Amphibian

Amphibians
Temporal range: Late Devonian–
present
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Scientific classification
Kingdom: Animalia
Phylum: Chordata
Superclass: Tetrapoda
Class: Amphibia
Linnaeus, 1758
Subclasses and Orders
Subclass
Labyrinthodontia –
extinct
Subclass Lepospondyli
– extinct
Subclass Lissamphibia
Order Anura
Order Caudata
Order Gymnophiona
Amphibians are members of the class Amphibia, a group of vertebrates whose living forms
include frogs, toads, salamanders, newts and caecilians. They are characterized as non-amniote,
ectothermic tetrapods, meaning their eggs are not surrounded by membranes, they are cold-
blooded, and they have four limbs. Most amphibians lay their eggs in water and the larvae
undergo metamorphosis from a juvenile form with gills to an adult air-breathing form with lungs.
Some, however, are paedomorphs that retain the juvenile water-breathing form throughout life.
Mudpuppies and olms are examples of this, retaining juvenile gills into adulthood. Adult
amphibians also use their skin for respiration.

The three modern orders of amphibians are placed in the subclass Lissamphibia and are the
Anura (frogs and toads), Caudata (salamanders and newts) and Gymnophiona (caecilians,
limbless amphibians that resemble snakes). The total number of known amphibian species is
approximately 6,500.They are superficially similar to reptiles, but reptiles, along with mammals
and birds, are amniotes, having impervious membranes that surround the egg. Amphibians are
ecological indicators,] and in recent decades there has been a dramatic decline in amphibian
populations around the globe. Many species are now threatened or extinct. The study of
amphibians is called batrachology.

The earliest amphibians evolved in the Devonian Period from Sarcopterygians, fish that had
lungs and bony fins, features that were helpful in adapting to dry land. They diversified and
became dominant during the Carboniferous and Permian periods] but were later displaced by
reptiles and other vertebrates. Over time, amphibians shrunk and decreased in diversity, leaving
the modern orders of Lissamphibia.

Contents
 1 Etymology
 2 Characteristics
o 2.1 Anura
o 2.2 Caudata
o 2.3 Gymnophiona
 3 Evolution
 4 Classification
 5 Anatomy and physiology
o 5.1 Integumentary system
o 5.2 Skeletal system
o 5.3 Circulatory and nervous systems
o 5.4 Digestive and excretory systems
o 5.5 Respiratory system
o 5.6 Sensory systems
 6 Reproduction
 7 Growth and development
o 7.1 Frogs and toads
o 7.2 Newts and salamanders
o 7.3 Caecilians
  8 Diet
 9 Vocalization
 10 Defense mechanisms
 11 Conservation
 12 See also
 13 References
 14 Further reading
 15 External links

Etymology
Amphibian is derived from the Ancient Greek term ἀμφίβιος (amphíbios), which means "both
kinds of life", amphi meaning "of both kinds" and bio meaning "life". The term was initially used
generally for animals that could live on land or in water, including seals and otters. It became
restricted to what we now call amphibians around 1600 and the taxon "Amphibia" was first used
in scientific classification circa 1819.[5]

Characteristics
The amphibians are a class of vertebrate animals with four limbs. They are non-amniotes which
means that their eggs are not surrounded by the several membranes, some impervious, which
enable mammals, reptiles and birds to reproduce on land. Amphibians typically reproduce in
fresh water and are not found in the sea, with the exception of one or two frogs that live in
brackish water in mangrove swamps.[6] Most amphibians lay eggs that have a gelatinous coating
which swells when it comes in contact with water. The larvae that hatch from the eggs are mostly
quite dissimilar to the adult form. In the case of frogs and toads they have a large head and a
dorsally flattened tail and are known as tadpoles. They are vegetarians and breathe with gills.
They have no limbs at first, the back limbs thrusting through then skin at a later stage, followed
by the fore limbs after which the tail is reabsorbed. After this metamorphosis the juveniles look
like miniature versions of the adult. Newt and salamander larvae have long bodies and feathery
gills. They are carnivorous and the front legs develop before the back ones. They do not undergo
metamorphosis in the same way that frogs and toads do.[7] The caecilians either produce live
young or lay eggs in damp positions in their burrows.[8]

Amphibians are cold blooded animals and unable to maintain their body temperature above that
of their surroundings. This means that they are only able to be active when the temperature is
high enough. There is great variability in the sensitivity of different species to cold. Many
species hibernate in winter, going into a state of torpor either in an underground chamber or
underwater. In colder climates, they may be in a state of hibernation for more than half the year.
In hot weather they may aestivate underground, sometimes buried in the mud of a dried up pond,
reviving when cooler weather and rain restores their habitat. An advantage of being cold blooded
is that little energy is required to provide body heat. This means that they can go without food for
prolonged periods without coming to harm.[7]
Anura

The order Anura includes the frogs and toads. Members of this order with smooth skins are
commonly referred to as frogs while those with warted skins are known as toads. There are no
taxonomic differences between these two groups but members of the family Bufonidae are
known as true toads. Frogs and toads have broad heads and plump bodies with short, stout fore
limbs and long hind limbs that fold underneath them. Although many species are associated with
water and damp habitats, some are specialised to live in trees and in deserts. They are found
worldwide.

Caudata

The order Caudata includes the salamanders and one of its constituent families, Salamandridae,
includes the true salamanders and the newts. Salamanders and newts have pointed heads, long
cylindrical bodies, four similar sized, short legs and long tails. They may be terrestrial or aquatic
but many spend part of the year in each habitat. When on land, they mostly spend the day hidden
under stones or logs or in dense vegetation, emerging in the evening and night to forage for
worms, insects and other invertebrates. They are found in the Holarctic region of the northern
hemisphere and in Central and South America north of the Amazon Basin.[7]

Gymnophiona

The order Gymnophiona includes the caecilians. These are long, cylindrical, limbless animals
that resemble snakes or worms. Their skin has circular folds which enhances their similarity to
the segments of earthworms. Some are aquatic but most live underground in burrows they
hollow out. Many caecilians give birth to live young, and in the animals that do not do this, the
eggs may undergo metamorphosis before they hatch. Caecilians are found in tropical Africa,
Asia and Central and South America.

Classification
Class Amphibia

 Subclass Lissamphibia
o Family Albanerpetontidae — Jurassic to Miocene (extinct)
o Superorder Salientia
 Genus Triadobatrachus — Triassic (extinct) — A stem Anuran
 Order Anura (frogs and toads): Jurassic to recent — 5,602 recent species
in 48 families
 Order Caudata or Urodela (salamanders, newts): Jurassic to recent — 571
recent species in 10 families
 Order Gymnophiona (caecilians): Jurassic to recent — 190 recent species
in 10 families
The actual number of species partly also depends on the taxonomic classification followed, the
two most common classifications being the classification of the website AmphibiaWeb,
University of California (Berkeley) and the classification by herpetologist Darrel Frost and The
American Museum of Natural History, available as the online reference database Amphibian
Species of the World.[13] The numbers of species cited above follow Frost.

Anatomy and physiology

Integumentary system

The fire salamander has brightly colored yellow spots, indicating that it secretes toxins.

Amphibian skin is permeable to water and contains many mucous glands which keep the skin
from drying out. Gas exchange can take place through the skin and this allows adult amphibians
to hibernate at the bottom of ponds.[7] To compensate for their thin and delicate skin, all
amphibians have evolved poison glands as a defense mechanism, although toxicity varies by
species. Some amphibian toxins can be lethal to humans while others have little effect. [14] The
main poison-producing glands, the paratoids, contain the neurotoxin bufotoxin and are located
behind the ears of certain frogs and toads and behind the eyes of salamanders.[15] The
integumentary structure contains some typical characteristics common to terrestrial vertebrates,
such as the presence of highly cornified outer layers, renewed periodically through a molting
process controlled by the pituitary and thyroid glands. Local thickenings (often called warts) are
common, such as those found on toads. The outside of the skin is shed periodically more or less
in one piece in contrast to mammals and birds where it is shed in flakes. Amphibians often eat
the sloughed skin.[7]

The skin color of amphibians is produced by three layers of pigment cells called
chromatophores. These three cell layers consist of the melanophores (occupying the deepest
layer), the guanophores (forming an intermediate layer and containing many granules, producing
a blue-green color) and the lipophores (yellow, the most superficial layer). The color change
experienced by many species is caused by secretions from the pituitary gland. Unlike bony fish,
there is no direct control by the nervous system of the pigment cells. Therefore, the color change
is slower. Bright colors usually indicate that the species produces an exceptionally toxic poison.

Circulatory and nervous systems

Amphibians have a juvenile stage and an adult stage and the circulatory systems of the two are
distinct. In the juvenile (or tadpole) stage, gills are used to oxygenate blood and movement is
similar to that of fish. In the adult stage, amphibians (especially frogs) lose their gills and
develop lungs. They have a heart that consists of a ventricle and two atria (it may be considered a
single atrium, if not totally or partially divided) that pumps oxygenated blood through arteries
and deoxygenated blood through veins to the lungs. Since amphibians are cold-blooded, they
must find ways to keep their blood at a constant temperature to maintain homeostasis.[18]

The nervous system is basically the same as in other vertebrates, with a central brain, a spinal
cord, and nerves throughout the body.[18] The amphibian brain is less developed compared to that
of reptiles, birds, and mammals. It consists of a cerebrum, midbrain, and cerebellum of similar
sizes. The olfactory lobe is the center of the sense of smell. The cerebrum integrates behavior
and learning. The optic lobe processes information from the eyes. The cerebellum is the center of
muscular coordination. The medulla oblongata controls some organ functions, such as heart rate
and respiration. The brain sends signals through the spinal cord and nerves to regulate activity in
the rest of the body. The pineal body, known to regulate sleep patterns in humans, is thought to
produce the hormones involved in hibernation and estivation in amphibians.[19]

Digestive and excretory systems

Amphibians possess a pancreas, liver and gall bladder. Like mammals, the liver functions as the
central metabolic organ that regulates blood sugar, and also produces the final metabolic
products and transports them through the vascular system to the kidneys, and finally to excretion.
The liver in most amphibians is large with two lobes. The size of the liver is determined by its
vital function as a glycogen and fat storage unit, and may change proportionally with the seasons
with increasing or decreasing activity. In aquatic amphibians, the liver plays only a small role in
processing nitrogen for excretion, and ammonia is diffused mainly through the skin and
excretion. The liver of terrestrial amphibians converts ammonia to urea, a less toxic, water
soluble nitrogenous compound, as a means of water conservation. In some species, urea is
further converted into uric acid. The liver secretions from the liver collect in the gall bladder, and
flow into the small intestine. Salamanders lack a valve separating the small intestine from the
large intestine. In the small intestine, enzymes digest carbohydrates, fats, and proteins. Salt and
water absorption occur in the large intestine, as well as mucous secretion to aid in the transport
of fecal matter, which is passed out through the cloaca. Amphibians have two kidneys located
dorsally, near the roof of the body cavity. Their job is to filter the blood of waste and transport it
to the urinary bladder where it passes out of the cloacal vent.[18]

Respiratory system

The lungs in amphibians are primitive compared to those of amniotes, possessing few internal
septa and large alveoli and consequently having a comparatively slow diffusion rate for oxygen
entering the blood. Ventilation is accomplished by buccal pumping. Most amphibians, however,
are able to exchange gases with the water or air via their skin. To enable sufficient cutaneous
respiration, the surface of their highly vascularized skin must remain moist in order to allow the
oxygen to diffuse at a sufficiently high rate.[18] Because oxygen concentration in the water
increases at both low temperatures and high flow rates, aquatic amphibians in these situations
can rely primarily on cutaneous respiration, as in the Titicaca water frog and the hellbender
salamander. In air, where oxygen is more concentrated, some small species can rely solely on
cutaneous gas exchange, most famously the plethodontid salamanders, which have neither lungs
nor gills. Many aquatic salamanders and all tadpoles have gills in their larval stage, with some
(such as the axolotl) retaining gills as aquatic adults.[18]

Sensory systems

The eyes of amphibians have lids and associated glands and ducts. They are an improvement on
invertebrate eyes and were a first step in the development of more advanced vertebrate eyes.
They allow colour vision and depth of focus. In the retinas are green rods which are receptive to
a wide range of wave lengths.[20]

The tympani, or eardrums, of many frogs are external and lie just behind the eyes. There is also a
patch of papilla amphibiorum in the ear which is unique to amphibians and which can detect low
frequency sounds. Another unique feature is the columella-opercular complex, adjoining the
auditory capsule, which is involved in the transmission of both airborne and seismic signals. The
ears of salamanders and caecilians are less highly developed as they do not normally
communicate with each other by sound.[20]

Reproduction

Frogspawn

For the purpose of reproduction most amphibians require fresh water. A few (e.g. Fejervarya
raja) can inhabit brackish water and even survive (though not thrive) in seawater, but there are
no true marine amphibians. However, there are reports of particular amphibian populations
invading marine waters where their species is normally unable to survive. Such is the case with
the Black Sea invasion of the natural hybrid Pelophylax esculentus reported in 2010.[21]

Several hundred frog species in adaptive radiations (e.g., Eleutherodactylus, the Pacific
Platymantines, the Australo-Papuan microhylids, and many other tropical frogs), however, do
not need any water for breeding in the wild. They reproduce via direct development, an
ecological and evolutionary adaptation that has allowed them to be completely independent from
free-standing water. Almost all of these frogs live in wet tropical rainforests and their eggs hatch
directly into miniature versions of the adult, passing through the tadpole stage within the egg.
Reproductive success of many amphibians is dependent not only on the quantity of rainfall, but
the seasonal timing.[22]
Many amphibians exhibit different kinds of parenting behaviour. After their hatching, the
tadpoles of different species of poison dart frogs (family Dendrobatidae) are carried by the
adults to a suitable place where they can pass metamorphosis. Such places are the rosettes of
many bromeliads in which water is gathered and used by the plant. The Surinam toad raises its
young in pores at its back and after enough time they appear out of these pores fully developed.
The ringed caecilian (Siphonops annulatus) has developed a unique adaptation for the purposes
of reproduction. The progeny feeds on a skin layer that is specially developed by the adult. This
phenomenon is known as maternal dermatophagy.

Several species have also adapted to arid and semi-arid environments, but most of them still need
water to lay their eggs. Symbiosis with single celled algae that lives in the jelly-like layer of the
eggs has evolved several times. The larvae of frogs (tadpoles or polliwogs) breathe with exterior
gills at the start, but soon a pouch is formed that covers the gills and the front legs. Lungs are
also formed quite early to assist in breathing. Newt larvae have large external gills that gradually
disappear and the larvae of newts are quite similar to the adult form from an early age.

Growth and development

Tadpoles

Most amphibians go through metamorphosis, a process of significant morphological change after

birth. In typical amphibian development, eggs are laid in water and larvae are adapted to an
aquatic lifestyle. Frogs, toads and salamanders all hatch from the egg as larvae with external
gills. Metamorphosis in amphibians is regulated by thyroxin concentration in the blood, which
stimulates metamorphosis, and prolactin, which counteracts its effect. Specific events are
dependent on threshold values for different tissues.[23] Because most embryonic development is
outside the parental body, development is subject to many adaptations due to specific ecological
circumstances. For this reason tadpoles can have horny ridges for teeth, whiskers and fins. They
also make use of the lateral line organ. After metamorphosis, these organs become redundant and
will be reabsorbed by controlled cell death, called apoptosis. The extent of adaptations to specific
ecological circumstances among amphibians is remarkable, with many discoveries still being
made.[24]

Frogs and toads

Life cycle of a typical frog

With frogs and toads, the external gills of the newly hatched tadpole are covered with a gill sac
after a few days, and lungs are quickly formed. Front legs are formed under the gill sac, and hind
legs are visible a few days later. Following that there is usually a longer stage during which the
tadpole has a vegetarian diet, scraping algae off plant surfaces with their horny tooth ridges.
Tadpoles have a relatively long, spiral-shaped gut to enable them to digest this diet.[25]

Rapid changes in the body can then be observed as the lifestyle of the frog changes completely.
The spiral‐shaped mouth with horny tooth ridges is reabsorbed together with the spiral gut. The
animal develops a big jaw, and its gills disappear along with its gill sac. Eyes and legs grow
quickly, a tongue is formed, and all this is accompanied by associated changes in the neural
networks (development of stereoscopic vision, loss of the lateral line system, etc.) All this can
happen in about a day, so it is truly a metamorphosis. It isn't until a few days later that the tail is
reabsorbed, due to the higher thyroxin concentrations required for tail resorption.[25]

Newts and salamanders

In newts and salamanders, there is no true metamorphosis because the larvae already feed as
predators and continue doing so as adults. Their gills are never covered by a gill sac and will be
reabsorbed only just before the animal leaves the water. Just as in tadpoles, their lungs are
functional early, but the larvae don't make as much use of them as tadpoles do. They often have
an aquatic phase in spring and summer, and a land phase in winter. For adaptation to a water
phase, prolactin is the required hormone, and for adaptation to the land phase, thyroxin. External
gills do not return in subsequent aquatic phases because these are completely absorbed upon
leaving the water for the first time.[26]

Caecilians

Most caecilians lay eggs in burrows formed in moist soil close to water. The eggs are watched
over by the female and, on hatching, the young make their way into the adjoining pond or
stream. They have gill slits but no external gills. In some species, the young are produced by
vivipary. Up to four eggs begin their development in the oviduct. When their yolk has been
consumed, the developing larvae feed on maternal secretions and on the cells lining the oviduct,
scraping them off with their deciduous teeth.[27]

Diet
Adult amphibians are predators, feeding mainly on live invertebrates. The diet mostly consists of
small items of prey that do not move too fast such as beetles, caterpillars, earthworms and
spiders. Many amphibians have extensible tongues with sticky tips which can be flicked forward
to catch the prey. They usually swallow the food whole but may chew it lightly first in order to
subdue it.[7] The larvae of frogs and toads have a mostly vegetarian diet, using their specialised
mouthparts, consisting of a horny beak edged by several rows of labial teeth, to feed on aquatic
plants.[7]

Vocalization
The noises made by newts, salamanders and caecilians are limited to occasional soft squeaks,
grunts or hisses. Frogs and toads however use their voice in the breeding season to attract mates.
Each call is characteristic of the species, the presence of which in an area may be easier to detect
by its voice than by a fleeting glimpse of the animal itself. In ponds where more than one species
breed, it is important for the female to be attracted to the correct partner as male frogs, in their
ardour, have been known to attempt to mate with other males, females of the wrong species or
inanimate objects.[7] Besides the attracting call, another vocalization is a release call, used by a
frog suffering in this way from advances by the wrong frog. Calling bears the risk of attracting
predators. Some males do not call at all but station themselves near others that are calling, with
the intention of intercepting a female attracted to the sound. Although calling mainly occurs in
the breeding season, sporadic calls may be heard at other times of year in some species.[7]

The calls are made by vibrations of the larynx but are often amplified by vocal sacs which act as
resonators. Sometimes these are fairly small air-filled pouches inside the mouth but in other
species they are much larger and cause a ballooning of the elastic skin of the throat and are
known as external vocal sacs. Many species have single sacs but members of the family Ranidae
have two, one on each side of the mouth.[7]

Defense mechanisms
Amphibians have soft bodies and are relatively helpless. They are eaten by birds and mammals
when on land and by fish when in the water. Many are nocturnal and hide during the day, thereby
avoiding predators that hunt by sight, apart from owls which are also nocturnal. Other
amphibians use camouflage to avoid being detected. They have various colorings such as mottled
browns, greys and olives to make themselves inconspicuous so that they blend into the
background. Other species contain poison glands and use bright colours to warn potential
predators of their toxicity. These are mostly black and yellow or black and red, like the fire
salamander (Salamandra salamandra). Once a predator has tried eating one of these, it is likely
to remember the coloration next time it encounters one. In some species, such as the fire-bellied
toad (Bombina spp.), the warning coloration is on the belly and these animals adopt a defensive
pose when attacked to exhibit the bright colors to the attacker. A few salamanders will
autotomise their tails when attacked, sacrificing this part of their body to enable the main part to
escape. The tail is regenerated later. Some frogs and toads inflate themselves to make themselves
look large and fierce, and some spadefoot toads (Pelobates spp) scream and leap towards the
attacker.[7]