and these help to retain prey. All types of teeth are resorbed and replaced at intervals throughout the life of the animal. [32] A terrestrial salamander captures its prey by flicking out its sticky tongue in an action that takes less than half a second. In some species, the tongue is anteriorly attached to the floor of the mouth, while in others, it is mounted on a pedicel. It is rendered sticky by secretions of mucus from the glands in its tip and on the roof of the mouth. [33] Fast cinematography shows how the tiger salamander (Ambystoma tigrinum) positions itself with its snout near its prey. Its mouth then gapes wide, the lower jaw remains stationary, and the tongue curves and changes shape as it pushes forward. The protruding tongue has a central depression, and the edge of this collapses inwardas the target is hit, catching the prey in a mucus-laden trough. Here it is kept while the animal's neck is bent, his tongue indented and its jaws closed. Large or resistant prey is retained by the teeth while repeated protrusions and withdrawals of the tongue pull it in. Swallowing involves alternative contraction and relaxation of muscles in the throat, assisted by depression of the eyeballs in the ceiling of the mouth. [34] Many lungless salamanders in the family have more elaborate feeding methods. Muscles that surround the hyoid bone contract to store elastic energy in springy connective tissue, and actually push the hyoid bone out of the mouth, causing the elongated tongue. [35] [36] Muscles originating in the pelvic region and inserting into the tongue are used to roll the tongue and hyoid back to their original positions. [37] An aquatic salamander lacks muscle in his tongue, and catches its prey in a completely different way. It grabs the food, grabs it with its teeth, and adopts a kind of inertial feeding. This involves throwing your head about, pulling water sharply in and out of your mouth, and snapping their jaws, all of which tend to tear and macerate the prey, which is then swallowed. [37] Though often feeding on slow-moving like snails, shrimps and worms, sirenids are unique among salamanders to have developed speciations toward herbivorous ones, such as beak-like jaw ends and extensive intestines. They feed on algae and other soft plants in nature, and easy eating is offered salad. [38] Defense Additional Information: Antipredator adaptations Salamanders have thin skins and soft bodies, and move quite slowly, and at first glance may seem vulnerable to opportunistic predation. However, they have several effective lines of defence. Mucus coating on damp skin makes them difficult to grip, and the slimy coating can have an offensive taste or be toxic. When attacked by a predator, a salamander can position itself to make the main venomary glands meet the aggressor. Often these are on the tail, which can be waved or turned up and arched over the animal's back. The sacrifice of the tail can be a valuable strategy, if the salamander escapes with his life and the predator learns to avoid that species of salamander in the future. [39] Aposematism Additional information: aposematism A fire salamanders striking black and yellow pattern warns of predators Skin secretions of the tiger salamander (Ambystoma tigrinum) fed to rats have been shown to produce aversion to the taste, and the rats avoided the presentation medium when it was offered them again. [40] The lighthouse salammandern (Salamandra salamandra) has a ridge of large granules down its spine able to spray a fine jet of poisonous liquid on its attacker. By angling its body appropriately, it can exactly direct the spray for a distance of up to 80 cm (31 in). [41] The Iberian bar (Pleurodeles waltz) has a different method of deterring attackers. Its skin radiates a toxic, viscous liquid and at the same time newt rotates its sharply pointed ribs through an angle between 27 and 92 °, and adopts an inflated posture. This action causes the ribs to puncture the body wall, each rib sticking out through an orange wart arranged in a lateral row. This can give an aposematic signal that makes the tags more When the danger has passed, the ribs retreat and the skin heals. [42] Camouflage and mimicking Additional information: camouflage and mimicking Although many salamanders have cryptic colors so that they are indisputable, others signal their toxicity through their vivid staining. Yellow, orange, and red are the colors that are generally used, often with black for greater contrast. Sometimes, the animal postures if attacked, revealing a flash of warning hue on its underside. The red eft, the brightly colored terrestrial juvenile form of the eastern newt (Notophthalmus viridescens), is highly toxic. It is avoided by birds and snakes, and can survive for up to 30 minutes after being swallowed (later regurgitated). [43] The red salamander (Pseudotriton ruber) is a tasty species with a similar color to the red eften. Predators that previously fed on it have been shown to avoid it after encountering red efts, an example of Batesian mimicry. [43] Other species exhibit similar mimics. In California the tasty yellow-eyed salamander (Ensatina eschscholtzii) resembles the poisonous California-newt (Taricha torosa) and the coarse-skinned newt (Taricha granulosa), whereas it in other parts of its range, cryptically colored. [44] A correlation exists between the toxicity of Californian salamander species and diurnal habits: relatively harmless species such as the Californian slender salamander (Batrachoseps attenuatus) are nocturnal and are eaten by snakes, while California's newt has many large venom glands in its skin, is diurnal, and is avoided by snakes. [45] Autotomy Additional information: autotomy Some salamander species use tail autotomy to escape predators. The tail falls off and winds around for a while after an attack, and salamander either runs away or still stops enough not to be noticed while the predator is distracted. The tail regrows over time, and salamanders routinely regenerate other complex tissues, including the lens or retina of the eye. Within just a few weeks of losing a piece of a limb, a salamander perfectly reforms the missing structure. [46] The range and habitat of Salamanders separated from the other amphibians during the mid to late , and initially resembled modern members of Cryptobranchoidea. Their similarity to lizards is the result of symplesiomorphy, their joint retention of the primitive tetrapod body plan, but they are no more closely related to lizards than they are to mammals. Their closest relatives are frogs and toads, within Batrachia. The earliest known salamander fossils have been found in geological deposits in China and Kazakhstan, dating to the middle of the Jurassic period about 164 million years ago. [47] Salamanders are found only in the Holarctic and neotropical regions, and do not reach the south of the Mediterranean Basin, himalayas, or South America in the Amazon. They extend north of about Arctic tree line, with the northernmost Asian species, Salamandrella keyserlingii occurs in the Siberian larch forests of Sakha and the northernmost species in North America, Ambystoma laterale, no further north than Labrador and Taricha granulosa do not reach beyond the Alaska Panhandle. [48] They had an exclusively Laurasian distribution until Bolitoglossa invaded South America from Central America, probably at the start of the early Miocene, some 23 million years ago. [49] They also lived on the Caribbean islands during the early Miocene era, which was confirmed by the discovery of Palaeoplethodon hispaniolae,[50] which was found trapped in amber in the Dominican Republic. But possible salamander fossils have been found in Australia at the Murgon fossil site, representing the only known salamanders known from the continent. [51] There are about 655 living species of salamander. [52] One-third of the known salamander species are found in North America. The highest concentration of these is found in the Appalachian Region, where Plethodontidae is believed to have originated in mountain streams. Here are vegetation zones and proximity to water of greater importance than height. Only species that adopted a more terrestrial way of life have been able to spread to other locations. The northern slimy salamander (Plethodon glutinosus) has a wide range and occupies a habitat similar to the southern grey-cheeked salamander (Plethodon metcalfi). The latter is limited to the slightly cooler and wetter conditions in north-facing bay forests in southern Appalachia, and to higher altitudes above 900 m (3000 feet), while the former is more adaptable, and would be fully capable of inhabiting these sites, but any unknown factor seems to prevent the two species from co-existing. [27] One species, Anderson's salamander, is one of the few species of live amphibians found in brackish water or salt water. [53] Reproduction and Development See also: Sexual selection in amphibians Sierra newt amplexus found in stream at Woolman Vacations in Nevada County, California Salamanders are not vocal and in most species the sexes look the same, so they use smell and tactile cues to identify potential mates, and sexual selection occurs. Pheromones play an important part in the process and can be produced by the abdominal gland in males and by the valve glands and skin of both sexes. Men are sometimes seen examining potential mates with their snouts. In Old World newts, Triturus spp., the males are sexually dimorphic and appear in front of the females. Visual cues are also believed to be important in some Plethodont species. [54] In about 90% of all species, fertilisation is internal. The male usually deposits a spermatophobe on the ground or in the water according to species, and the female picks this up with its valve. It is Has a package of sperm supported on a conical gelatina base, and often an elaborate courtship behavior is involved in its deposition and collection. Once inside the sewer, spermatozoa moves to spermatheca, one or more chambers in the ceiling of kloaca, where they are stored for sometimes long periods until the eggs are laid. In the most primitive salamanders, like the Asian salamanders and the giant salamanders, external fertilization takes place instead. In these species, the male releases sperm on the egg mass in a reproductive process similar to typical frogs. [54] Three different types of egg deposition occur. Ambystoma and Taricha spp. Spawn large numbers of small eggs in calm ponds where many large predators are unlikely. Most obscure salamanders (Dismature thus) and Pacific giant salamanders (Dicamptodon) lay smaller batches of medium-sized eggs in a hidden place in flowing water, and these are usually guarded by an adult, normally female. Many of the tropical climbing salamander (Bolitoglossa) and lungless salamanders (Plethodontinae) lay a small number of large eggs on land in a well-hidden place, where they are also guarded by the mother. [54] Some species such as the Fire Salamander (Salamandra) are ovoviviparous, where the female retains the eggs inside the body until they hatch, either to larvae to be deposited in a water body, or in fully formed juveniles. [3] Spawning embryonic development of a salamander, filmed in the 1920s In temperate regions, reproduction is usually seasonal and salamanders can migrate to breeding grounds. Males arrive usually first and in some instanceset up territories. Typically, a larval stage follows where the organism is completely aquatic. Tadpoles have three pairs of outer gills, no eyelids, a long body, a lateral flattened tail with dorsal and ventral fins and in some species limb-buds or limbs. Dust-type larvae can have a pair of rod-like balancing apparatus on either side of the head, long gill filaments and wide fins. Stream-type larvae are narrower with short gill filaments, narrower fins and no balancers, but instead the hind legs have already evolved when they hatch. [55] Tadpoles are carnivorous and the larval stage can last from day to year, depending on the species. Sometimes this stage is completely past, and the eggs of most lungless salamanders (Plethodontidae) develop directly into miniature versions of the adult without an intermediate larval stage. [56] At the end of the larval stage, the tadpoles already have limbs and metamorphosis normally takes place. In salamanders, this occurs over a short period of time and involves the closure of gill slits and the loss of gill and tail fin structures that are not required as adults. At the same time, eyelids develop, the mouth becomes wider, a tongue appears, and teeth are formed. The aqueous appear on land as a terrestrial adult. [57] Neotenic axolotl, which shows external gills Not all species of salamanders follow this path. Neoteny, also known as paedomorphosis, has been observed in all salamander families, and may be generally possible in all salamander species. In this condition, an individual can retain gills or other childish characteristics while achieving reproductive maturity. The changes that occur at metamorphosis are under the control of thyroid hormones and in obligate neotenes such as axolotl (Ambystoma mexicanum), the tissues are seemingly unresponsive to the hormones. In other species, the changes cannot be triggered due to the underactivity of the hypothalamus-pituitary-thyroid mechanism that can occur when conditions in the terrestrial environment are too inhospitable. [57] This may be due to cold or wildly fluctuating temperatures, aridity, lack of food, lack of coverage, or insufficient iodine for the formation of thyroid hormones. Genetics can also play a role. The larvae of tiger salamanders (Ambystoma tigrinum), for example, develop limbs shortly after hatching and in seasonal pools immediately undergo metamorphosis. Other larvae, especially in permanent pools and warmer climates, may not undergo metamorphosis until fully adult in size. Other populations in colder climates may not be metamorphosis at all, and become sexually mature while in their larval forms. Neoteny allows the species to survive even when the terrestrial environment is too harsh for the adults to thrive on land. [55] Conservation The endangered hell-bone rara A general decline in live amphibian species has been linked to the fungal disease chytridiomycosis. A higher proportion of salamander species than frogs or caecilian sits in one of the risk categories established by the IUCN. Salamanders showed a significant decrease in numbers in recent decades of the 20th century, although no direct link between the fungus and population decline has yet to be found. [58] The IUCN made further efforts in 2005 when they established the Amphibian Conservation Action Plan (ACAP), which was subsequently followed by the Amphibian Ark (AArk), the Amphibian Specialist Group (ASG), and finally the umbrella organisation known as ACAPC. [59] The researchers also cite deforestation, resulting in fragmentation of suitable habitats, and climate change as possible contributing factors. Species such as Pseudoeurycea brunnata and Pseudoeurycea goebeli that had been abundant in the cloud forests of Guatemala and Mexico in the 1970s were found to be rare in 2009. [60] However, few data have been collected on population sizes over the years, and through intensive mapping of historical and appropriate new sites, it has been possible to locate individuals of other species such as Parvimolge townsendi, which had been thought to be extinct. [58] At present, lines of defence for the conservation of Salamanders include both in situ and ex situ conservation methods. There are efforts for some members of the Salamander family to be preserved under a conservation breeding program (CBP) but it is important to note that there should be research in advance to determine whether salamander species will actually value from CBP, which scientists have noted that some species of amphibians completely fail in this environment. [59] Various conservation initiatives are being deprised around the world. The Chinese giant salamander, at 1.8 m (6 feet) the largest amphibian in the world, is critically endangered, as it is collected for food and for use in traditional Chinese medicine. An environmental training programme is being implemented to encourage the sustainable management of wild populations in the Qinling Mountains and captive breeding programmes have been established. [61] The Hellbender is another large, long-lived species with dwindling numbers and fewer juveniles reaching maturity than before. [62] Another alarming finding is the increase in anomalies in up to 90% of the Hellbender population in the Spring River watershed in Arkansas. [63] Habitat loss, waterdegradation, pollution and disease have all been involved in the decline, and a captive breeding program at the Saint Louis Zoo has been successfully established. [64] Of the 20 species of minute salamanders (Thorius spp.) in Mexico, half are believed to have been extinct and most of the others are critically endangered. Specific causes of the decline may include climate change, chytridiomycosis, or volcanic activity, but the biggest threat is habitat destruction such as logging, agricultural activities and human settlement reducing their often small, fragmented ranges. Research work is underway to assess the status of these salamanders, and to better understand the factors involved in their population declines, with a view to taking action. Ambystoma mexicanum is a species protected under the Mexican Unit for Management and Conservation of Wildlife (UMA) from April 1994. Another harmful factor is that axolotl has lost its role as a top predator since the introduction of locally exotic species such as Nile tilapia and carp. Tilapia and carp compete directly with axolotls by consuming their eggs, larvae and juveniles. Climate change has also greatly affected axolotls and their populations throughout the southern Mexico area. Because of its proximity to Mexico City, officials are currently working on programs at Lake Xochimilco to bring in tourism and educate the local population about the restoration of the natural habitat of these creatures. [66] This proximity is a major factor that has affected the survival of the axolotl, as the city has expanded to take over the Xochimilco region to make resources for water and disposal and sewerage. [67] However, Axolotlen has the advantage of being housed on farms for research premises. So there is still a chance that they might be able to return to their natural habitat. The recent population decline has significantly affected the genetic diversity of populations of axolotl, making it difficult to further progress scientifically. It is important to note that although there is a level of limited genetic diversity due to Ambystoma populations, such as axolotl, paedeomorphic species, it does not take into account the overall lack of diversity. There is evidence to point to a historical bottleneck of Ambystoma that contributes to the variation issues. Unfortunately, there is no large genetic pool for the species to draw from unlike in historical times. Thus, there are serious concerns about inbreeding due to lack of gene flow. [68] One way that scientists are investigating to maintain genetic diversity within the population is via cryopreservation of the spermatophores from the male axolotl. It is a safe and non-invasive method that requires the collection of the spermatophores and places them in a deep freeze for preservation. Most importantly, they have found that in only limited damage to the spermatophores when dewed and thus it is a viable option. As of 2013, it is a method used to save not only axolotl but also many other members of the salamander family. [67] [69] [70] Research is being done on the environmental signals that need to be replicated before captive animals can be persuaded to breed. Common species such as tiger salamander and mudpuppy are given hormones to stimulate the production of sperm and eggs, and arginine vasotocin's role in courtship behavior is investigated. Another line of research is artificial insemination, either in vitro or by inserting spermatophores into females clonacae. The results of this research can be used in captive breeding programmes for endangered species. [71] Taxonomy Contradiction exists among different authorities as to the definition of terms Caudata and Urodela. Some argue that Urodela should be limited to the crown group, with Caudata used for the overall group. Others limit the name Caudata to the crown group and use Urodela for the total group. [72] [73] The previous procedure appears to be most generally adopted and is used in this article. [52] The 10 families belonging to Urodela are divided into three sub-orders. [72] Kladen Neocaudata is often used to separate the Cryptobranchoidea and the Salamandroidea from the Sirenoidea. Cryptobranchoidea (Giant salamanders) Family Common names Examples species Example image Cryptobranchidae Giant salamanders Hellbender (Cryptobranchus alleganiensis) Hynobiidae Asiasalamanders Hida salamander (Hynobius Salamandroidea (Advanced Salamanders) Ambystomatidae Mole Salamanders Marbled Salamanders (Ambystoma opacum) Amphiumidae Amphiumas or Congo eels Two-to- (Amphiuma agent) Plethodontidae Lungless salamanders Red-backed salamander (Plethodon cinereus) Mudpuppies and Olms Olm (Proteus anguinus) Rhyacotritonidae Torrent salamanders Southern torrent salamander (Rhyacotriton variegatus) Salamandridae Newts and true salamanders Alpine newt (Ichthyosaura alpestris) Sirenoidea (Sirens) Sirens Major Siren (Siren lacertina) Phylogeny and evolution Origin and evolutionary relationships between the three main groups of amphibians (gymnophionans urodeles and anurans) is a matter of debate. A 2005 molecular phylogeny, based on rDNA analysis, suggested that the first divergence between these three groups took place shortly after they had branched out from lobe-finned fish in (about 360 million years ago), and before the dissolution of the supercontinent Pangaea. The short period, and the rate at which radiation took place, may help account for the relative lack of amphibian fossils that appear to be closely related to lissamphibians. [74] However, recent studies have generally found newer (Late [75] to Permian[76]) age for the basalmost divergence among lissamphibians. The first known fossil salamanders are Kokartus honorarius from the middle Jura in Kyrgyzstan and two species of the seemingly neotenan, aquatic Marmorerpeton from England[77] of a similar date. [78] They looked superficially like robust modern salamanders but lacked a number of anatomical properties that were later developed. Karaurus sharovi from the upper Jurassic of Kazakhstan resembled modern mole salamanders in morphology and probably had a similar digging lifestyle. [52] The two main groups of extant salamanders are Cryptobranchoidea (primitive salamanders) and Salamandroidea (advanced salamanders), also known as Diadectosalamandroidei, both seem to have appeared before the end of the Jura, the former as exemplified by chunerpeton tianyiensis, Pangerpeton sinensis, Jeholotriton paradoxus, Regalerpeton weichangensis, Liaoxitriton daohugouensis and Iridotriton hechti, and the latter by Beiyanerpeton jianpingensis. By the Upper , most or all of the living salamander families had probably appeared. [52] The following kladogram shows the relationships between salamander families based on the molecular analysis of the Pyron and Wiens (2011). [79] Sirenidae's position is disputed, but the position of sister of Salamandroidea fits best with the molecular and fossil evidence. [52] Cryptobranchoidea Hynobiidae Cryptobranchidae Sirenoidea Sirenidae Salamandroidea Treptobranchia Ambystomatidae Dicamptodontidae Proteidae Plethosalamandroidei Rhyacotritonidae Xenosalamandroidei Amphiumidae Plethodontidae Genome and genetics salamanders possess gigantic genomes, spanning the range from 14 Gb to 120 Gb[80] (the human genome is 3.2 Gb long). Pleurodele's genome (20 Gb) and Ambystoma mexicanum (32 Gb) have been sequenced. [81] [82] In the human community Myth and legend Main articles: Salamanders in folklore and legend and Legendary salamander in popular cultureA A salamander unscathed in the fire, 1350 Legends have evolved around the salamander over the centuries, many related to fire. This connection probably originates from the tendency of many salamanders to live inside rotting logs. When the log was placed in a fire, salamander would try to escape, lending credence to the belief that salamanders were created from flames. [83] The association of salamander with fire first appeared in ancient Rome, with Pliny the Elder writing in his Natural History that A Salamander is so cold that it extinguishes fire upon contact. It vomits from its mouth a milky liquid; if this fluid touches any part of the human body it causes all hair to fall off, and the skin to change color and break out into a rash. [84] The ability to extinguish fire is repeated by Saint Augustine in the fifth century and Isidore of Seville in the seventh century. [85] Ukiyo-e-print by Utagawa Kuniyoshi (1797–1861) depicting a giant salamander stabbed by the samurai Hanagami Danjō no jō Arakage The mythical ruler Prester John is to have a robe made of salamander hair; The Emperor of India possessed a suit made of a thousand skins; Pope Alexander III had a tunic that he valued highly and William Caxton (1481) wrote: This Salemandre berithe wulle, of which is made fabric and gyrdles that do not get brenne in fyre. [87] The salamander was said to be so poisonous that by twisting around a tree, the poisoned fruit could kill everyone who ate them and by falling into a well could kill everyone who drank from it. [87] The Japanese giant salamander has been the subject of legend and artwork in Japan, in the ukiyo-e work of Utagawa Kuniyoshi. The well-known Japanese mythological creature known as the coat can be inspired by this salamander. [88] Limb regeneration applied to human Salamandrar limb regeneration has long been the focus of interest among scientists. Researchers have been trying to find out the conditions required for the growth of new limbs and hope that such a renewal could be replicated in humans using stem cells. Axolotls have been used in research and have been genetically manipulated so that a fluorescent protein is present in cells in the bone, allowing the cell division process to be traced under the microscope. It seems that after the loss of a limb, cells pull together to form lump called a blastema. This seems superficially undifferentiated, but cells originating in the skin later develop into new skin, muscle cells to new muscles and cartilage cells to new cartilage. It is only the cells from just below the surface of the skin that are pluripotent and able to develop into any type of cell. [89] Researchers from the Australian Regenerative Medicine Institute have found that when macrophages were removed, salamanders lost their ability to regenerate and instead formed scar tissue. If the processes involved in forming new tissue can be reverse engineered to humans, it may be possible to heal damage to the spinal cord or brain, repair damaged organs and reduce scarring and fibrosis after surgery. [90] Salamander brandy A 1995 article in the Slovenian weekly newspaper Mladina published Salamander brandy, a liquor supposedly native to Slovenia. 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Stebbins, Robert C.; Cohen, Nathan W. (1995). A natural history of amphibians. Princeton University Press. ISBN 978-0-691-03281-8. External links Wikispecies has information related to Urodela Wikimedia Commons has media related to Urodela. Tree of Life: Caudata Salamander Gallery Caudata Culture Critter Crossings: Salamander Tunnels at the U.S. Department of Transportation ArchéoZooThèque : Urodele Skeleton Drawing : available in vector, image and PDF format Taken from

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