Snake

I can absolutely teach you a lot about salamanders — but instead of dumping a giant wall of
text (which usually doesn’t stick), let’s build it together so you actually learn it well.

Here’s a quick, clear starting point. After each small chunk, I’ll ask you one question so you
stay in control of what we go deeper into.

🦎 Quick Starter: What a Salamander Is

Salamanders are amphibians (like frogs), and most have:

●​ Moist, smooth skin​

●​ A long tail (unlike frogs)​

●​ A need for wet or damp environments​

●​ A life cycle that usually starts in water​

Question:​
Do you want to start by exploring (A) their body + adaptations, (B) life cycle, (C) habitat +
ecology, or (D) types of salamanders?​
Pick one, and we’ll go deep step-by-step.

Salamanders are a group of amphibians typically characterized by their lizard-like

appearance, with slender bodies, blunt snouts, short limbs projecting at right angles
to the body, and the presence of a tail in both larvae and adults. All ten extant
salamander families are grouped together under the order Urodela, the sole
[2]
surviving order from the group Caudata. Urodela is a scientific Latin term based on
the Ancient Greek οὐρά δήλη: ourà dēlē "conspicuous tail". Caudata is the Latin for
"tailed ones", from cauda: "tail".

Salamander diversity is highest in eastern North America, especially in the

Appalachian Mountains; most species are found in the Holarctic realm, with some
species present in the Neotropical realm. Salamanders never have more than four
[3]
toes on their front legs and five on their rear legs, but some species have fewer
digits and others lack hind limbs. Their permeable skin usually makes them reliant on
habitats in or near water or other cool, damp places. Some salamander species are
fully aquatic throughout their lives, some take to the water intermittently, and others
are entirely terrestrial as adults.

This group of amphibians is capable of regenerating lost limbs as well as other

damaged parts of their bodies. Researchers hope to reverse engineer the
regenerative processes for potential human medical applications, such as brain and
spinal cord injury treatment or preventing harmful scarring during heart surgery
[4]
recovery. The remarkable ability of salamanders to regenerate is not just limited to
limbs but extends to vital organs such as the heart, jaw, and parts of the spinal cord,
showing their uniqueness compared to different types of vertebrates. ⁤This ability is
most remarkable for occurring without any type of scarring. ⁤This has made
salamanders an invaluable model organism in scientific research aimed at
understanding and achieving regenerative processes for medical advancements in
[5]
human and animal biology.

Members of the family Salamandridae are mostly known as newts and lack the
costal grooves along the sides of their bodies typical of other groups. The skin of
some species contains the powerful poison tetrodotoxin; these salamanders tend to
be slow-moving and have bright warning coloration to advertise their toxicity.
Salamanders typically lay eggs in water and have aquatic larvae, but great variation
occurs in their lifecycles. Some species in harsh environments reproduce while still
in the larval state.

Etymology[edit]
The word salamander comes from Old French salamandre from Latin salamandra
[6]
from Greek σαλαμάνδρα : salamándra, of uncertain, possibly, pre-Greek origin.
[7]
The Greek word is used for the fire salamander.

Description[edit]

X-ray image of salamander

The skin lacks scales and is moist and smooth to the touch, except in newts of the
Salamandridae, which may have velvety or warty skin, wet to the touch. The skin
may be drab or brightly colored, exhibiting various patterns of stripes, bars, spots,
blotches, or dots. Male newts become dramatically colored during the breeding
season. Cave species dwelling in darkness lack pigmentation and have a translucent
[8]
pink or pearlescent appearance.

Salamanders range in size from the minute salamanders, with a total length of 27
mm (1

1
⁄8 in), including the tail, to the Chinese giant salamander which reaches 1.8 m (6 ft)
and weighs up to 65 kg (145 lb). All the largest species are found in the four families
giant salamanders, sirens, Congo eels and Proteidae, which are all aquatic and
[9][10]
obligate paedomorphs. Some of the largest terrestrial salamanders, which go
through full metamorphosis, belong to the family of Pacific giant salamanders, and
[11]
are much smaller. Most salamanders are between 10 and 20 cm (4 and 8 in) in
[12]
length.

Trunk, limbs and tail[edit]

An adult salamander generally resembles a small lizard, having a basal tetrapod

body form 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 which run from the mid-dorsal region to the ventral area and are known as
costal grooves. Their function seems to be to help keep the skin moist by channeling
[13]
water over the surface of the body.

Sirens have an eel-like appearance.

Some aquatic species, such as sirens and amphiumas, have reduced or absent hind
limbs, giving them an eel-like appearance, but in most species, the front and rear
limbs are about the same length and project sideward, barely raising the trunk off the
ground. The feet are broad with short digits, usually four on the front feet and five on
the rear. Salamanders do not have claws, and the shape of the foot varies according
to the animal's habitat. Climbing species have elongated, square-tipped toes, while
rock-dwellers have larger feet with short, blunt toes. The tree-climbing salamander
(Bolitoglossa sp.) has plate-like webbed feet which adhere to smooth surfaces by
suction, while the rock-climbing Hydromantes species from California have feet with
fleshy webs and short digits and use their tails as an extra limb. When ascending,
the tail props up the rear of the body, while one hind foot moves forward and then
[14]
swings to the other side to provide support as the other hind foot advances.

In larvae and aquatic salamanders, the tail is laterally flattened, has dorsal and
ventral fins, and undulates from side to side to propel the animal through the water.
In the families Ambystomatidae and Salamandridae, the male's tail, which is larger
than that of the female, is used during the amplexus embrace to propel the mating
couple to a secluded location. In terrestrial species, the tail moves to counterbalance
the animal as it runs, while in the arboreal salamander and other tree-climbing
species, it is prehensile. The tail is also used by certain plethodontid salamanders
[14]
that can jump, to help launch themselves into the air. The tail is used in courtship
and as a storage organ for proteins and lipids. It also functions as a defense against
predation, when it may be lashed at the attacker or autotomised when grabbed.
Unlike frogs, an adult salamander is able to regenerate limbs and its tail when these
[14]
are lost.

Skin[edit]

Rough-skinned newt

The skin of salamanders, in common with other amphibians, is thin, permeable to

water, serves as a respiratory membrane, and is well-supplied with glands. It has
highly cornified outer layers, renewed periodically through a skin shedding process
controlled by hormones from the pituitary and thyroid glands. During moulting, the
skin initially breaks around the mouth, and the animal moves forward through the
gap to shed the skin. When the front limbs have been worked clear, a series of body
ripples pushes the skin toward the rear. The hind limbs are extracted and push the
skin farther back, before it is eventually freed by friction as the salamander moves
[15]
forward with the tail pressed against the ground. The animal often then eats the
[8]
resulting sloughed skin.

Glands in the skin discharge mucus which keeps the skin moist, an important factor
in skin respiration and thermoregulation. The sticky layer helps protect against
bacterial infections and molds, reduces friction when swimming, and makes the
animal slippery and more difficult for predators to catch. Granular glands scattered
on the upper surface, particularly the head, back, and tail, produce repellent or toxic
[15]
secretions. Some salamander toxins are particularly potent. The rough-skinned
newt (Taricha granulosa) produces the neurotoxin tetrodotoxin, the most toxic
nonprotein substance known. Handling the newts does no harm, but ingestion of
even a minute fragment of skin is deadly. In feeding trials, fish, frogs, reptiles, birds,
[16]
and mammals were all found to be susceptible.

Mature adults of some salamander species have "nuptial" glandular tissue in their
cloacae, at the base of their tails, on their heads or under their chins. Some females
release chemical substances, possibly from the ventral cloacal gland, to attract
[17]
males, but males do not seem to use pheromones for this purpose. In some
plethodonts, males have conspicuous mental glands on the chin which are pressed
against the females' nostrils during the courtship ritual. They may function to speed
up the mating process, reducing the risk of its being disrupted by a predator or rival
[18]
male. The gland at the base of the tail in Plethodon cinereus is used to mark fecal
[17]
pellets to proclaim territorial ownership.

Senses[edit]

The front part of the olm's head carries sensitive chemo-, mechano-, and electroreceptors.
 Biofluorescence can be observed across various salamander species

Smell[edit]

Olfaction in salamanders plays a role in territory maintenance, the 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 adjoining vomeronasal organs detect nonvolatile
chemical cues, such as tastes in the mouth. In plethodonts, the sensory epithelium of
the vomeronasal organs extends to the nasolabial grooves, which stretch from the
nostrils to the corners of the mouth. These extended areas seem to be associated
with the identification of prey items, the recognition of conspecifics, and the
[19]
identification of individuals.

Vision[edit]

The eyes of most salamanders are adapted primarily for vision at night. In some
permanently aquatic species, they are reduced in size and have a simplified retinal
structure, and in cave dwellers such as the Georgia blind salamander, they are
absent or covered with a layer of skin. In amphibious species, the eyes are a
compromise and are nearsighted in air and farsighted in water. Fully terrestrial
species such as the fire salamander have a flatter lens which can focus over a much
[20]
wider range of distances. To find their prey, salamanders use trichromatic color
vision extending into the ultraviolet range, based on three photoreceptor types that
[21]
are maximally sensitive around 450, 500, and 570 nm. The larvae, and the adults
of some highly aquatic species, also have a lateral line organ, similar to that of fish,
[8]
which can detect changes in water pressure.

Hearing[edit]

All salamanders lack middle ear cavity, eardrum and eustachian tube, but have an
[22][23]
opercularis system like frogs, and are still able to detect airborne sound. The
opercularis system consists of two ossicles: the columella (equivalent to the stapes
of higher vertebrates) which is fused to the skull, and the operculum. An opercularis
muscle connects the latter to the pectoral girdle, and is kept under tension when the
[24]
animal is alert. The system seems able to detect low-frequency vibrations
(500–600 Hz), which may be picked up from the ground by the fore limbs and
transmitted to the inner ear. These may serve to warn the animal of an approaching
[25]
predator.

Vocalization[edit]

Salamanders are usually considered to have no voice and do not use sound for
communication in the way that frogs do. Before mating, they communicate by
pheromone signaling; some species make quiet ticking, clicking, squeaks or popping
[26]
noises, perhaps by the opening and closing of valves in the nose. Most
salamanders lack vocal cords, but a larynx is present in the mudpuppy (Necturus)
and some other species, and the Pacific giant salamanders and a few others have a
[27]
large larynx and bands known as plicae vocales. The California giant salamander
can produce a bark or rattle, and a few species can squeak by contracting muscles
in the throat. The arboreal salamander can squeak using a different mechanism; it
retracts its eyes into its head, forcing air out of its mouth. The ensatina salamander
occasionally makes a hissing sound, while the sirens sometimes produce quiet
clicks, and can resort to faint shrieks if attacked. Similar clicking behaviour was
observed in two European newts Lissotriton vulgaris and Ichthyosaura alpestris in
[26]
their aquatic phase. Vocalization in salamanders has been little studied and the
[28]
purpose of these sounds is presumed to be the startling of predators.
 Salamanders need moist environments to respire through their skin.

Respiration[edit]

Respiration differs among the different species of salamanders, and can involve gills,
lungs, skin, and the membranes of mouth and throat. Larval salamanders breathe
primarily by means of gills, which are usually external and feathery in appearance.
Water is drawn in through the mouth and flows out through the gill slits. Some
neotenic species such as the mudpuppy (Necturus maculosus) retain their gills
throughout their lives, but most species lose them at metamorphosis. The embryos
of some terrestrial lungless salamanders, such as Ensatina, that undergo direct
[29]
development, have large gills that lie close to the egg's surface.

When present in adult salamanders, lungs vary greatly among different species in
size and structure. In aquatic, cold-water species like the torrent salamanders
(Rhyacotriton), the lungs are very small with smooth walls, while species living in
warm water with little dissolved oxygen, such as the lesser siren (Siren intermedia),
have large lungs with convoluted surfaces. In the lungless salamanders (family
Plethodontidae and the clawed salamanders in the family of Asiatic salamanders),
no lungs or gills are present, and gas exchange mostly takes place through the skin,
known as cutaneous respiration, supplemented by the tissues lining the mouth. To
facilitate this, these salamanders have a dense network of blood vessels just under
[29][30][31]
the skin and in the mouth.

In the amphiumas, metamorphosis is incomplete, and they retain one pair of gill slits
[32]
as adults, with fully functioning internal lungs. Some species that lack lungs
respire through gills. In most cases, these are external gills, visible as tufts on either
side of the head. Some terrestrial salamanders have lungs used in respiration,
although these are simple and sac-like, unlike the more complex organs found in
[8]
mammals. Many species, such as the olm, have both lungs and gills as adults.
 A dissected view of the levatores arcuum muscles in a Necturus maculosus specimen. These
(shown in the purple circles) move the external gills, as a means of respiration.

In the Necturus, external gills begin to form as a means of combating hypoxia in the
[33]
egg as egg yolk is converted into metabolically active tissue. Molecular changes
in the mudpuppy during post-embryonic development primarily due to the thyroid
gland prevent the internalization of the external gills as seen in most salamanders
[34]
that undergo metamorphosis. The external gills seen in salamanders differs
greatly from that of amphibians with internalized gills. Unlike amphibians with
internalized gills which typically rely on the changing of pressures within the buccal
and pharyngeal cavities to ensure diffusion of oxygen onto the gill curtain, neotenic
salamanders such as Necturus use specified musculature, such as the levatores
arcuum, to move external gills to keep the respiratory surfaces constantly in contact
[35][36]
with new oxygenated water.

Feeding and diet[edit]

Salamanders are opportunistic predators. They are generally not restricted to
[37]
specific foods, but feed on almost any organism of a reasonable size. Large
species such as the Japanese giant salamander (Andrias japonicus) eat crabs, fish,
[38]
small mammals, amphibians, and aquatic insects. In a study of smaller dusky
salamanders (Desmognathus) in the Appalachian Mountains, their diet includes
earthworms, flies, beetles, beetle larvae, leafhoppers, springtails, moths, spiders,
[37]
grasshoppers, and mites. Cannibalism sometimes takes place, 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
[39]
individuals. Adult blackbelly salamanders (Desmognathus quadramaculatus) prey
on adults and young of other species of salamanders, while their larvae sometimes
[40]
cannibalise smaller larvae.

The head of a tiger salamander

Most species of salamander have small teeth in both their upper and lower jaws.
[8]
Unlike frogs, even the larvae of salamanders possess these teeth. Although larval
teeth are shaped like pointed cones, the teeth of adults are adapted to enable them
to readily grasp prey. The crown, which has two cusps (bicuspid), is attached to a
pedicel by collagenous fibers. The joint formed between the bicuspid and the pedicel
is partially flexible, as it can bend inward, but not outward. When struggling prey is
advanced into the salamander's mouth, the teeth tips relax and bend in the same
direction, encouraging movement toward the throat, and resisting the prey's
[41]
escape. Many salamanders have patches of teeth attached to the vomer and the
palatine bones in the roof of the mouth, and these help to retain prey. All types of
[42]
teeth are resorbed and replaced at intervals throughout the animal's life.

A terrestrial salamander catches its prey by flicking out its sticky tongue in an action
that takes less than half a second. In some species, the tongue is attached anteriorly
to the floor of the mouth, while in others, it is mounted on a pedicel. It is rendered
[43]
sticky by secretions of mucus from glands in its tip and on the roof of the mouth.
High-speed cinematography shows how the tiger salamander (Ambystoma tigrinum)
positions itself with its snout close to its prey. Its mouth then gapes widely, the lower
jaw remains stationary, and the tongue bulges and changes shape as it shoots
forward. The protruded tongue has a central depression, and the rim of this
collapses inward as the target is struck, trapping the prey in a mucus-laden trough.
Here it is held while the animal's neck is flexed, the tongue retracted and jaws
closed. Large or resistant prey is retained by the teeth while repeated protrusions
and retractions of the tongue draw it in. Swallowing involves alternate contraction
and relaxation of muscles in the throat, assisted by depression of the eyeballs into
[44]
the roof of the mouth. Many lungless salamanders of the family Plethodontidae
have more elaborate feeding methods. Muscles surrounding the hyoid bone contract
to store elastic energy in springy connective tissue, and actually "shoot" the hyoid
[45][46]
bone out of the mouth, thus elongating the tongue. Muscles that originate in
the pelvic region and insert in the tongue are used to reel the tongue and the hyoid
[47]
back to their original positions.

An aquatic salamander lacks muscles in the tongue, and captures its prey in an
entirely different manner. It grabs the food item, grasps it with its teeth, and adopts a
kind of inertial feeding. This involves tossing its head about, drawing water sharply in
and out of its mouth, and snapping its jaws, all of which tend to tear and macerate
[47]
the prey, which is then swallowed.

Though frequently feeding on slow-moving animals like snails, shrimps and worms,
sirenids are unique among salamanders for having developed herbivory speciations,
such as beak-like jaw ends and extensive intestines. They feed on algae and other
[48]
soft-plants in the wild, and easily eat offered lettuce.

Defense[edit]
Further information: Antipredator adaptations
Salamanders have thin skins and soft bodies, move rather slowly and might appear
vulnerable to opportunistic predation, but have several effective lines of defense.
Mucus coating on damp skin makes them difficult to grasp, and the slimy coating
may have an offensive taste or be toxic. When attacked by a predator, a salamander
may position itself to make the main poison glands face the aggressor. Often, these
are on the tail, which may be waggled or turned up and arched over the animal's
back. The sacrifice of the tail may be a worthwhile strategy, if the salamander
escapes with its life and the predator learns to avoid that species of salamander in
[49]
the future.

Aposematism[edit]
Further information: Aposematism

A fire salamander's striking black and yellow pattern warns off predators
Skin secretions of the tiger salamander (Ambystoma tigrinum) fed to rats have been
shown to produce aversion to the flavor, and the rats avoided the presentational
[50]
medium when it was offered to them again. The fire salamander (Salamandra
salamandra) has a ridge of large granular glands down its spine which are able to
squirt a fine jet of toxic fluid at its attacker. By angling its body appropriately, it can
[51]
accurately direct the spray for a distance of up to 80 cm (30 in).

The Iberian ribbed newt (Pleurodeles waltl) has another method of deterring
aggressors. Its skin exudes a poisonous, viscous fluid and at the same time, the
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 protruding through an orange wart arranged in a lateral row. This may
provide an aposematic signal that makes the spines more visible. When the danger
[52]
has passed, the ribs retract and the skin heals.

Camouflage and mimicry[edit]

Further information: Camouflage and mimicry
Although many salamanders have cryptic colors so as to be unnoticeable, others
signal their toxicity by their vivid coloring. Yellow, orange, and red are the colors
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 poisonous. It is avoided by birds and snakes, and can survive for up to 30
[53]
minutes after being swallowed (later being regurgitated). The red salamander
(Pseudotriton ruber) is a palatable species with a similar coloring to the red eft.
Predators that previously fed on it have been shown to avoid it after encountering
[53]
red efts, an example of Batesian mimicry. Other species exhibit similar mimicry. In
California, the palatable yellow-eyed salamander (Ensatina eschscholtzii) closely
resembles the toxic California newt (Taricha torosa) and the rough-skinned newt
[54]
(Taricha granulosa), whereas in other parts of its range, it is cryptically colored. A
correlation exists between the toxicity of Californian salamander species and diurnal
habits: relatively harmless species like the California slender salamander
(Batrachoseps attenuatus) are nocturnal and are eaten by snakes, while the
California newt has many large poison glands in its skin, is diurnal, and is avoided by
[55]
snakes.

Autotomy[edit]
Further information: Autotomy
Some salamander species use tail autotomy to escape predators. The tail drops off
and wriggles around for a while after an attack, and the salamander either runs away
or stays still enough not to be noticed while the predator is distracted. The tail
regrows with time, and salamanders routinely regenerate other complex tissues,
including the lens or retina of the eye. Within only a few weeks of losing a piece of a
[56]
limb, a salamander perfectly reforms the missing structure.

Distribution and habitat[edit]

Salamanders split off from the other amphibians during the mid- to late Permian, and
initially were similar to modern members of the Cryptobranchoidea. Their
resemblance to lizards is the result of symplesiomorphy, their common retention of
the primitive tetrapod body plan, but they are no more closely related to lizards than
they are to mammals. Their nearest relatives are the frogs and toads, within
Batrachia.

The oldest known total-group (Caudata) salamander is Triassurus from the Triassic
[57]
of Kyrgyzstan. Further salamander fossils are known from the Middle Jurassic of
[58] [59] [60] [61]
England, Scotland, China, and Kazakhstan. The oldest known
crown-group salamander (Urodela) remains uncertain but recent analyses suggest it
[59]
is Valdotriton from the Late Jurassic of Spain.

Salamanders are found only in the Holarctic and Neotropical regions, not reaching
south of the Mediterranean Basin, the Himalayas, or in South America the Amazon
Basin. They do not extend north of the Arctic tree line, with the northernmost Asian
species, Salamandrella keyserlingii, which can survive long-term freezing at −55
[62]
°C, occurring in the Siberian larch forests of Sakha and the most northerly
species in North America, Ambystoma laterale, reaching no farther north than
[63]
Labrador and Taricha granulosa not beyond the Alaska Panhandle. They had an
exclusively Laurasian distribution until Bolitoglossa invaded South America from
Central America, probably by the start of the Early Miocene, about 23 million years
[64]
ago. They also lived on the Caribbean Islands during the early Miocene epoch,
[65]
confirmed by the discovery of Palaeoplethodon hispaniolae, found trapped in
amber in the Dominican Republic. Vertebrae fossils recovered from the Murgon fossil
[66]
site have been tentatively attributed to that of a salamander, though its true
identity is disputed. If the vertebrae truly belong to a salamander, they would
represent the only salamanders in Australia.

[67][68]
There are about 760 living species of salamander. One-third of the known
salamander species are found in North America. The highest concentration of these
is found in the Appalachian Mountains region, where the Plethodontidae are thought
to have originated in mountain streams. Here, vegetation zones and proximity to
water are of greater importance than altitude. Only species that adopted a more
terrestrial mode of life have been able to disperse to other localities. The northern
slimy salamander (Plethodon glutinosus) has a wide range and occupies a habitat
similar to that of the southern gray-cheeked salamander (Plethodon metcalfi). The
latter is restricted to the slightly cooler and wetter conditions in north-facing cove
forests in the southern Appalachians, and to higher elevations above 900 m (3,000
ft), while the former is more adaptable, and would be perfectly able to inhabit these
locations, but some unknown factor seems to prevent the two species from
[37]
co-existing.

One species, the Anderson's salamander, is one of the few species of living
[69]
amphibians to occur in brackish or salt water.

Reproduction and development[edit]

See also: Sexual selection in amphibians

Sierra newt amplexus found in stream at Woolman Semester in Nevada County, California

[70]
Many salamanders do not use vocalisations, and in most species the sexes look
alike, so they use olfactory and tactile cues to identify potential mates, and sexual
selection occurs. Pheromones play an important part in the process and may be
produced by the abdominal gland in males and by the cloacal glands and skin in
both sexes. Males are sometimes to be seen investigating potential mates with their
snouts. In Old World newts, Triturus spp., the males are sexually dimorphic and
display in front of the females. Visual cues are also thought to be important in some
[71]
plethodont species.

Except for terrestrial species in the three families Plethodontidae, Ambystomatidae,

[72]
and Salamandridae, salamanders mate in water. The mating varies from
[73]
courtship between a single male and female to explosive group breeding. In the
clade Salamandroidea, which makes up about 90% of all species, fertilization is
 [74]
internal. As a general rule, salamanders with internal fertilization have indirect
sperm transfer, but in species like the Sardinian brook salamander, the Corsican
brook salamander, the Caucasian salamander and the Pyrenean brook salamander,
[75][76][77]
the male transfers his sperm directly into the female cloaca. For the species
with indirect sperm transfer, the male deposits a spermatophore on the ground or in
the water according to species, and the female picks this up with her vent. The
spermatophore has a packet of sperm supported on a conical gelatinous base, and
often an elaborate courtship behavior is involved in its deposition and collection.
Once inside the cloaca, the spermatozoa move to the spermatheca, one or more
chambers in the roof of the cloaca, where they are stored for sometimes lengthy
periods until the eggs are laid. In the Asiatic salamanders, the giant salamanders
and Sirenidae, which are the most primitive groups, the fertilization is external. In a
reproductive process similar to that of typical frogs, the male releases sperm onto
the egg mass. These salamanders also have males that exhibit parental care, which
[71][78]
otherwise only occur in females with internal fertilization.

Three different types of egg deposition occur. Ambystoma and Taricha spp. spawn
large numbers of small eggs in quiet ponds where many large predators are unlikely.
Most dusky salamanders (Desmognathus) and Pacific giant salamanders
(Dicamptodon) lay smaller batches of medium-sized eggs in a concealed site in
flowing water, and these are usually guarded by an adult, normally the female. Many
of the tropical climbing salamanders (Bolitoglossa) and lungless salamanders
(Plethodontinae) lay a small number of large eggs on land in a well-hidden spot,
[71]
where they are also guarded by the mother. Some species such as the fire
salamanders (Salamandra) are ovoviviparous, with the female retaining the eggs
inside her body until they hatch, either into larvae to be deposited in a water body, or
[8]
into fully formed juveniles.

Duration: 14 minutes and 53 seconds.

14:53

Embryonic development of a salamander, filmed in the 1920s

In temperate regions, reproduction is usually seasonal and salamanders may

migrate to breeding grounds. Males usually arrive first and in some instances set up
territories. Typically, a larval stage follows in which the organism is fully aquatic. The
tadpole has three pairs of external gills, no eyelids, a long body, a laterally flattened
tail with dorsal and ventral fins and in some species limb-buds or limbs. Pond-type
larvae may have a pair of rod-like balancers on either side of the head, long gill
filaments and broad fins. Stream-type larvae are more slender with short gill
filaments—in Rhyacotriton and Onychodactylus, and some species in
[79]
Batrachuperus, the gills and gill rakers are extremely reduced, narrower fins and
 [80]
no balancers, but instead have hind limbs already developed when they hatch.
The tadpoles are carnivorous and the larval stage may last from days to years,
depending on species. Sometimes this stage is completely bypassed, and the eggs
of most lungless salamanders (Plethodontidae) develop directly into miniature
[81]
versions of the adult without an intervening larval stage.

By the end of the larval stage, the tadpoles already have limbs and metamorphosis
takes place normally. In salamanders, this occurs over a short period of time and
involves the closing of the gill slits and the loss of structures such as gills and tail fins
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 larva
[82]
emerges onto land as a terrestrial adult.

Neotenic axolotl, showing 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
universally possible in all salamander species. In this state, an individual may retain
gills or other juvenile features while attaining reproductive maturity. The changes that
take place at metamorphosis are under the control of thyroid hormones and in
obligate neotenes such as the axolotl (Ambystoma mexicanum), the tissues are
seemingly unresponsive to the hormones. In other species, the changes may not be
triggered because of underactivity of the hypothalamus-pituitary-thyroid mechanism
which may occur when conditions in the terrestrial environment are too
[82]
inhospitable. This may be due to cold or wildly fluctuating temperatures, aridity,
lack of food, lack of cover, or insufficient iodine for the formation of thyroid hormones.
Genetics may also play a part. The larvae of tiger salamanders (Ambystoma
tigrinum), for example, develop limbs soon after hatching and in seasonal pools
promptly 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 metamorphose at all, and become sexually
mature while in their larval forms. Neoteny allows the species to survive even when
[80]
the terrestrial environment is too harsh for the adults to thrive on land.

Conservation[edit]

The threatened hellbender

A general decline in living amphibian species has been linked with the fungal
disease chytridiomycosis. A higher proportion of salamander species than of frogs or
caecilians are in one of the at-risk categories established by the IUCN. Salamanders
showed a significant diminution in numbers in the last few decades of the 20th
century, although no direct link between the fungus and the population decline has
[83]
yet been found. The IUCN made further efforts in 2005 as they established the
Amphibian Conservation Action Plan (ACAP), which was subsequently followed by
Amphibian Ark (AArk), Amphibian Specialist Group (ASG), and finally the umbrella
[84]
organization known as the Amphibian Survival Alliance (ASA). Researchers also
cite deforestation, resulting in fragmentation of suitable habitats, and climate change
as possible contributory factors. Species such as Pseudoeurycea brunnata and
Pseudoeurycea goebeli that had been abundant in the cloud forests of Guatemala
[85]
and Mexico during the 1970s were found by 2009 to be rare. Few data have been
gathered on population sizes over the years and, by intensive surveying of historic
and suitable new locations, it has been possible to locate individuals of other
[83]
species, such as Parvimolge townsendi, which had been thought to be extinct.
Currently, the major lines of defense for the conservation of salamanders includes
both in situ and ex situ conservation methods. There are efforts in place for certain
members of the salamander family to be conserved under a conservation breeding
program (CBP) but there should be research done ahead of time to determine if the
salamander species is actually going to value from the CBP, as researchers have
[84]
noted that some species of amphibians completely fail in this environment.

Various conservation initiatives are being attempted around the world. The Chinese
giant salamander, at 1.8 m (6 ft) the largest amphibian in the world, is critically
endangered, as it is collected for food and for use in traditional Chinese medicine. An
environmental education programme is being undertaken to encourage sustainable
management of wild populations in the Qinling Mountains and captive breeding
[86]
programmes have been set up. The hellbender is another large, long-lived
species with dwindling numbers and fewer juveniles reaching maturity than
[87]
previously. Another alarming finding is the increase in abnormalities in up to 90%
[88]
of the hellbender population in the Spring River watershed in Arkansas. Habitat
loss, silting of streams, pollution and disease have all been implicated in the decline
and a captive breeding programme at Saint Louis Zoo has been successfully
[89]
established. Of the 20 species of minute salamanders (Thorius spp.) in Mexico,
half are believed to have become extinct and most of the others are critically
endangered. Specific reasons for the decline may include climate change,
chytridiomycosis, or volcanic activity, but the main threat is habitat destruction as
logging, agricultural activities, and human settlement reduce their often tiny,
fragmented ranges. Survey work is being undertaken to assess the status of these
salamanders, and to better understand the factors involved in their population
[90]
declines, with a view to taking action.

Ambystoma mexicanum, an aquatic salamander, is a species protected under the

Mexican UMA (Unit for Management and conservation of wildlife) as of April 1994.
Another detrimental factor is that the axolotl lost their role as a top predator since the
introduction of locally exotic species such as Nile tilapia and carp. Tilapia and carp
directly compete with axolotls by consuming their eggs, larvae, and juveniles.
Climate change has also immensely affected axolotls and their populations
throughout the southern Mexico area. Due to its proximity to Mexico City, officials are
currently working on programs at Lake Xochimilco to bring in tourism and educate
[91]
the local population on the restoration of the natural habitat of these creatures.
This proximity is a large factor that has impacted the survival of the axolotl, as the
city has expanded to take over the Xochimilco region in order to make use of its
[92]
resources for water and provision and sewage. It is farmed for use in research
facilities and so may one day return to its natural habitat. The recent decline in
population has substantially impacted genetic diversity among populations, making it
difficult to further progress scientifically. Some genetic indiversity due to
paedeomorphism in Ambystoma species such as the axolotl does not account for the
overall lack of diversity. Evidence points toward a historical bottlenecking of
Ambystoma that contributes to the variation issues and no longer a large genetic
[93]
pool for it to pull from, thus raising concern for inbreeding due to lack of gene flow.
One way researchers are looking into maintaining 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 into a deep freeze for preservation. Most importantly, they have
found that there is only limited damage done to the spermatophores upon thawing
and thus it is a viable option. As of 2013, it is a method that is being used to save not
[92][94][95]
only the axolotl but also numerous other members of the salamander family.

Research is being done on the environmental cues that have to be replicated before
captive animals can be persuaded to breed. Common species such as the tiger
salamander and the mudpuppy are being given hormones to stimulate the
production of sperm and eggs, and the role of arginine vasotocin in courtship
behaviour is being investigated. Another line of research is artificial insemination,
either in vitro or by inserting spermatophores into the cloacae of females. The results
of this research may be used in captive-breeding programmes for endangered
[96]
species.

Taxonomy[edit]
The order name Urodela comes from the name Urodèles given by André Marie
[2]
Constant Duméril in 1805, it is derived from the Greek words οὐρά ourā́ "tail" and
[97]
δῆλος dēlos "visible, conspicuous" because of their "persistent" tails.

Disagreement exists among different authorities as to the definition of the terms

Caudata and Urodela. Some maintain that the Urodela should be restricted to the
[2][citation needed]
crown group, with the Caudata being used for the total group. Others
restrict the name Caudata to the crown group and use Urodela for the total
[98][99]
group. The former approach seems to be most widely adopted and is used in
[68]
this article.

[98]
The ten families belonging to Urodela are divided into three suborders. The clade
Neocaudata is often used to separate the Cryptobranchoidea and Salamandroidea
from the Sirenoidea.