{{taxobox
| color=Pink
| name=Amphibians
| fossil_range=
Late Devonian - Recent
| image = Spea hammondii 1.jpg
| image_width = 230px
| image_caption =
Western Spadefoot Toad, ''Spea hammondii''
| regnum =
Animalia
| phylum =
Chordata
| subphylum =
Vertebrata
| classis =
Amphibia
| classis_authority =
Linnaeus, 1758
| subdivision_ranks = Subclasses and Orders
| subdivision =
Order
Temnospondyli - ''extinct''
Subclass
Lepospondyli - ''extinct''
Subclass
Lissamphibia
Order
Anura
Order
Caudata
Order
Gymnophiona}}
Amphibians (
class Amphibia; from
Greek ''αμφις'' "both" and ''βιος'' "life") are a
taxon of
animals that include all living
tetrapods (four-legged
vertebrates) that do not have
amniotic eggs, are
ectothermic (term for the animals whose body heat is regulated by the external environment; previously known as cold-blooded), and generally spend part of their time on land. Most amphibians do not have the adaptations to an entirely terrestrial existence found in most other modern tetrapods (
amniotes). There are around 6,000 described, living
species of amphibians. The study of amphibians and
reptiles is known as
herpetology. Amphibians are able to breathe through their skin.
Classification
Traditionally the amphibians have included all
tetrapods that are not
amniotes. They are divided into three
subclasses:
Subclass Labyrinthodontia (diverse Paleozoic and early Mesozoic group)
Subclass Lepospondyli (small Paleozoic group)
Subclass Lissamphibia (frogs, salamanders, etc)
Of these only the last includes recent species.
With the
cladistic revolution, this classification has been modified, and the Labyrinthodontia discarded as being a
paraphyletic 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 node-based classification. Generally amphibians are defined as the group that includes the common ancestors of all living amphibians (frogs, salamanders, etc) and all their descendants. This may also include extinct groups like the
temnospondyls (traditionally placed in the disbanded subclass "labyrinthodontia"), and the Lepospondyls. This means that there are a now large number of
basal Devonian and
Carboniferous tetrapod groups, described as "amphibians" in earlier books, that are no longer placed in the formal Amphibia.
All recent amphibians are included in the Lissamphibia, which is usually considered a
clade (which means that it is thought that all Lissamphibians evolved from a common ancestor apart from other extinct groups), although it has also been suggested also that salamanders arose separately from a temnospondyl-like ancestor (Carroll, 2007).
Authorities also disagree on whether Salientia is a Superorder that includes the order Anura, or whether Anura is a sub-order of the order Salientia. In effect Salientia includes all the Anura plus a single
Triassic proto-frog species, ''
Triadobatrachus''. Practical considerations seem to favour using the former arrangement now.
The Lissamphibia are traditionally divided into three
orders, but an extinct salamander-like group, the Albanerpetontidae, is now considered in addition to the other three groups.
Family ''Albanerpetontidae'' - Jurassic to Miocene (extinct)
Superorder ''Salientia''
''Triadobatrachus'' (Triassic)
Order ''Anura'' (frogs and toads): Jurassic to recent - 5,362 recent species
Order ''Caudata'' or ''Urodela'' (salamanders): Jurassic to recent - 556 recent species
Order ''Gymnophiona'' or ''Apoda'' (caecilians): Jurassic to recent - 173 recent species
Systems
Reproductive
For the purpose of
reproduction most amphibians are bound to
fresh water. A few tolerate
brackish water, but there are no true
seawater amphibians. Several hundred frog species in adaptive radiations (e.g., ''
Eleutherodactylus'', the Pacific Platymantines, the Australo-Papuan microhylids, and many other tropical frogs), however, do not need any water whatsoever. They reproduce via direct development, an ecological and
evolutionary adaptation that has allowed them to be completely independent from free-standing water. Almost all of these frogs live in wet
tropical rainforests and their eggs hatch directly into miniature versions of the adult, passing through the
tadpole stage within the egg. Several species have also adapted to arid and semi-arid environments, but most of them still need water to lay their eggs.
Symbiosis with single celled
algae that lives in the jelly-like layer of the eggs has evolved several times. The larvae (tadpoles or polliwogs) breathe with exterior
gills. After hatching, they start to transform gradually into the adult's appearance. This process is called
metamorphosis. Typically, the animals then leave the water and become terrestrial adults, but there are many interesting exceptions to this general way of reproduction.
The most obvious part of the amphibian metamorphosis is the formation of four legs in order to support the body on land. But there are several other changes:
The gills are replaced by other respiratory organs, i.e., lungs.
The skin changes and develops glands to avoid dehydration.
The eyes develop eyelids and adapt to vision outside the water.
An eardrum is developed to lock the middle ear.
In frogs and toads, the tail disappears.
Conservation
Dramatic declines in amphibian populations, including population crashes and mass localized
extinction, have been noted in the past two decades from locations all over the world, and amphibian declines are thus perceived as one of the most critical threats to global
biodiversity. A number of causes are believed to be involved, including
habitat destruction and modification, over-exploitation,
pollution,
introduced species,
climate change, 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 amphibian declines are currently a topic of much ongoing research.
Evolutionary history
The first major groups of amphibians developed in the Devonian Period from fishes similar to the modern coelocanth where the fins had evolved into legs. These amphibians were around five meters long in length, which is rare now except for some species of Japanese Salamander. The land was safe as the giant fishes and sharks in the ocean could not come onto land. However, there were two problems with living out their entire lives on land. Primarily, the food that these amphibians consumed was in the water, but also at this point the skin on most of these amphibians was not water-tight.
In the Carboniferous Period, the amphibians moved up in the food chain and began to occupy the ecological position where we now find crocodiles. These amphibians were notable for eating the mega-insects on land and many types of fishes in the water. Towards the end of the Permian Period and the Triassic Period, the amphibians started having competition with proto-crocodiles which led to their drop in size in the temperate zones or leaving for the poles. (Amphibians were able to hibernate during the winter whereas crocodiles could not, allowing the amphibians in higher latitudes protection from the reptiles.)
The modern
mudskipper provides a rough glimpse into the kind of lifestyle and adaptations that proto-amphibians may have taken.(Mudskippers are not closely related to coelocanths.)
Further reading
{{cite book
| last = Carroll
| first = Robert L.
| title = Vertebrate Paleontology and Evolution
| year = 1988
| publisher = W.H. Freeman & Co.
| location = New York
}}
{{cite book
| last = Duellman
| first = William E.
| coauthors = Linda Trueb
| title = Biology of Amphibians
| year = 1994
| publisher = Johns Hopkins University Press
| isbn = 978-0801847806
}}
{{cite journal
| last = Frost
| first = Darrel R.
| title = The Amphibian Tree of Life
| url = http://digitallibrary.amnh.org/dspace/handle/2246/5781
| journal = Bulletin of the American Museum of Natural History
| volume = 297
| pages = 1-291
| year = 2006
| month = March
| coauthors = Taran Grant, Julián Faivovich, Raoul H. Bain, Alexander Haas, Célio F.B. Haddad, Rafael O. De Sá, Alan Channing, Mark Wilkinson, Stephen C. Donnellan, Christopher J. Raxworthy, Jonathan A. Campbell, Boris L. Blotto, Paul Moler, Robert C. Drewes, Ronald A. Nussbaum, John D. Lynch, David M. Green, Ward C. Wheeler
}}
{{cite journal
| last = Pounds
| first = J. Alan
| title = Widespread amphibian extinctions from epidemic disease driven by global warming
| url = http://www.nature.com/nature/journal/v439/n7073/full/nature04246.html
| journal = Nature
| volume = 439
| pages = 161-167
| year = 2006
| month = January
| doi = 10.1038/nature04246
| coauthors = Martín R. Bustamante, Luis A. Coloma, Jamie A. Consuegra, Michael P. L. Fogden, Pru N. Foster, Enrique La Marca, Karen L. Masters, Andrés Merino-Viteri, Robert Puschendorf, Santiago R. Ron, G. Arturo Sánchez-Azofeifa, Christopher J. Still and Bruce E. Young
}}
{{cite journal
| last = San Mauro
| first = Diego
| coauthors = Miguel Vences, Marina Alcobendas, Rafael Zardoya and Axel Meyer
| title = Initial diversification of living amphibians predated the breakup of Pangaea
| url = http://www.journals.uchicago.edu/AN/journal/issues/v165n5/40546/40546.html
| journal = American Naturalist
| volume = 165
| pages = 590-599
| year = 2005
| month = May
}}
Solomon Berg Martin, ''Biology''
{{cite journal
| last = Stuart
| first = Simon N.
| coauthors = Janice S. Chanson, Neil A. Cox, Bruce E. Young, Ana S. L. Rodrigues, Debra L. Fischman, Robert W. Waller
| title = Status and trends of amphibian declines and extinctions worldwide
| url = http://www.sciencemag.org/cgi/content/full/306/5702/1783
| journal = Science
| volume = 306
| issue = 5702
| pages = 1783-1786
| year = 2004
| month = December
| doi = 10.1126/science.1103538
}}