ectotherm
An ectotherm (from the Greek () "outside" and () "heat") is an organism in which internal physiological sources of heat are of relatively small or of quite negligible importance in controlling body temperature.Davenport, John. Animal Life a ...
ic
vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s of the
class
Class or The Class may refer to:
Common uses not otherwise categorized
* Class (biology), a taxonomic rank
* Class (knowledge representation), a collection of individuals or objects
* Class (philosophy), an analytical concept used differentl ...
Amphibia. All living amphibians belong to the group
Lissamphibia
The Lissamphibia is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia (frogs, toads, and their extinct relatives), the Caudata (salamanders, newts, and their extinct relatives), ...
. They inhabit a wide variety of
habitat
In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s, with most species living within
terrestrial
Terrestrial refers to things related to land or the planet Earth.
Terrestrial may also refer to:
* Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
,
fossorial
A fossorial () animal is one adapted to digging which lives primarily, but not solely, underground. Some examples are badgers, naked mole-rats, clams, meerkats, and mole salamanders, as well as many beetles, wasps, and bees.
Prehistoric eviden ...
,
arboreal
Arboreal locomotion is the Animal locomotion, locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. Th ...
or freshwater
aquatic ecosystems
An aquatic ecosystem is an ecosystem formed by surrounding a body of water, in contrast to land-based terrestrial ecosystems. Aquatic ecosystems contain communities of organisms that are dependent on each other and on their environment. The tw ...
. Thus amphibians typically start out as
larva
A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle.
The ...
e living in water, but some species have developed behavioural adaptations to bypass this.
The young generally undergo
metamorphosis
Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
from larva with gills to an adult air-breathing form with
lung
The lungs are the primary organs of the respiratory system in humans and most other animals, including some snails and a small number of fish. In mammals and most other vertebrates, two lungs are located near the backbone on either side of t ...
s. Amphibians use their skin as a secondary respiratory surface and some small terrestrial
salamander
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 ...
s and frogs lack lungs and rely entirely on their skin. They are superficially similar to
reptile
Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s like
lizard
Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...
s but, along with mammals and birds, reptiles are
amniote
Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...
s and do not require water bodies in which to breed. With their complex reproductive needs and permeable skins, amphibians are often
ecological indicator
Ecological indicators are used to communicate information about ecosystems and the impact human activity has on ecosystems to groups such as the public or government policy makers. Ecosystems are complex and ecological indicators can help describe ...
s; in recent decades there has been a dramatic
decline in amphibian populations
Since the 1980s, decreases in amphibian populations, including population decline and localized mass extinctions, have been observed in locations all over the world. These declines are known as one of the most critical threats to global biodiversi ...
for many species around the globe.
The earliest amphibians
evolved
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation t ...
in the
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
period from
sarcopterygian
Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includi ...
fish with lungs and bony-limbed fins, features that were helpful in adapting to dry land. They diversified and became dominant during the
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
and
Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
periods, but were later displaced by reptiles and other vertebrates. The origin of modern amphibians belonging to Lissamphibia, which first appeared during the Early Triassic, around 250 million years ago, has long been contentious. However the emerging consensus is that they likely originated from
temnospondyls
Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carbo ...
, the most diverse group of prehistoric amphibians, during the Permian period.
The three modern orders of amphibians are Anura (the frogs),
Urodela
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 ...
(the salamanders), and Apoda (the caecilians), a fourth group, the
Albanerpetontidae
The Albanerpetontidae are an extinct family of small amphibians, native to the Northern Hemisphere during the Mesozoic and Cenozoic. The only members of the order Allocaudata, they are thought to be allied with living amphibians belonging to L ...
, became extinct around 2 million years ago. The number of known amphibian species is approximately 8,000, of which nearly 90% are frogs. The smallest amphibian (and vertebrate) in the world is a frog from
New Guinea
New Guinea (; Hiri Motu
Hiri Motu, also known as Police Motu, Pidgin Motu, or just Hiri, is a language of Papua New Guinea, which is spoken in surrounding areas of Port Moresby (Capital of Papua New Guinea).
It is a simplified version of ...
(''
Paedophryne amauensis
''Paedophryne amauensis'' is a species of microhylid frog endemic to eastern Papua New Guinea. At in snout-to-vent length, it is considered the world's smallest known vertebrate. (See also Ecological guild.)
The species was listed in the '' ...
'') with a length of just . The largest living amphibian is the
South China giant salamander
The South China giant salamander (''Andrias sligoi'') may be the largest species of salamander and the largest amphibian in the world. It is Endemism, endemic to southern China, mainly in the Pearl River (China), Pearl River basin south of the Na ...
(''Andrias sligoi''), but this is dwarfed by prehistoric temnospondyls such as ''
Mastodonsaurus
''Mastodonsaurus'' (meaning "teat tooth lizard") is an extinct genus of temnospondyl amphibian from the Middle Triassic of Europe. It belongs to a Triassic group of temnospondyls called Capitosauria, characterized by their large body size and pre ...
'' which could reach up to 6 metres in length. The study of amphibians is called
batrachology
Batrachology is the branch of zoology concerned with the study of amphibians including frogs and toads, salamanders, newts, and caecilians. It is a sub-discipline of herpetology, which also includes non-avian reptiles (snakes, lizards, amphisbaen ...
, while the study of both reptiles and amphibians is called
herpetology
Herpetology (from Greek ἑρπετόν ''herpetón'', meaning "reptile" or "creeping animal") is the branch of zoology concerned with the study of amphibians (including frogs, toads, salamanders, newts, and caecilians (gymnophiona)) and rept ...
.
Classification
The word ''amphibian'' is derived from the
Ancient Greek
Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
term (), which means 'both kinds of life', meaning 'of both kinds' and meaning 'life'. The term was initially used as a general adjective for animals that could live on land or in water, including seals and otters. Traditionally, the class Amphibia includes all tetrapod vertebrates that are not amniotes. Amphibia in its widest sense () was divided into three subclasses, two of which are extinct:
*Subclass
Lepospondyli
Lepospondyli is a diverse taxon of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco (''Diplocaulus minumus''), lepospondyls lived from the Early Carboniferous ( Mississippian) to the Early Per ...
† (small Paleozoic group, which are more closely related to amniotes than Lissamphibia)
* Subclass
Temnospondyli
Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carb ...
† (diverse Paleozoic and early Mesozoic grade)
* Subclass
Lissamphibia
The Lissamphibia is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia (frogs, toads, and their extinct relatives), the Caudata (salamanders, newts, and their extinct relatives), ...
(all modern amphibians, including frogs, toads, salamanders, newts and caecilians)
** Salientia (frogs,
toad
Toad is a common name for certain frogs, especially of the family Bufonidae, that are characterized by dry, leathery skin, short legs, and large bumps covering the parotoid glands.
A distinction between frogs and toads is not made in scientif ...
s and relatives): Jurassic to present—7,360 current species in 53 families
** Caudata (
salamander
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 ...
s,
newt
A newt is a salamander in the subfamily Pleurodelinae. The terrestrial juvenile phase is called an eft. Unlike other members of the family Salamandridae, newts are semiaquatic, alternating between aquatic and terrestrial habitats. Not all aqua ...
s and relatives): Jurassic to present—764 current species in 9 families
** Gymnophiona (
caecilian
Caecilians (; ) are a group of limbless, vermiform or serpentine amphibians. They mostly live hidden in the ground and in stream substrates, making them the least familiar order of amphibians. Caecilians are mostly distributed in the tropics of ...
s and relatives): Jurassic to present—215 current species in 10 families
**Allocaudata† (
Albanerpetontidae
The Albanerpetontidae are an extinct family of small amphibians, native to the Northern Hemisphere during the Mesozoic and Cenozoic. The only members of the order Allocaudata, they are thought to be allied with living amphibians belonging to L ...
) Middle Jurassic – Early Pleistocene
The actual number of species in each group depends on the taxonomic classification followed. The two most common systems are the classification adopted by the website AmphibiaWeb,
University of California, Berkeley
The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California) is a public land-grant research university in Berkeley, California. Established in 1868 as the University of California, it is the state's first land-grant u ...
, and the classification by
herpetologist
Herpetology (from Greek ἑρπετόν ''herpetón'', meaning "reptile" or "creeping animal") is the branch of zoology concerned with the study of amphibians (including frogs, toads, salamanders, newts, and caecilians (gymnophiona)) and rept ...
Darrel Frost
Darrel Richmond Frost (born 1951) is an American herpetologist and systematist. He was previously head curator of herpetology at the American Museum of Natural History, as well as president of both the Society for the Study of Amphibians and Re ...
and the
American Museum of Natural History
The American Museum of Natural History (abbreviated as AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. In Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 26 inter ...
, available as the online reference database "Amphibian Species of the World". The numbers of species cited above follows Frost and the total number of known amphibian species as of March 31, 2019, is exactly 8,000, of which nearly 90% are frogs.
With the
phylogenetic
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
classification, the taxon
Labyrinthodontia
"Labyrinthodontia" (Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally consid ...
has been discarded as it is a polyparaphyletic group without unique defining features apart from shared primitive characteristics. Classification varies according to the preferred phylogeny of the author and whether they use a stem-based or a node-based classification. Traditionally, amphibians as a class are defined as all tetrapods with a larval stage, while the group that includes the common ancestors of all living amphibians (frogs, salamanders and caecilians) and all their descendants is called Lissamphibia. The phylogeny of Paleozoic amphibians is uncertain, and Lissamphibia may possibly fall within extinct groups, like the Temnospondyli (traditionally placed in the subclass Labyrinthodontia) or the Lepospondyli, and in some analyses even in the amniotes. This means that advocates of
phylogenetic nomenclature
Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional approach, in which taxon names are defined by a '' type'', which ...
have removed a large number of basal Devonian and Carboniferous amphibian-type tetrapod groups that were formerly placed in Amphibia in
Linnaean taxonomy
Linnaean taxonomy can mean either of two related concepts:
# The particular form of biological classification (taxonomy) set up by Carl Linnaeus, as set forth in his ''Systema Naturae'' (1735) and subsequent works. In the taxonomy of Linnaeus t ...
, and included them elsewhere under
cladistic taxonomy
Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived chara ...
. If the common ancestor of amphibians and amniotes is included in Amphibia, it becomes a paraphyletic group.
All modern amphibians are included in the subclass Lissamphibia, which is usually considered a
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
, a group of species that have evolved from a common ancestor. The three modern orders are Anura (the frogs), Caudata (or Urodela, the salamanders), and Gymnophiona (or Apoda, the caecilians). It has been suggested that salamanders arose separately from a Temnospondyl-like ancestor, and even that caecilians are the sister group of the advanced
reptiliomorph
Reptiliomorpha (meaning reptile-shaped; in PhyloCode known as ''Pan-Amniota'') is a clade containing the amniotes and those tetrapods that share a more recent common ancestor with amniotes than with living amphibians ( lissamphibians). It was de ...
amphibians, and thus of amniotes. Although the fossils of several older proto-frogs with primitive characteristics are known, the oldest "true frog" is '' Prosalirus bitis'', from the
Early Jurassic
The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
Kayenta Formation
The Kayenta Formation is a geological formation in the Glen Canyon Group that is spread across the Colorado Plateau province of the United States, including northern Arizona, northwest Colorado, Nevada, and Utah. Traditionally has been suggested ...
of Arizona. It is anatomically very similar to modern frogs. The oldest known caecilian is another Early Jurassic species, ''
Eocaecilia micropodia
''Eocaecilia'' is an extinct genus of gymnophionan amphibian from the early Jurassic Kayenta Formation of Arizona, United States. One species is described, ''Eocaecilia micropodia''.
''Eocaecilia'' shared some characteristics with salamanders ...
'', also from Arizona. The earliest salamander is ''
Beiyanerpeton jianpingensis
''Beiyanerpeton'' is an extinct genus of salamandroid amphibians known from the Late Jurassic of western Liaoning Province, China. It contains a single species, ''B. jianpingensis''.
Discovery
''Beiyanerpeton'' is known from the holotype specime ...
'' from the
Late Jurassic
The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987.
In European lithostratigraphy, the name ...
of northeastern China.
Authorities disagree as to whether Salientia is a superorder that includes the order Anura, or whether Anura is a sub-order of the order Salientia. The Lissamphibia are traditionally divided into three
orders
Order, ORDER or Orders may refer to:
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
, but an extinct salamander-like family, the Albanerpetontidae, is now considered part of Lissamphibia alongside the superorder Salientia. Furthermore, Salientia includes all three recent orders plus the Triassic proto-frog, ''
Triadobatrachus
''Triadobatrachus'' is an extinct genus of salientian frog-like amphibians, including only one known species, ''Triadobatrachus massinoti''. It is the oldest member of the frog lineage known, and an excellent example of a transitional fossil. It ...
''.
Evolutionary history
The first major groups of amphibians developed in the
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
period, around 370 million years ago, from
lobe-finned fish
Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includ ...
which were similar to the modern
coelacanth
The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
and
lungfish
Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...
. These ancient lobe-finned fish had evolved multi-jointed leg-like fins with digits that enabled them to crawl along the sea bottom. Some fish had developed primitive lungs that help them breathe air when the stagnant pools of the Devonian swamps were low in oxygen. They could also use their strong fins to hoist themselves out of the water and onto dry land if circumstances so required. Eventually, their bony fins would evolve into limbs and they would become the ancestors to all
tetrapod
Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...
s, including modern amphibians, reptiles, birds, and
mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s. Despite being able to crawl on land, many of these prehistoric
tetrapodomorph
The Tetrapodomorpha (also known as Choanata) are a clade of vertebrates consisting of tetrapods (four-limbed vertebrates) and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish. Advance ...
fish still spent most of their time in the water. They had started to develop lungs, but still breathed predominantly with gills.
Many examples of species showing transitional features have been discovered. ''
Ichthyostega
''Ichthyostega'' (from el, ἰχθῦς , 'fish' and el, στέγη , 'roof') is an extinct genus of limbed tetrapodomorphs from the Late Devonian of Greenland. It was among the earliest four-limbed vertebrates in the fossil record, and was on ...
'' was one of the first primitive amphibians, with nostrils and more efficient lungs. It had four sturdy limbs, a neck, a tail with fins and a skull very similar to that of the lobe-finned fish, ''
Eusthenopteron
''Eusthenopteron'' (from el, εὖ , 'good', el, σθένος , 'strength', and el, πτερόν 'wing' or 'fin') is a genus of prehistoric sarcopterygian (often called lobe-finned fishes) which has attained an iconic status from its close ...
''. Amphibians evolved adaptations that allowed them to stay out of the water for longer periods. Their lungs improved and their skeletons became heavier and stronger, better able to support the weight of their bodies on land. They developed "hands" and "feet" with five or more digits; the skin became more capable of retaining body fluids and resisting desiccation. The fish's
hyomandibula
The hyomandibula, commonly referred to as hyomandibular one( la, os hyomandibulare, from el, hyoeides, "upsilon-shaped" (υ), and Latin: mandibula, "jawbone") is a set of bones that is found in the hyoid region in most fishes. It usually plays ...
bone in the
hyoid
The hyoid bone (lingual bone or tongue-bone) () is a horseshoe-shaped bone situated in the anterior midline of the neck between the chin and the thyroid cartilage. At rest, it lies between the base of the mandible and the third cervical vertebra. ...
region behind the gills diminished in size and became the
stapes
The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foot ...
of the amphibian ear, an adaptation necessary for hearing on dry land. An affinity between the amphibians and the
teleost
Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tel ...
fish is the multi-folded structure of the teeth and the paired supra-occipital bones at the back of the head, neither of these features being found elsewhere in the animal kingdom.
At the end of the Devonian period (360 million years ago), the seas, rivers and lakes were teeming with life while the land was the realm of early plants and devoid of vertebrates, though some, such as ''Ichthyostega'', may have sometimes hauled themselves out of the water. It is thought they may have propelled themselves with their forelimbs, dragging their hindquarters in a similar manner to that used by the
elephant seal
Elephant seals are very large, oceangoing earless seals in the genus ''Mirounga''. Both species, the northern elephant seal (''M. angustirostris'') and the southern elephant seal (''M. leonina''), were hunted to the brink of extinction for oil ...
. In the early
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
(360 to 345 million years ago), the climate became wet and warm. Extensive swamps developed with
moss
Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
es,
fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
s,
horsetail
''Equisetum'' (; horsetail, snake grass, puzzlegrass) is the only living genus in Equisetaceae, a family of ferns, which reproduce by spores rather than seeds.
''Equisetum'' is a "living fossil", the only living genus of the entire subclass Eq ...
s and
calamites
''Calamites'' is a genus of extinct arborescent (tree-like) horsetails to which the modern horsetails (genus ''Equisetum'') are closely related. Unlike their herbaceous modern cousins, these plants were medium-sized trees, growing to heights o ...
. Air-breathing
arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s evolved and invaded the land where they provided food for the
carnivorous
A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
amphibians that began to adapt to the terrestrial environment. There were no other tetrapods on the land and the amphibians were at the top of the food chain, occupying the ecological position currently held by the crocodile. Though equipped with limbs and the ability to breathe air, most still had a long tapering body and strong tail. They were the top land predators, sometimes reaching several metres in length, preying on the large insects of the period and the many types of fish in the water. They still needed to return to water to lay their shell-less eggs, and even most modern amphibians have a fully aquatic larval stage with gills like their fish ancestors. It was the development of the amniotic egg, which prevents the developing embryo from drying out, that enabled the reptiles to reproduce on land and which led to their dominance in the period that followed.
After the Carboniferous rainforest collapse amphibian dominance gave way to reptiles, and amphibians were further devastated by the
Permian–Triassic extinction event
The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, as ...
. During the
Triassic Period
The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
(250 to 200 million years ago), the reptiles continued to out-compete the amphibians, leading to a reduction in both the amphibians' size and their importance in the
biosphere
The biosphere (from Greek βίος ''bíos'' "life" and σφαῖρα ''sphaira'' "sphere"), also known as the ecosphere (from Greek οἶκος ''oîkos'' "environment" and σφαῖρα), is the worldwide sum of all ecosystems. It can also be ...
. According to the fossil record,
Lissamphibia
The Lissamphibia is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia (frogs, toads, and their extinct relatives), the Caudata (salamanders, newts, and their extinct relatives), ...
, which includes all modern amphibians and is the only surviving lineage, may have branched off from the extinct groups
Temnospondyli
Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carb ...
and
Lepospondyli
Lepospondyli is a diverse taxon of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco (''Diplocaulus minumus''), lepospondyls lived from the Early Carboniferous ( Mississippian) to the Early Per ...
at some period between the Late Carboniferous and the Early Triassic. The relative scarcity of fossil evidence precludes precise dating, but the most recent molecular study, based on
multilocus sequence typing
Multilocus sequence typing (MLST) is a technique in molecular biology for the typing of multiple loci, using DNA sequences of internal fragments of multiple housekeeping genes to characterize isolates of microbial species.
The first MLST scheme ...
, suggests a Late Carboniferous/
Early Permian 01 or '01 may refer to:
* The year 2001, or any year ending with 01
* The month of January
* 1 (number)
Music
* '01 (Richard Müller album), 01'' (Richard Müller album), 2001
* 01 (Son of Dave album), ''01'' (Son of Dave album), 2000
* 01 (Urban ...
origin for extant amphibians.
The origins and evolutionary relationships between the three main groups of amphibians is a matter of debate. A 2005 molecular phylogeny, based on rDNA analysis, suggests that salamanders and caecilians are more closely related to each other than they are to frogs. It also appears that the divergence of the three groups took place in the
Paleozoic
The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon.
The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838
by combining the Greek words ''palaiós'' (, "old") and ' ...
or early
Mesozoic
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
(around 250 million years ago), before the breakup of the supercontinent
Pangaea
Pangaea or Pangea () was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous approximately 335 million y ...
and soon after their divergence from the lobe-finned fish. The briefness of this period, and the swiftness with which radiation took place, would help account for the relative scarcity of primitive amphibian fossils. There are large gaps in the
fossil record
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
, the discovery of the
dissorophoid
Dissorophoideans are a clade of medium-sized, temnospondyl amphibians that appeared during the Moscovian in Euramerica, and continued through to the Late Permian and the Early Triassic of Gondwana. They are distinguished by various details of the ...
temnospondyl ''Gerobatrachus'' from the Early Permian in Texas in 2008 provided a missing link with many of the characteristics of modern frogs.Molecular analysis suggests that the frog–salamander divergence took place considerably earlier than the
palaeontological
Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (geology), epoch (roughly 11,700 years before present). It includes t ...
evidence indicates. One study suggested suggested that the last common ancestor of all modern amphibians lived about 315 million years ago, and that stereospondyl temnospondyls are the closest relatives to the caecilians. However, most studies support a single
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
origin of all modern amphibians within the dissorophoid temnospondyls.
As they evolved from lunged fish, amphibians had to make certain adaptations for living on land, including the need to develop new means of locomotion. In the water, the sideways thrusts of their tails had propelled them forward, but on land, quite different mechanisms were required. Their vertebral columns, limbs, limb girdles and musculature needed to be strong enough to raise them off the ground for locomotion and feeding. Terrestrial adults discarded their
lateral line
The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial ...
systems and adapted their sensory systems to receive stimuli via the medium of the air. They needed to develop new methods to regulate their body heat to cope with fluctuations in ambient temperature. They developed behaviours suitable for reproduction in a terrestrial environment. Their skins were exposed to harmful
ultraviolet
Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nanometer, nm (with a corresponding frequency around 30 Hertz, PHz) to 400 nm (750 Hertz, THz), shorter than that of visible light, but longer than ...
rays that had previously been absorbed by the water. The skin changed to become more protective and prevent excessive water loss.
Characteristics
The superclass Tetrapoda is divided into four classes of vertebrate animals with four limbs. Reptiles, birds and mammals are amniotes, the eggs of which are either laid or carried by the female and are surrounded by several membranes, some of which are impervious. Lacking these membranes, amphibians require water bodies for reproduction, although some species have developed various strategies for protecting or bypassing the vulnerable aquatic larval stage. They are not found in the sea with the exception of one or two frogs that live in
brackish water
Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estua ...
in
mangrove
A mangrove is a shrub or tree that grows in coastal saline water, saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evoluti ...
swamps; the
Anderson's salamander
Anderson's salamander (''Ambystoma andersoni'') is a neotenic salamander from Zacapu Lagoon in the Mexican state of Michoacán.
This salamander is a relatively recent discovery, first described by Branden and Krebs in 1984. ''Ambystoma andersoni ...
meanwhile occurs in brackish or salt water lakes. On land, amphibians are restricted to moist habitats because of the need to keep their skin damp.
Modern amphibians have a simplified anatomy compared to their ancestors due to
paedomorphosis
Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compare ...
, caused by two evolutionary trends: miniaturization and an unusually large genome, which result in a slower growth and development rate compared to other vertebrates. Another reason for their size is associated with their rapid metamorphosis, which seems to have evolved only in the ancestors of lissamphibia; in all other known lines the development was much more gradual. Because a remodeling of the feeding apparatus means they don't eat during the metamorphosis, the metamorphosis has to go faster the smaller the individual is, so it happens at an early stage when the larvae are still small. (The largest species of salamanders don't go through a metamorphosis.) Amphibians that lay eggs on land often go through the whole metamorphosis inside the egg. An anamniotic terrestrial egg is less than 1 cm in diameter due to diffusion problems, a size which puts a limit on the amount of posthatching growth.
The smallest amphibian (and vertebrate) in the world is a microhylid frog from
New Guinea
New Guinea (; Hiri Motu
Hiri Motu, also known as Police Motu, Pidgin Motu, or just Hiri, is a language of Papua New Guinea, which is spoken in surrounding areas of Port Moresby (Capital of Papua New Guinea).
It is a simplified version of ...
(''
Paedophryne amauensis
''Paedophryne amauensis'' is a species of microhylid frog endemic to eastern Papua New Guinea. At in snout-to-vent length, it is considered the world's smallest known vertebrate. (See also Ecological guild.)
The species was listed in the '' ...
'') first discovered in 2012. It has an average length of and is part of a genus that contains four of the world's ten smallest frog species. The largest living amphibian is the
Chinese giant salamander
The Chinese giant salamander (''Andrias davidianus'') is one of the largest salamanders and one of the largest amphibians in the world. but this is a great deal smaller than the largest amphibian that ever existed—the extinct ''
Prionosuchus
''Prionosuchus'' is an extinct genus of large temnospondyl. A single species ''P. plummeri'', is recognized from the Early Permian (some time between 299 and 272 million years ago). Its fossils have been found in what is now northeastern Brazil.
...
'', a crocodile-like temnospondyl dating to 270 million years ago from the middle Permian of Brazil. The largest frog is the African
Goliath frog
The goliath frog (''Conraua goliath''), otherwise known commonly as the giant slippery frog and the goliath bullfrog, is a species of frog in the family Conrauidae. The goliath frog is the largest living frog. Specimens can grow up to in lengt ...
(''Conraua goliath''), which can reach and weigh .
Amphibians are ectothermic (cold-blooded) vertebrates that do not maintain their body temperature through internal
physiological
Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical ...
processes. Their
metabolic rate
Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
is low and as a result, their food and energy requirements are limited. In the adult state, they have tear ducts and movable eyelids, and most species have ears that can detect airborne or ground vibrations. They have muscular tongues, which in many species can be protruded. Modern amphibians have fully
ossified
Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by Cell (biology), cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes ...
vertebrae with
articular processes
The articular processes or zygapophyses (Greek ζυγον = "yoke" (because it links two vertebrae) + απο = "away" + φυσις = "process") of a vertebra are projections of the vertebra that serve the purpose of fitting with an adjacent vertebr ...
. Their ribs are usually short and may be fused to the vertebrae. Their skulls are mostly broad and short, and are often incompletely ossified. Their skin contains little
keratin
Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, ho ...
and lacks scales, apart from a few fish-like scales in certain caecilians. The skin contains many
mucous gland
Mucous gland, also known as muciparous glands, are found in several different parts of the body, and they typically stain lighter than serous glands during standard histological preparation. Most are multicellular, but goblet cells are single-cel ...
s and in some species, poison glands (a type of granular gland). The hearts of amphibians have three chambers, two atria and one ventricle. They have a
urinary bladder
The urinary bladder, or simply bladder, is a hollow organ in humans and other vertebrates that stores urine from the kidneys before disposal by urination. In humans the bladder is a distensible organ that sits on the pelvic floor. Urine enters ...
and nitrogenous waste products are excreted primarily as
urea
Urea, also known as carbamide, is an organic compound with chemical formula . This amide has two amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest amide of carbamic acid.
Urea serves an important r ...
. Most amphibians lay their eggs in water and have aquatic larvae that undergo metamorphosis to become terrestrial adults. Amphibians breathe by means of a pump action in which air is first drawn into the buccopharyngeal region through the nostrils. These are then closed and the air is forced into the lungs by contraction of the throat. They supplement this with
gas exchange
Gas exchange is the physical process by which gases move passively by Diffusion#Diffusion vs. bulk flow, diffusion across a surface. For example, this surface might be the air/water interface of a water body, the surface of a gas bubble in a liqui ...
through the skin.
Anura
The order Anura (from the Ancient Greek '' a(n)-'' meaning "without" and ''oura'' meaning "tail") comprises the frogs and toads. They usually have long hind limbs that fold underneath them, shorter forelimbs, webbed toes with no claws, no tails, large eyes and glandular moist skin. Members of this order with smooth skins are commonly referred to as frogs, while those with warty skins are known as toads. The difference is not a formal one taxonomically and there are numerous exceptions to this rule. Members of the family Bufonidae are known as the "true toads". Frogs range in size from the
Goliath frog
The goliath frog (''Conraua goliath''), otherwise known commonly as the giant slippery frog and the goliath bullfrog, is a species of frog in the family Conrauidae. The goliath frog is the largest living frog. Specimens can grow up to in lengt ...
(''Conraua goliath'') of West Africa to the ''
Paedophryne amauensis
''Paedophryne amauensis'' is a species of microhylid frog endemic to eastern Papua New Guinea. At in snout-to-vent length, it is considered the world's smallest known vertebrate. (See also Ecological guild.)
The species was listed in the '' ...
'', first described in Papua New Guinea in 2012, which is also the smallest known vertebrate. Although most species are associated with water and damp habitats, some are specialised to live in trees or in deserts. They are found worldwide except for polar areas.
Anura is divided into three suborders that are broadly accepted by the scientific community, but the relationships between some families remain unclear. Future molecular genetics, molecular studies should provide further insights into their evolutionary relationships. The suborder Archaeobatrachia contains four families of primitive frogs. These are Ascaphidae, Bombinatoridae, Discoglossidae and Leiopelmatidae which have few derived features and are probably paraphyletic with regard to other frog lineages. The six families in the more evolutionarily advanced suborder Mesobatrachia are the
fossorial
A fossorial () animal is one adapted to digging which lives primarily, but not solely, underground. Some examples are badgers, naked mole-rats, clams, meerkats, and mole salamanders, as well as many beetles, wasps, and bees.
Prehistoric eviden ...
Megophryidae, Pelobatidae, Pelodytidae, Scaphiopodidae and Rhinophrynidae and the obligatorily aquatic Pipidae. These have certain characteristics that are intermediate between the two other suborders. Neobatrachia is by far the largest suborder and includes the remaining families of modern frogs, including most common species. Ninety-six percent of the over 5,000 extant species of frog are neobatrachians.
Caudata
The order Caudata (from the Latin ''cauda'' meaning "tail") consists of the salamanders—elongated, low-slung animals that mostly resemble lizards in form. This is a Symplesiomorphy, symplesiomorphic trait and they are no more closely related to lizards than they are to mammals. Salamanders lack claws, have scale-free skins, either smooth or covered with tubercles, and tails that are usually flattened from side to side and often finned. They range in size from the
Chinese giant salamander
The Chinese giant salamander (''Andrias davidianus'') is one of the largest salamanders and one of the largest amphibians in the world. South America was apparently invaded from Central America by about the start of the Miocene, 23 million years ago. Urodela is a name sometimes used for all the extant taxon, extant species of salamanders. Members of several salamander families have become Neoteny, paedomorphic and either fail to complete their metamorphosis or retain some larval characteristics as adults. Most salamanders are under long. They may be terrestrial or aquatic and 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.
The suborder Cryptobranchoidea contains the primitive salamanders. A number of fossil cryptobranchids have been found, but there are only three living species, the Chinese giant salamander (''Andrias davidianus''), the Japanese giant salamander (''Andrias japonicus'') and the hellbender (''Cryptobranchus alleganiensis'') from North America. These large amphibians retain several larval characteristics in their adult state; gills slits are present and the eyes are unlidded. A unique feature is their ability to feed by suction, depressing either the left side of their lower jaw or the right. The males excavate nests, persuade females to lay their egg strings inside them, and guard them. As well as breathing with lungs, they respire through the many folds in their thin skin, which has Capillary, capillaries close to the surface.
The suborder Salamandroidea contains the advanced salamanders. They differ from the cryptobranchids by having fused Mandible, prearticular bones in the lower jaw, and by using internal fertilisation. In salamandrids, the male deposits a bundle of sperm, the spermatophore, and the female picks it up and inserts it into her cloaca where the sperm is stored until the eggs are laid. The largest family in this group is Plethodontidae, the lungless salamanders, which includes 60% of all salamander species. The Family (biology), family Salamandridae includes the true salamanders and the name "
newt
A newt is a salamander in the subfamily Pleurodelinae. The terrestrial juvenile phase is called an eft. Unlike other members of the family Salamandridae, newts are semiaquatic, alternating between aquatic and terrestrial habitats. Not all aqua ...
" is given to members of its subfamily Pleurodelinae.
The third suborder, Sirenoidea, contains the four species of sirens, which are in a single family, Sirenidae. Members of this order are eel-like aquatic salamanders with much reduced forelimbs and no hind limbs. Some of their features are primitive while others are derived. Fertilisation is likely to be external as sirenids lack the cloacal glands used by male salamandrids to produce spermatophores and the females lack spermathecae for sperm storage. Despite this, the eggs are laid singly, a behaviour not conducive for external fertilisation.
Gymnophiona
The order Gymnophiona (from the Greek ''gymnos'' meaning "naked" and ''ophis'' meaning "serpent") or Apoda comprises the caecilians. These are long, cylindrical, limbless animals with a snake- or Annelid, worm-like form. The adults vary in length from 8 to 75 centimetres (3 to 30 inches) with the exception of Caecilia thompsoni, Thomson's caecilian (''Caecilia thompsoni''), which can reach . A caecilian's skin has a large number of transverse folds and in some species contains tiny embedded dermal scales. It has rudimentary eyes covered in skin, which are probably limited to discerning differences in light intensity. It also has a pair of short tentacles near the eye that can be extended and which have Somatosensory system, tactile and Olfaction, olfactory functions. Most caecilians live underground in burrows in damp soil, in rotten wood and under plant debris, but some are aquatic. Most species lay their eggs underground and when the larvae hatch, they make their way to adjacent bodies of water. Others brood their eggs and the larvae undergo metamorphosis before the eggs hatch. A few species give birth to live young, nourishing them with glandular secretions while they are in the oviduct. Caecilians have a mostly Gondwanan distribution, being found in tropical regions of Africa, Asia and Central and South America.
Anatomy and physiology
Skin
The Wikt:integument, integumentary structure contains some typical characteristics common to terrestrial vertebrates, such as the presence of highly Keratin#Cornification, cornified outer layers, renewed periodically through a moulting process controlled by the pituitary gland, 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 mostly in one piece, in contrast to mammals and birds where it is shed in flakes. Amphibians often eat the sloughed skin. Caecilians are unique among amphibians in having mineralized dermal scales embedded in the dermis between the furrows in the skin. The similarity of these to the scales of bony fish is largely superficial. Squamata, Lizards and some frogs have somewhat similar osteoderms forming bony deposits in the dermis, but this is an example of convergent evolution with similar structures having arisen independently in diverse vertebrate lineages.
Amphibian skin is permeable to water. Gas exchange can take place through the skin (cutaneous respiration) and this allows adult amphibians to respire without rising to the surface of water and to hibernate at the bottom of ponds. To compensate for their thin and delicate skin, amphibians have evolved mucous glands, principally on their heads, backs and tails. The secretions produced by these help keep the skin moist. In addition, most species of amphibian have granular glands that secrete distasteful or poisonous substances. Some amphibian toxins can be lethal to humans while others have little effect. The main poison-producing glands, the Parotoid gland, parotoids, produce the neurotoxin bufotoxin and are located behind the ears of toads, along the backs of frogs, behind the eyes of salamanders and on the upper surface of caecilians.
The skin colour 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 colour) and the lipophores (yellow, the most superficial layer). The colour change displayed by many species is initiated by hormones secreted by the pituitary gland. Unlike bony fish, there is no direct control of the pigment cells by the nervous system, and this results in the colour change taking place more slowly than happens in fish. A vividly coloured skin usually indicates that the species is toxic and is a warning sign to predators.
Skeletal system and locomotion
Amphibians have a skeletal system that is structurally Homology (biology), homologous to other tetrapods, though with a number of variations. They all have four limbs except for the legless caecilians and a few species of salamander with reduced or no limbs. The bones are hollow and lightweight. The musculoskeletal system is strong to enable it to support the head and body. The bones are fully ossification, ossified and the vertebrae interlock with each other by means of overlapping processes. The pectoral girdle is supported by muscle, and the well-developed pelvic girdle is attached to the backbone by a pair of sacral ribs. The ilium (bone), ilium slopes forward and the body is held closer to the ground than is the case in mammals.
In most amphibians, there are four digits on the fore foot and five on the hind foot, but no claws on either. Some salamanders have fewer digits and the amphiumas are eel-like in appearance with tiny, stubby legs. The Siren (genus), sirens are aquatic salamanders with stumpy forelimbs and no hind limbs. The caecilians are limbless. They burrow in the manner of earthworms with zones of muscle contractions moving along the body. On the surface of the ground or in water they move by undulating their body from side to side.
In frogs, the hind legs are larger than the fore legs, especially so in those species that principally move by jumping or swimming. In the walkers and runners the hind limbs are not so large, and the burrowers mostly have short limbs and broad bodies. The feet have adaptations for the way of life, with webbing between the toes for swimming, broad adhesive toe pads for climbing, and keratinised tubercles on the hind feet for digging (frogs usually dig backwards into the soil). In most salamanders, the limbs are short and more or less the same length and project at right angles from the body. Locomotion on land is by walking and the tail often swings from side to side or is used as a prop, particularly when climbing. In their normal gait, only one leg is advanced at a time in the manner adopted by their ancestors, the lobe-finned fish. Some salamanders in the genus ''Aneides'' and certain lungless salamander, plethodontids climb trees and have long limbs, large toepads and prehensile tails. In aquatic salamanders and in frog tadpoles, the tail has dorsal fin, dorsal and ventral fin, ventral fins and is moved from side to side as a means of propulsion. Adult frogs do not have tails and caecilians have only very short ones.
Salamanders use their tails in defence and some are prepared to jettison them to save their lives in a process known as autotomy. Certain species in the Plethodontidae have a weak zone at the base of the tail and use this strategy readily. The tail often continues to twitch after separation which may distract the attacker and allow the salamander to escape. Both tails and limbs can be regenerated. Adult frogs are unable to regrow limbs but tadpoles can do so.
Circulatory system
Amphibians have a juvenile stage and an adult stage, and the circulatory systems of the two are distinct. In the juvenile (or tadpole) stage, the circulation is similar to that of a fish; the two-chambered heart pumps the blood through the gills where it is oxygenated, and is spread around the body and back to the heart in a single loop. In the adult stage, amphibians (especially frogs) lose their gills and develop lungs. They have a heart that consists of a single ventricle and two atria. When the ventricle starts contracting, deoxygenated blood is pumped through the pulmonary artery to the lungs. Continued contraction then pumps oxygenated blood around the rest of the body. Mixing of the two bloodstreams is minimized by the anatomy of the chambers.
Nervous and sensory systems
The nervous system is basically the same as in other vertebrates, with a central brain, a spinal cord, and nerves throughout the body. The amphibian brain is relatively simple but broadly the same structurally as in reptiles, birds and mammals. Their brains are elongated, except in caecilians, and contain the usual motor and sensory areas of tetrapods. The pineal body, known to regulate sleep patterns in humans, is thought to produce the hormones involved in hibernation and aestivation in amphibians.
Tadpoles retain the lateral line system of their ancestral fishes, but this is lost in terrestrial adult amphibians. Some caecilians possess Electroreception, electroreceptors that allow them to locate objects around them when submerged in water. The ears are well developed in frogs. There is no external ear, but the large circular Tympanum (anatomy), eardrum lies on the surface of the head just behind the eye. This vibrates and sound is transmitted through a single bone, the
stapes
The ''stapes'' or stirrup is a bone in the middle ear of humans and other animals which is involved in the conduction of sound vibrations to the inner ear. This bone is connected to the oval window by its annular ligament, which allows the foot ...
, to the inner ear. Only high-frequency sounds like mating calls are heard in this way, but low-frequency noises can be detected through another mechanism. There is a patch of specialized haircells, called ''papilla amphibiorum'', in the inner ear capable of detecting deeper sounds. Another feature, unique to frogs and salamanders, is the columella-operculum 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 than those of frogs as they do not normally communicate with each other through the medium of sound.
The eyes of tadpoles lack lids, but at metamorphosis, the cornea becomes more dome-shaped, the Lens (anatomy), lens becomes flatter, and eyelids and associated glands and ducts develop. The adult eyes 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 wavelengths.
Digestive and excretory systems
Many amphibians catch their prey by flicking out an elongated tongue with a sticky tip and drawing it back into the mouth before seizing the item with their jaws. Some use inertial feeding to help them swallow the prey, repeatedly thrusting their head forward sharply causing the food to move backwards in their mouth by inertia. Most amphibians swallow their prey whole without much chewing so they possess voluminous stomachs. The short Esophagus, oesophagus is lined with Cilium, cilia that help to move the food to the stomach and mucus produced by glands in the mouth and pharynx eases its passage. The enzyme chitinase produced in the stomach helps digest the chitinous cuticle of arthropod prey.
Amphibians possess a pancreas, liver and gall bladder. The liver is usually large with two lobes. Its size is determined by its function as a glycogen and fat storage unit, and may change with the seasons as these reserves are built or used up. Adipose tissue is another important means of storing energy and this occurs in the abdomen (in internal structures called fat bodies), under the skin and, in some salamanders, in the tail.
There are two kidneys located dorsally, near the roof of the body cavity. Their job is to filter the blood of metabolic waste and transport the urine via ureters to the urinary bladder where it is stored before being passed out periodically through the cloacal vent. Larvae and most aquatic adult amphibians excrete the nitrogen as ammonia in large quantities of dilute urine, while terrestrial species, with a greater need to conserve water, excrete the less toxic product urea. Some tree frogs with limited access to water excrete most of their metabolic waste as uric acid.
Respiratory system
The lungs in amphibians are primitive compared to those of amniotes, possessing few internal alveolar septum, septa and large Pulmonary alveolus, 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 vascularised skin must remain moist to allow the oxygen to diffuse at a sufficiently high rate. 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 Telmatobius culeus, Titicaca water frog and the Cryptobranchus, hellbender salamander. In air, where oxygen is more concentrated, some small species can rely solely on cutaneous gas exchange, most famously the Plethodontidae, 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.
Reproduction
For the purpose of Biological reproduction, reproduction most amphibians require fresh water although some lay their eggs on land and have developed various means of keeping them moist. A few (e.g. ''Fejervarya raja'') can inhabit brackish water, but there are no true marine (ocean), marine amphibians. There are reports, however, of particular amphibian populations unexpectedly invading marine waters. Such was the case with the Black Sea invasion of the natural hybrid ''Pelophylax esculentus'' reported in 2010.
Several hundred frog species in adaptive radiations (e.g., ''Eleutherodactylus'', the Pacific ''Platymantis'', 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 and subtropical moist broadleaf forests, 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.
In the tropics, many amphibians breed continuously or at any time of year. In temperate regions, breeding is mostly seasonal, usually in the spring, and is triggered by increasing day length, rising temperatures or rainfall. Experiments have shown the importance of temperature, but the trigger event, especially in arid regions, is often a storm. In anurans, males usually arrive at the breeding sites before females and the vocal chorus they produce may stimulate ovulation in females and the endocrine activity of males that are not yet reproductively active.
In caecilians, fertilisation is internal, the male extruding an intromittent organ, the , and inserting it into the female cloaca. The paired Müllerian glands inside the male cloaca secrete a fluid which resembles that produced by mammalian prostate glands and which may transport and nourish the sperm. Fertilisation probably takes place in the oviduct.
The majority of salamanders also engage in internal fertilisation. In most of these, the male deposits a spermatophore, a small packet of sperm on top of a gelatinous cone, on the Substrate (biology), substrate either on land or in the water. The female takes up the sperm packet by grasping it with the lips of the cloaca and pushing it into the vent. The spermatozoa move to the spermatheca in the roof of the cloaca where they remain until ovulation which may be many months later. Courtship rituals and methods of transfer of the spermatophore vary between species. In some, the spermatophore may be placed directly into the female cloaca while in others, the female may be guided to the spermatophore or restrained with an embrace called amplexus. Certain primitive salamanders in the families Sirenidae, Hynobiidae and Cryptobranchidae practice external fertilisation in a similar manner to frogs, with the female laying the eggs in water and the male releasing sperm onto the egg mass.
With a few exceptions, frogs use external fertilisation. The male grasps the female tightly with his forelimbs either behind the arms or in front of the back legs, or in the case of ''Epipedobates tricolor'', around the neck. They remain in amplexus with their cloacae positioned close together while the female lays the eggs and the male covers them with sperm. Roughened nuptial pads on the male's hands aid in retaining grip. Often the male collects and retains the egg mass, forming a sort of basket with the hind feet. An exception is the granular poison frog (''Oophaga granulifera'') where the male and female place their cloacae in close proximity while facing in opposite directions and then release eggs and sperm simultaneously. The tailed frog (''Ascaphus truei'') exhibits internal fertilisation. The "tail" is only possessed by the male and is an extension of the cloaca and used to inseminate the female. This frog lives in fast-flowing streams and internal fertilisation prevents the sperm from being washed away before fertilisation occurs. The sperm may be retained in storage tubes attached to the oviduct until the following spring.
Most frogs can be classified as either prolonged or explosive breeders. Typically, prolonged breeders congregate at a breeding site, the males usually arriving first, calling and setting up territories. Other satellite males remain quietly nearby, waiting for their opportunity to take over a territory. The females arrive sporadically, mate selection takes place and eggs are laid. The females depart and territories may change hands. More females appear and in due course, the breeding season comes to an end. Explosive breeders on the other hand are found where temporary pools appear in dry regions after rainfall. These frogs are typically wikt:fossorial, fossorial species that emerge after heavy rains and congregate at a breeding site. They are attracted there by the calling of the first male to find a suitable place, perhaps a pool that forms in the same place each rainy season. The assembled frogs may call in unison and frenzied activity ensues, the males scrambling to mate with the usually smaller number of females.
There is a direct competition between males to win the attention of the females in salamanders and newts, with elaborate courtship displays to keep the female's attention long enough to get her interested in choosing him to mating, mate with. Some species store sperm through long breeding seasons, as the extra time may allow for interactions with rival sperm.
Life cycle
Most amphibians go through
metamorphosis
Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
, 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 thyroxine concentration in the blood, which stimulates metamorphosis, and prolactin, which counteracts thyroxine's effect. Specific events are dependent on threshold values for different tissues. Because most embryonic development is outside the parental body, it is subject to many adaptations due to specific environmental circumstances. For this reason tadpoles can have horny ridges instead of Teeth, whisker-like skin extensions or fins. They also make use of a sensory lateral line organ similar to that of fish. After metamorphosis, these organs become redundant and will be reabsorbed by controlled cell death, called apoptosis. The variety of adaptations to specific environmental circumstances among amphibians is wide, with many discoveries still being made.
Eggs
In the egg, the embryo is suspended in Perivitelline space, perivitelline fluid and surrounded by semi-permeable gelatinous capsules, with the yolk mass providing nutrients. As the larvae hatch, the capules are dissolved by enzymes secreted from gland at the tip of the snout. The eggs of some salamanders and frogs contain unicellular green algae. These penetrate the jelly envelope after the eggs are laid and may increase the supply of oxygen to the embryo through photosynthesis. They seem to both speed up the development of the larvae and reduce mortality. In the wood frog (''Rana sylvatica''), the interior of the globular egg cluster has been found to be up to warmer than its surroundings, which is an advantage in its cool northern habitat.
The eggs may be deposited singly, in cluster or in long strands. Sites for laying eggs include water, mud, burrows, debris and on plants or under logs or stones. The Eleutherodactylus planirostris, greenhouse frog (''Eleutherodactylus planirostris'') lays eggs in small groups in the soil where they develop in about two weeks directly into juvenile frogs without an intervening larval stage. The tungara frog (''Physalaemus pustulosus'') builds a floating nest from foam to protect its eggs. First a raft is built, then eggs are laid in the centre, and finally a foam cap is overlaid. The foam has anti-microbial properties. It contains no detergents but is created by whipping up proteins and lectins secreted by the female.
Larvae
The eggs of amphibians are typically laid in water and hatch into free-living larvae that complete their development in water and later transform into either aquatic or terrestrial adults. In many species of frog and in most lungless salamanders (Plethodontidae), direct development takes place, the larvae growing within the eggs and emerging as miniature adults. Many caecilians and some other amphibians lay their eggs on land, and the newly hatched larvae wriggle or are transported to water bodies. Some caecilians, the alpine salamander (''Salamandra atra'') and some of the Nectophrynoides, African live-bearing toads (''Nectophrynoides spp.'') are Viviparity, viviparous. Their larvae feed on glandular secretions and develop within the female's oviduct, often for long periods. Other amphibians, but not caecilians, are Ovoviviparity, ovoviviparous. The eggs are retained in or on the parent's body, but the larvae subsist on the yolks of their eggs and receive no nourishment from the adult. The larvae emerge at varying stages of their growth, either before or after metamorphosis, according to their species. The toad genus ''Nectophrynoides'' exhibits all of these developmental patterns among its dozen or so members. Amphibian larvae are known as tadpoles. They have thick, rounded bodies with powerful muscular tails.
Frogs
Unlike in other amphibians, frog tadpoles do not resemble adults. The free-living larvae are normally fully aquatic, but the tadpoles of some species (such as ''Nannophrys ceylonensis'') are semi-terrestrial and live among wet rocks. Tadpoles have cartilaginous skeletons, gills for respiration (external gills at first, internal gills later), lateral line systems and large tails that they use for swimming. Newly hatched tadpoles soon develop gill pouches that cover the gills. The lungs develop early and are used as accessory breathing organs, the tadpoles rising to the water surface to gulp air. Some species complete their development inside the egg and hatch directly into small frogs. These larvae do not have gills but instead have specialised areas of skin through which respiration takes place. While tadpoles do not have true teeth, in most species, the jaws have long, parallel rows of small keratinized structures called keradonts surrounded by a horny beak. Front legs are formed under the gill sac and hind legs become visible a few days later.
Iodine and T4 (over stimulate the spectacular apoptosis [programmed cell death] of the cells of the larval gills, tail and fins) also stimulate the evolution of nervous systems transforming the aquatic, vegetarian tadpole into the terrestrial, carnivorous frog with better neurological, visuospatial, olfactory and cognitive abilities for hunting.
In fact, tadpoles developing in ponds and streams are typically herbivore, herbivorous. Pond tadpoles tend to have deep bodies, large caudal fins and small mouths; they swim in the quiet waters feeding on growing or loose fragments of vegetation. Stream dwellers mostly have larger mouths, shallow bodies and caudal fins; they attach themselves to plants and stones and feed on the surface films of algae and bacteria. They also feed on diatoms, filtered from the water through the gills, and stir up the sediment at bottom of the pond, ingesting edible fragments. They have a relatively long, spiral-shaped gut to enable them to digest this diet. Some species are carnivorous at the tadpole stage, eating insects, smaller tadpoles and fish. Young of the Cuban tree frog (''Osteopilus septentrionalis'') can occasionally be Cannibalism (zoology), cannibalistic, the younger tadpoles attacking a larger, more developed tadpole when it is undergoing metamorphosis.
At metamorphosis, rapid changes in the body take place 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 large jaw, and its gills disappear along with its gill sac. Eyes and legs grow quickly, and a tongue is formed. There are associated changes in the neural networks such as development of stereoscopic vision and loss of the lateral line system. All this can happen in about a day. A few days later, the tail is reabsorbed, due to the higher thyroxine concentration required for this to take place.
Salamanders
At hatching, a typical salamander larva has eyes without lids, teeth in both upper and lower jaws, three pairs of feathery external gills, and a long tail with Anatomical terms of location#Dorsal and ventral, dorsal and Anatomical terms of location#Dorsal and ventral, ventral fins. The forelimbs may be partially developed and the hind limbs are rudimentary in pond-living species but may be rather more developed in species that reproduce in moving water. Pond-type larvae often have a pair of balancers, rod-like structures on either side of the head that may prevent the gills from becoming clogged up with sediment. Both of these are able to breed. Some have larvae that never fully develop into the adult form, a condition known as neoteny. Neoteny occurs when the animal's growth rate is very low and is usually linked to adverse conditions such as low water temperatures that may change the response of the tissues to the hormone thyroxine. as well as lack of food. There are fifteen species of wikt:obligate, obligate neotenic salamanders, including species of ''Necturus'', ''Olm, Proteus'' and ''Amphiuma'', and many examples of wikt:facultative, facultative ones, such as the northwestern salamander (''Ambystoma gracile'') and the tiger salamander (''A. tigrinum'') that adopt this strategy under appropriate environmental circumstances.
Lungless salamanders in the family Plethodontidae are terrestrial and lay a small number of unpigmented eggs in a cluster among damp leaf litter. Each egg has a large yolk sac and the larva feeds on this while it develops inside the egg, emerging fully formed as a juvenile salamander. The female salamander often broods the eggs. In the genus ''Ensatinas'', the female has been observed to coil around them and press her throat area against them, effectively massaging them with a mucous secretion.
In newts and salamanders, metamorphosis is less dramatic than in frogs. This is because the larvae are already carnivorous and continue to feed as predators when they are adults so few changes are needed to their digestive systems. Their lungs are functional early, but the larvae do not make as much use of them as do tadpoles. Their gills are never covered by gill sacs and are reabsorbed just before the animals leave the water. Other changes include the reduction in size or loss of tail fins, the closure of gill slits, thickening of the skin, the development of eyelids, and certain changes in dentition and tongue structure. Salamanders are at their most vulnerable at metamorphosis as swimming speeds are reduced and transforming tails are encumbrances on land. Adult salamanders 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, thyroxine. External gills do not return in subsequent aquatic phases because these are completely absorbed upon leaving the water for the first time.
Caecilians
Most terrestrial caecilians that lay eggs do so in burrows or moist places on land near bodies of water. The development of the young of ''Ichthyophis glutinosus'', a species from Sri Lanka, has been much studied. The eel-like larvae hatch out of the eggs and make their way to water. They have three pairs of external red feathery gills, a blunt head with two rudimentary eyes, a lateral line system and a short tail with fins. They swim by undulating their body from side to side. They are mostly active at night, soon lose their gills and make sorties onto land. Metamorphosis is gradual. By the age of about ten months they have developed a pointed head with sensory tentacles near the mouth and lost their eyes, lateral line systems and tails. The skin thickens, embedded scales develop and the body divides into segments. By this time, the caecilian has constructed a burrow and is living on land.
In the majority of species of caecilians, the young are produced by viviparity. ''Typhlonectes compressicauda'', a species from South America, is typical of these. Up to nine larvae can develop in the oviduct at any one time. They are elongated and have paired sac-like gills, small eyes and specialised scraping teeth. At first, they feed on the yolks of the eggs, but as this source of nourishment declines they begin to rasp at the ciliated epithelial cells that line the oviduct. This stimulates the secretion of fluids rich in lipids and mucoproteins on which they feed along with scrapings from the oviduct wall. They may increase their length sixfold and be two-fifths as long as their mother before being born. By this time they have undergone metamorphosis, lost their eyes and gills, developed a thicker skin and mouth tentacles, and reabsorbed their teeth. A permanent set of teeth grow through soon after birth.
The ringed caecilian (''Siphonops annulatus'') has developed a unique adaptation for the purposes of reproduction. The progeny feed on a skin layer that is specially developed by the adult in a phenomenon known as maternal dermatophagy. The brood feed as a batch for about seven minutes at intervals of approximately three days which gives the skin an opportunity to regenerate. Meanwhile, they have been observed to ingest fluid exuded from the maternal cloaca.
Parental care
The care of offspring among amphibians has been little studied but, in general, the larger the number of eggs in a batch, the less likely it is that any degree of parental care takes place. Nevertheless, it is estimated that in up to 20% of amphibian species, one or both adults play some role in the care of the young. Those species that breed in smaller water bodies or other specialised habitats tend to have complex patterns of behaviour in the care of their young.
Many woodland salamanders lay clutches of eggs under dead logs or stones on land. The black mountain salamander (''Desmognathus welteri'') does this, the mother brooding the eggs and guarding them from predation as the embryos feed on the yolks of their eggs. When fully developed, they break their way out of the egg capsules and disperse as juvenile salamanders. The male hellbender, a primitive salamander, excavates an underwater nest and encourages females to lay there. The male then guards the site for the two or three months before the eggs hatch, using body undulations to fan the eggs and increase their supply of oxygen.
The male ''Colostethus subpunctatus'', a tiny frog, protects the egg cluster which is hidden under a stone or log. When the eggs hatch, the male transports the tadpoles on his back, stuck there by a mucous secretion, to a temporary pool where he dips himself into the water and the tadpoles drop off. The male Common midwife toad, midwife toad (''Alytes obstetricans'') winds egg strings round his thighs and carries the eggs around for up to eight weeks. He keeps them moist and when they are ready to hatch, he visits a pond or ditch and releases the tadpoles. The female gastric-brooding frog (''Rheobatrachus spp.'') reared larvae in her stomach after swallowing either the eggs or hatchlings; however, this stage was never observed before the species became extinct. The tadpoles secrete a hormone that inhibits digestion in the mother whilst they develop by consuming their very large yolk supply. The pouched frog (''Assa darlingtoni'') lays eggs on the ground. When they hatch, the male carries the tadpoles around in brood pouches on his hind legs. The aquatic Surinam toad (''Pipa pipa'') raises its young in pores on its back where they remain until metamorphosis. The granular poison frog (''Oophaga granulifera'') is typical of a number of tree frogs in the poison dart frog family Dendrobatidae. Its eggs are laid on the forest floor and when they hatch, the tadpoles are carried one by one on the back of an adult to a suitable water-filled crevice such as the wikt:axil, axil of a leaf or the Rosette (botany), rosette of a Bromeliaceae, bromeliad. The female visits the nursery sites regularly and deposits unfertilised eggs in the water and these are consumed by the tadpoles.
Genetics and genomics
Amphibians are notable among vertebrates for their diversity of chromosomes and genomes. The karyotypes (chromosomes) have been determined for at least 1,193 (14.5%) of the ≈8,200 known (Ploidy, diploid) species, including 963 Frog, anurans, 209
salamander
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 ...
s, and 21
caecilian
Caecilians (; ) are a group of limbless, vermiform or serpentine amphibians. They mostly live hidden in the ground and in stream substrates, making them the least familiar order of amphibians. Caecilians are mostly distributed in the tropics of ...
s. Generally, the karyotypes of diploid amphibians are characterized by 20–26 bi-armed chromosomes. Amphibians have also very large genomes compared to other taxa of vertebrates and corresponding variation in genome size (C-value: picograms of DNA in Ploidy, haploid nuclei). The genome sizes range from 0.95 to 11.5 pg in frogs, from 13.89 to 120.56 pg in salamanders, and from 2.94 to 11.78 pg in caecilians.
The large genome sizes have prevented whole-Whole genome sequencing, genome sequencing of amphibians although a number of genomes have been published recently. The 1.7GB draft genome of ''Western clawed frog, Xenopus tropicalis'' was the first to be reported for amphibians in 2010. Compared to some salamanders this frog genome is tiny. For instance, the genome of the Axolotl, Mexican axolotl turned out to be 32 Gb, which is more than 10 times larger than the human genome (3GB).
Feeding and diet
With a few exceptions, adult amphibians are Predation, predators, feeding on virtually anything that moves that they can swallow. The diet mostly consists of small prey that do not move too fast such as beetles, caterpillars, earthworms and spiders. The sirens (''Siren (genus), Siren spp.'') often ingest aquatic plant material with the invertebrates on which they feed and a Brazilian tree frog (''Xenohyla truncata'') includes a large quantity of fruit in its diet. The Mexican burrowing toad (''Rhinophrynus dorsalis'') has a specially adapted tongue for picking up ants and termites. It projects it with the tip foremost whereas other frogs flick out the rear part first, their tongues being hinged at the front.
Food is mostly selected by sight, even in conditions of dim light. Movement of the prey triggers a feeding response. Frogs have been caught on fish hooks baited with red flannel and Rana clamitans, green frogs (''Rana clamitans'') have been found with stomachs full of elm seeds that they had seen floating past. Toads, salamanders and caecilians also use smell to detect prey. This response is mostly secondary because salamanders have been observed to remain stationary near odoriferous prey but only feed if it moves. Cave-dwelling amphibians normally hunt by smell. Some salamanders seem to have learned to recognize immobile prey when it has no smell, even in complete darkness.
Amphibians usually swallow food whole but may chew it lightly first to subdue it. They typically have small hinged pedicellate teeth, a feature unique to amphibians. The base and crown of these are composed of dentine separated by an Calcification, uncalcified layer and they are replaced at intervals. Salamanders, caecilians and some frogs have one or two rows of teeth in both jaws, but some frogs (''Rana spp.'') lack teeth in the lower jaw, and toads (''Bufo spp.'') have no teeth. In many amphibians there are also vomerine teeth attached to a facial bone in the roof of the mouth.
The tiger salamander (''Ambystoma tigrinum'') is typical of the frogs and salamanders that hide under cover ready to ambush unwary invertebrates. Others amphibians, such as the ''Bufo spp.'' toads, actively search for prey, while the Argentine horned frog (''Ceratophrys ornata'') lures inquisitive prey closer by raising its hind feet over its back and vibrating its yellow toes. Among leaf litter frogs in Panama, frogs that actively hunt prey have narrow mouths and are slim, often brightly coloured and toxic, while ambushers have wide mouths and are broad and well-camouflaged. Caecilians do not flick their tongues, but catch their prey by grabbing it with their slightly backward-pointing teeth. The struggles of the prey and further jaw movements work it inwards and the caecilian usually retreats into its burrow. The subdued prey is gulped down whole.
When they are newly hatched, frog larvae feed on the yolk of the egg. When this is exhausted some move on to feed on bacteria, algal crusts, detritus and raspings from submerged plants. Water is drawn in through their mouths, which are usually at the bottom of their heads, and passes through branchial food traps between their mouths and their gills where fine particles are trapped in mucus and filtered out. Others have specialised mouthparts consisting of a horny beak edged by several rows of labial teeth. They scrape and bite food of many kinds as well as stirring up the bottom sediment, filtering out larger particles with the papillae around their mouths. Some, such as the spadefoot toads, have strong biting jaws and are carnivorous or even cannibalistic.
file:Sophisticated-Communication-in-the-Brazilian-Torrent-Frog-Hylodes-japi-pone.0145444.s001.oga, left, Audio showing Brazilian torrent frog males executing advertisement, peep, and squeal calls.
Vocalization
The calls made by caecilians and salamanders are limited to occasional soft squeaks, grunts or hisses and have not been much studied. A clicking sound sometimes produced by caecilians may be a means of orientation, as in bats, or a form of communication. Most salamanders are considered voiceless, but the California giant salamander (''Dicamptodon ensatus'') has vocal cords and can produce a rattling or barking sound. Some species of salamander emit a quiet squeak or yelp if attacked.
Frogs are much more vocal, especially during the breeding season when they use their voices to attract mates. The presence of a particular species in an area may be more easily discerned by its characteristic call than by a fleeting glimpse of the animal itself. In most species, the sound is produced by expelling air from the lungs over the vocal cords into an Gular skin, air sac or sacs in the throat or at the corner of the mouth. This may distend like a balloon and acts as a resonator, helping to transfer the sound to the atmosphere, or the water at times when the animal is submerged. The main vocalisation is the male's loud advertisement call which seeks to both encourage a female to approach and discourage other males from intruding on its territory. This call is modified to a quieter courtship call on the approach of a female or to a more aggressive version if a male intruder draws near. Calling carries the risk of attracting predators and involves the expenditure of much energy. Other calls include those given by a female in response to the advertisement call and a release call given by a male or female during unwanted attempts at amplexus. When a frog is attacked, a distress or fright call is emitted, often resembling a scream. The usually nocturnal Cuban tree frog (''Osteopilus septentrionalis'') produces a rain call when there is rainfall during daylight hours.
Territorial behaviour
Little is known of the territorial behaviour of caecilians, but some frogs and salamanders defend home ranges. These are usually feeding, breeding or sheltering sites. Males normally exhibit such behaviour though in some species, females and even juveniles are also involved. Although in many frog species, females are larger than males, this is not the case in most species where males are actively involved in territorial defence. Some of these have specific adaptations such as enlarged teeth for biting or spines on the chest, arms or thumbs.
In salamanders, defence of a territory involves adopting an aggressive posture and if necessary attacking the intruder. This may involve snapping, chasing and sometimes biting, occasionally causing the loss of a tail. The behaviour of red back salamanders (''Plethodon cinereus'') has been much studied. 91% of marked individuals that were later recaptured were within a metre (yard) of their original daytime retreat under a log or rock. A similar proportion, when moved experimentally a distance of , found their way back to their home base. The salamanders left odour marks around their territories which averaged in size and were sometimes inhabited by a male and female pair. These deterred the intrusion of others and delineated the boundaries between neighbouring areas. Much of their behaviour seemed stereotyped and did not involve any actual contact between individuals. An aggressive posture involved raising the body off the ground and glaring at the opponent who often turned away submissively. If the intruder persisted, a biting lunge was usually launched at either the tail region or the naso-labial grooves. Damage to either of these areas can reduce the fitness of the rival, either because of the need to regenerate tissue or because it impairs its ability to detect food.
In frogs, male territorial behaviour is often observed at breeding locations; calling is both an announcement of ownership of part of this resource and an advertisement call to potential mates. In general, a deeper voice represents a heavier and more powerful individual, and this may be sufficient to prevent intrusion by smaller males. Much energy is used in the vocalization and it takes a toll on the territory holder who may be displaced by a fitter rival if he tires. There is a tendency for males to tolerate the holders of neighbouring territories while vigorously attacking unknown intruders. Holders of territories have a "home advantage" and usually come off better in an encounter between two similar-sized frogs. If threats are insufficient, chest to chest tussles may take place. Fighting methods include pushing and shoving, deflating the opponent's vocal sac, seizing him by the head, jumping on his back, biting, chasing, splashing, and ducking him under the water.
Defence mechanisms
Amphibians have soft bodies with thin skins, and lack claws, defensive armour, or spines. Nevertheless, they have evolved various defence mechanisms to keep themselves alive. The first line of defence in salamanders and frogs is the mucous secretion that they produce. This keeps their skin moist and makes them slippery and difficult to grip. The secretion is often sticky and distasteful or toxic. Snakes have been observed yawning and gaping when trying to swallow African clawed frogs (''Xenopus laevis''), which gives the frogs an opportunity to escape. Caecilians have been little studied in this respect, but the Cayenne caecilian (''Typhlonectes compressicauda'') produces toxic mucus that has killed predatory fish in a feeding experiment in Brazil. In some salamanders, the skin is poisonous. The rough-skinned newt (''Taricha granulosa'') from North America and other members of its genus contain the neurotoxin tetrodotoxin (TTX), the most toxic non-protein substance known and almost identical to that produced by pufferfish. Handling the newts does not cause harm, but ingestion of even the most minute amounts of the skin is deadly. In feeding trials, fish, frogs, reptiles, birds and mammals were all found to be susceptible. The only predators with some tolerance to the poison are certain populations of Common Garter Snake, common garter snake (''Thamnophis sirtalis'').
In locations where both snake and salamander co-exist, the snakes have developed immunity through genetic changes and they feed on the amphibians with impunity. Coevolution occurs with the newt increasing its toxic capabilities at the same rate as the snake further develops its immunity. Some frogs and toads are toxic, the main poison glands being at the side of the neck and under the warts on the back. These regions are presented to the attacking animal and their secretions may be foul-tasting or cause various physical or neurological symptoms. Altogether, over 200 toxins have been isolated from the limited number of amphibian species that have been investigated.
Poisonous species often use bright colouring to warn potential predators of their toxicity. These warning colours tend to be red or yellow combined with black, with the fire salamander (''Salamandra salamandra'') being an example. Once a predator has sampled one of these, it is likely to remember the colouration next time it encounters a similar animal. In some species, such as the fire-bellied toad (''Bombina spp.''), the warning colouration is on the belly and these animals adopt a defensive pose when attacked, exhibiting their bright colours to the predator. The frog ''Allobates zaparo'' is not poisonous, but Batesian mimicry, mimics the appearance of other toxic species in its locality, a strategy that may deceive predators.
Many amphibians are nocturnal and hide during the day, thereby avoiding diurnal predators that hunt by sight. Other amphibians use camouflage to avoid being detected. They have various colourings such as mottled browns, greys and olives to blend into the background. Some salamanders adopt defensive poses when faced by a potential predator such as the North American northern short-tailed shrew (''Blarina brevicauda''). Their bodies writhe and they raise and lash their tails which makes it difficult for the predator to avoid contact with their poison-producing granular glands. A few salamanders will autotomise their tails when attacked, sacrificing this part of their anatomy to enable them to escape. The tail may have a constriction at its base to allow it to be easily detached. The tail is regenerated later, but the energy cost to the animal of replacing it is significant.
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. Giant salamanders of the genus ''Andrias'', as well as Ceratophryinae, Ceratophrine and ''Pyxicephalus'' frogs possess sharp teeth and are capable of drawing blood with a defensive bite. The blackbelly salamander (''Desmognathus quadramaculatus'') can bite an attacking common garter snake (''Thamnophis sirtalis'') two or three times its size on the head and often manages to escape.
Cognition
In amphibians, there is evidence of habituation, associative learning through both Classical conditioning, classical and instrumental learning, and discrimination abilities.
In one experiment, when offered live fruit flies (''Drosophila virilis''), salamanders chose the larger of 1 vs 2 and 2 vs 3. Frogs can distinguish between low numbers (1 vs 2, 2 vs 3, but not 3 vs 4) and large numbers (3 vs 6, 4 vs 8, but not 4 vs 6) of prey. This is irrespective of other characteristics, i.e. surface area, volume, weight and movement, although discrimination among large numbers may be based on surface area.
Conservation
Dramatic declines in amphibian populations, including population crashes and mass localized extinction, have been noted since the late 1980s from locations all over the world, and amphibian declines are thus perceived to be one of the most critical threats to global biodiversity. In 2004, the International Union for Conservation of Nature (IUCN) reported stating that currently birds, mammals, and amphibians extinction rates were at minimum 48 times greater than natural extinction rates—possibly 1,024 times higher. In 2006, there were believed to be 4,035 species of amphibians that depended on water at some stage during their life cycle. Of these, 1,356 (33.6%) were considered to be threatened and this figure is likely to be an underestimate because it excludes 1,427 species for which there was insufficient data to assess their status. A number of causes are believed to be involved, including habitat destruction and modification, over-exploitation, pollution, introduced species, global warming, endocrine-disrupting chemical, endocrine-disrupting pollutants, destruction of the ozone layer (ultraviolet radiation has shown to be especially damaging to the skin, eyes, and eggs of amphibians), and diseases like chytridiomycosis. However, many of the causes of amphibian declines are still poorly understood, and are a topic of ongoing discussion.
Food webs and predation
Any decline in amphibian numbers will affect the patterns of predation. The loss of carnivorous species near the top of the food chain will upset the delicate ecosystem balance and may cause dramatic increases in opportunistic species.
Predators that feed on amphibians are affected by their decline. The western terrestrial garter snake (''Thamnophis elegans'') in California is largely aquatic and depends heavily on two species of frog that are decreasing in numbers, the Yosemite toad (''Bufo canorus'') and the mountain yellow-legged frog (''Rana muscosa''), putting the snake's future at risk. If the snake were to become scarce, this would affect birds of prey and other predators that feed on it. Meanwhile, in the ponds and lakes, fewer frogs means fewer tadpoles. These normally play an important role in controlling the growth of algae and also forage on detritus that accumulates as sediment on the bottom. A reduction in the number of tadpoles may lead to an overgrowth of algae, resulting in depletion of oxygen in the water when the algae later die and decompose. Aquatic invertebrates and fish might then die and there would be unpredictable ecological consequences.
Pollution and pesticides
The decline in amphibian and reptile populations has led to an awareness of the effects of pesticides on reptiles and amphibians. In the past, the argument that amphibians or reptiles were more susceptible to any chemical contamination than any land aquatic vertebrate was not supported by research until recently. Amphibians and reptiles have complex life cycles, live in different climate and ecological zones, and are more vulnerable to chemical exposure. Certain pesticides, such as organophosphates, neonicotinoids, and carbamates, react via cholinesterase inhibition. Cholinesterase is an enzyme that causes the hydrolysis of acetylcholine, an excitatory neurotransmitter that is abundant in the nervous system. AChE inhibitors are either reversible or irreversible, and carbamates are safer than organophosphorus insecticides, which are more likely to cause cholinergic poisoning. Reptile exposure to an AChE inhibitory pesticide may result in disruption of neural function in reptiles. The buildup of these inhibitory effects on motor performance, such as food consumption and other activities.
Conservation and protection strategies
The Amphibian Specialist Group of the IUCN is spearheading efforts to implement a comprehensive global strategy for amphibian conservation. Amphibian Ark is an organization that was formed to implement the ex-situ conservation recommendations of this plan, and they have been working with zoos and aquaria around the world, encouraging them to create assurance colonies of threatened amphibians. One such project is the Panama Amphibian Rescue and Conservation Project that built on existing conservation efforts in Panama to create a country-wide response to the threat of chytridiomycosis.
Another measure would be to stop exploitation of frogs for human consumption. In the Middle East, a growing appetite for eating frog legs and the consequent gathering of them for food was already linked to an increase in mosquitoes and thus has direct consequences for human health.
See also
* Cultural depictions of amphibians
* List of amphibians
* List of amphibian genera
* List of threatened reptiles and amphibians of the United States