Frog

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For other uses, see Frog (disambiguation).

Frogs are amphibians in the order Anura (meaning "tail-less", from Greek an-, without + oura,
tail), formerly referred to as Salientia (Latin salere (salio), "to jump"). Most frogs are
characterized by a short body, webbed digits (fingers or toes), protruding eyes and the absence of
a tail. Frogs are widely known as exceptional jumpers, and many of the anatomical
characteristics of frogs, particularly their long, powerful legs, are adaptations to improve
jumping performance. Due to their permeable skin, frogs are often semi-aquatic or inhabit humid
areas, but move easily on land. They typically lay their eggs in puddles, ponds or lakes, and their
larvae, called tadpoles, have gills and develop in water. Adult frogs follow a carnivorous diet,
mostly of arthropods, annelids and gastropods. Frogs are most noticeable by their call, which can
be widely heard during the night or day, mainly in their mating season.

The distribution of frogs ranges from tropic to subarctic regions, but most species are found in
tropical rainforests. Consisting of more than 5,000 species described, they are among the most
diverse groups of vertebrates. However, populations of certain frog species are declining
significantly.

A popular distinction is often made between frogs and toads on the basis of their appearance, but
this has no taxonomic basis. (Members of the anuran family Bufonidae are called true toads, but
many species from other families are also called toads.) In addition to their ecological
importance, frogs have many cultural roles, such as in literature, symbolism and religion, and
they are also valued as food and as pets.

Contents
[hide]

 1 Etymology and terminology

 2 Taxonomy
 3 Morphology and physiology
o 3.1 Feet and legs
o 3.2 Jumping
o 3.3 Skin
o 3.4 Poison
o 3.5 Respiration and circulation
o 3.6 Digestion and excretion
o 3.7 Nervous system
 4 Natural history
 o 4.1 Life cycle
o 4.2 Reproduction of frogs
o 4.3 Parental care
o 4.4 Call
 5 Distribution and conservation status
 6 Evolution
 7 Uses in agriculture and research
 8 Cultural beliefs
 9 See also
 10 References
o 10.1 Notes
o 10.2 Bibliography
 11 External links

Etymology and terminology

The name frog derives from Old English frogga, (compare Old Norse frauki, German Frosch,
older Dutch spelling kikvorsch), cognate with Sanskrit plava (frog), probably deriving from
Proto-Indo-European praw = "to jump".[1]

A distinction is often made between frogs and toads on the basis of their appearance, caused by
the convergent adaptation among so-called toads to dry environments; however, this distinction
has no taxonomic basis. The only family exclusively given the common name "toad" is
Bufonidae, but many species from other families are also called "toads," and the species within
the toad genus Atelopus are referred to as "harlequin frogs".

Taxonomy
For more details on this topic, see List of Anuran families.

The order Anura contains 4,810 species[2] in 33 families, of which the Leptodactylidae (1100
spp.), Hylidae (800 spp.) and Ranidae (750 spp.) are the richest in species. About 88% of
amphibian species are frogs.
European Fire-bellied Toad (Bombina bombina)

Young American bullfrog found in a stream in New Jersey

The use of the common names "frog" and "toad" has no taxonomic justification. From a
taxonomic perspective, all members of the order Anura are frogs, but only members of the
family Bufonidae are considered "true toads". The use of the term "frog" in common names
usually refers to species that are aquatic or semi-aquatic with smooth and/or moist skins, and the
term "toad" generally refers to species that tend to be terrestrial with dry, warty skin. An
exception is the fire-bellied toad (Bombina bombina): while its skin is slightly warty, it prefers a
watery habitat.

Frogs and toads are broadly classified into three suborders: Archaeobatrachia, which includes
four families of primitive frogs; Mesobatrachia, which includes five families of more
evolutionary intermediate frogs; and Neobatrachia, by far the largest group, which contains the
remaining 24 families of "modern" frogs, including most common species throughout the world.
Neobatrachia is further divided into the Hyloidea and Ranoidea.[3] This classification is based on
such morphological features as the number of vertebrae, the structure of the pectoral girdle, and
the morphology of tadpoles. While this classification is largely accepted, relationships among
families of frogs are still debated. Future studies of molecular genetics should soon provide
further insights to the evolutionary relationships among anuran families.[4]

Some species of anurans hybridise readily. For instance, the Edible Frog (Rana esculenta) is a
hybrid of the Pool Frog (R. lessonae) and the Marsh Frog (R. ridibunda). Bombina bombina and
Bombina variegata similarly form hybrids, although these are less fertile, giving rise to a hybrid
zone.

Morphology and physiology

Skeleton of Rana

The morphology of frogs is unique among amphibians. Compared with the other two groups of
amphibians, (salamanders and caecilians), frogs are unusual because they lack tails as adults and
their legs are more suited to jumping than walking. The physiology of frogs is generally like that
of other amphibians (and differs from other terrestrial vertebrates) because oxygen can pass
through their highly permeable skin. This unique feature allows frogs to "breathe" largely
through their skin.[citation needed] Because the oxygen is dissolved in an aqueous film on the skin and
passes from there to the blood, the skin must remain moist at all times; this makes frogs
susceptible to many toxins in the environment, some of which can similarly dissolve in the layer
of water and be passed into their bloodstream. This may be the cause of the decline in frog
populations.[5][6][7][8][9][10][11][12][13]

Many characteristics are not shared by all of the approximately 5,250 described frog species.
However, some general characteristics distinguish them from other amphibians. Frogs are
usually well suited to jumping, with long hind legs and elongated ankle bones. They have a short
vertebral column, with no more than ten free vertebrae, followed by a fused tailbone (urostyle or
coccyx), typically resulting in a tailless phenotype.[citation needed]

Frogs range in size from 10 mm (0.39 in) (Brachycephalus didactylus of Brazil and
Eleutherodactylus iberia of Cuba) to 300 mm (12 in) (goliath frog, Conraua goliath, of
Cameroon). The skin hangs loosely on the body because of the lack of loose connective tissue.
Skin texture varies: it can be smooth, warty or folded. Frogs have three eyelid membranes: one is
transparent to protect the eyes underwater, and two vary from translucent to opaque. Frogs have
a tympanum on each side of the head, which is involved in hearing and, in some species, is
covered by skin. Most frogs have teeth, specifically pedicellate teeth in which the crow is
separated from the root by fibrous tissue. Most only have teeth on the edge of the upper jaw
(maxillary teeth) as well as vomerine teeth on the roof of their mouth. They do not have any teeth
on their lower jaw, so they usually swallow their food whole. The teeth are mainly used to hold
the prey and keep it in place till they can get a good grip on it and swallow their meal, assisted by
retracting their eyes into their head.[14] True toads lack any teeth at all, and some species
(Pyxicephalus) which prey on relatively large organisms (including mice and other frogs) have
cone shaped projections of bone, called odontoid processes, at the front of the lower jaw which
function like teeth.[2]

Feet and legs

Tyler's Tree Frog (Litoria tyleri) illustrates large toe pads and webbed feet

A bullfrog skeleton, showing elongate limb bones and extra joints. Red marks indicate bones
which have been substantially elongated in frogs and joints which have become mobile. Blue
indicates joints and bones which have not been modified or only somewhat elongated.

The structure of the feet and legs varies greatly among frog species, depending in part on
whether they live primarily on the ground, in water, in trees, or in burrows. Frogs must be able to
move quickly through their environment to catch prey and escape predators, and numerous
adaptations help them do so.

Many frogs, especially those that live in water, have webbed toes. The degree to which the toes
are webbed is directly proportional to the amount of time the species lives in the water. For
example, the completely aquatic African dwarf frog (Hymenochirus sp.) has fully webbed toes,
whereas the toes of White's tree frog (Litoria caerulea), an arboreal species, are only a half or a
quarter webbed.

Arboreal frogs have "toe pads" to help grip vertical surfaces. These pads, located on the ends of
the toes, do not work by suction. Rather, the surface of the pad consists of interlocking cells,
with a small gap between adjacent cells. When the frog applies pressure to the toe pads, the
interlocking cells grip irregularities on the substrate. The small gaps between the cells drain
away all but a thin layer of moisture on the pad, and maintain a grip through capillarity. This
allows the frog to grip smooth surfaces, and does not function when the pads are excessively wet.
[15]

In many arboreal frogs, a small "intercalary structure" in each toe increases the surface area
touching the substrate. Furthermore, since hopping through trees can be dangerous, many
arboreal frogs have hip joints that allow both hopping and walking. Some frogs that live high in
trees even possess an elaborate degree of webbing between their toes, as do aquatic frogs. In
these arboreal frogs, the webs allow the frogs to "parachute" or control their glide from one
position in the canopy to another.[16]

Ground-dwelling frogs generally lack the adaptations of aquatic and arboreal frogs. Most have
smaller toe pads, if any, and little webbing. Some burrowing frogs have a toe extension—a
metatarsal tubercle—that helps them to burrow. The hind legs of ground dwellers are more
muscular than those of aqueous and tree-dwelling frogs.

Sometimes during the tadpole stage, one of the animal's rear leg stubs is eaten by a dragonfly
nymph. In some of these cases, the full leg grows anyway, and in other cases, it does not,
although the frog may still live out its normal lifespan with only three legs. Other times, a
parasitic flatworm called Riberoria trematodes digs into the rear of a tadpole, where it rearranges
the limb bud cells, which sometimes causes the frog to have extra legs.[17]

Jumping

Rainforest Rocket Frog jumping

Frogs are generally recognized as exceptional jumpers, and the best jumper of all vertebrates.
The Australian rocket frog, Litoria nasuta, can leap over 50 times its body length (5.5 cm),
resulting in jumps of over 2 meters. The acceleration of the jump may be up to twice gravity.
There are tremendous differences between species in jumping capability, but within a species,
jump distance increases with increasing size, but relative jumping distance (body-lengths
jumped) decreases.

While frog species can use a variety of locomotor modes (running, walking, gliding, swimming,
and climbing), more are either proficient at jumping or descended from ancestors who were, with
much of the musculo-skeletal morphology modified for this purpose. The tibia, fibula and tarsals
have been fused into a single, strong bone, as have the radius and ulna in the forelimbs (which
must absorb the impact of landing). The metatarsals have become elongated to add to the leg
length and allow the frog to push against the ground for longer during a jump. The illium has
elongated and formed a mobile joint with the sacrum which, in specialist jumpers such as Ranids
or Hylids, functions as an additional limb joint to further power the leaps. This elongation of the
limbs results in the frog being able to apply force to the ground for longer during a jump, which
in turn results in a longer, faster jump.[citation needed]

The muscular system has been similarly modified. The hind limbs of the ancestor of frogs
presumably contained pairs of muscles which would act in opposition (one muscle to flex the
knee, a different muscle to extend it), as is seen in most other limbed animals. However, in
modern frogs, almost all muscles have been modified to contribute to the action of jumping, with
only a few small muscles remaining to bring the limb back to the starting position and maintain
posture. The muscles have also been greatly enlarged, with the muscles involved in jumping
accounting for over 17% of the total mass of the frog.

In some extremely capable jumpers, such as the cuban tree frog, the peak power exerted during a
jump can exceed what muscle is capable of producing. Currently, it is hypothesized that frogs are
storing muscular energy by stretching their tendons like springs, then triggering the release all at
once, allowing the frog to increase the energy of its jump beyond the limits of muscle-powered
acceleration. A similar mechanism has already been documented in locusts and grasshoppers.[18]

Skin

Pouched Frog (Assa darlingtoni) camouflaged against leaf litter.

Microscopic view of frog skin

Many frogs are able to absorb water and oxygen directly through the skin, especially around the
pelvic area. However, the permeability of a frog's skin can also result in water loss. Some tree
frogs reduce water loss with a waterproof layer of skin. Others have adapted behaviours to
conserve water, including engaging in nocturnal activity and resting in a water-conserving
position. This position involves the frog lying with its toes and fingers tucked under its body and
chin, respectively, with no gap between the body and substrate. Some frog species will also rest
in large groups, touching the skin of the neighbouring frog. This reduces the amount of skin
exposed to the air or a dry surface, and thus reduces water loss. These adaptations only reduce
water loss enough for a predominantly arboreal existence, and are not suitable for arid
conditions.

Camouflage is a common defensive mechanism in frogs. Most camouflaged frogs are nocturnal,
which adds to their ability to hide. Nocturnal frogs usually find the ideal camouflaged position
during the day to sleep. Some frogs have the ability to change colour, but this is usually
restricted to shades of one or two colours. For example, White's tree frog varies in shades of
green and brown. Features such as warts and skin folds are usually found on ground-dwelling
frogs, where a smooth skin would not disguise them effectively. Arboreal frogs usually have
smooth skin, enabling them to disguise themselves as leaves.[citation needed]

Certain frogs change colour between night and day, as light and moisture stimulate the pigment
cells and cause them to expand or contract.

Poison

Many frogs contain mild toxins that make them unpalatable to potential predators. For example,
all toads have large poison glands—the parotoid glands—located behind the eyes, on the top of
the head. Some frogs, such as some poison dart frogs, are especially toxic. The chemical makeup
of toxins in frogs varies from irritants to hallucinogens, convulsants, nerve poisons, and
vasoconstrictors. Many predators of frogs have adapted to tolerate high levels of these poisons.
Others, including humans, may be severely affected.

Oophaga pumilio, a poison dart frog, contains numerous alkaloids which deter predators

Some frogs obtain poisons from the ants and other arthropods they eat;[19] others, such as the
Australian Corroboree Frogs (Pseudophryne corroboree and Pseudophryne pengilleyi), can
manufacture an alkaloid not derived from their diet.[20] Some native people of South America
extract poison from the poison dart frogs and apply it to their darts for hunting,[21] although few
species are toxic enough to be used for this purpose. It was previously a misconception the
poison was placed on arrows rather than darts. The common name of these frogs was thus
changed from "poison arrow frog" to "poison dart frog" in the early 1980s. Poisonous frogs tend
to advertise their toxicity with bright colours, an adaptive strategy known as aposematism. There
are at least two non-poisonous species of frogs in tropical America (Eleutherodactylus gaigei
and Lithodytes lineatus) that mimic the colouration of dart poison frogs' coloration for self-
protection (Batesian mimicry).[22][23]

Because frog toxins are extraordinarily diverse, they have raised the interest of biochemists as a
"natural pharmacy". The alkaloid epibatidine, a painkiller 200 times more potent than morphine,
is found in some species of poison dart frogs. Other chemicals isolated from the skin of frogs
may offer resistance to HIV infection.[24] Arrow and dart poisons are under active investigation
for their potential as therapeutic drugs.[25]

The skin secretions of some toads, such as the Colorado River toad and cane toad, contain
bufotoxins, some of which, such as bufotenin, are psychoactive, and have therefore been used as
recreational drugs. Typically, the skin secretions are dried and smoked. Skin licking is especially
dangerous, and appears to constitute an urban myth. See psychoactive toad.

Respiration and circulation

The skin of a frog is permeable to oxygen and carbon dioxide, as well as to water. There are a
number of blood vessels near the surface of the skin. When a frog is underwater, oxygen is
transmitted through the skin directly into the bloodstream. On land, adult frogs use their lungs to
breathe. Their lungs are similar to those of humans, but the chest muscles are not involved in
respiration, and there are no ribs or diaphragm to support breathing. Frogs breathe by taking air
in through the nostrils (which often have valves which close when the frog is submerged),
causing the throat to puff out, then compressing the floor of the mouth, which forces the air into
the lungs. In August 2007 an aquatic frog named Barbourula kalimantanensis was discovered in
a remote part of Indonesia. The Bornean Flat-headed Frog (B. kalimantanensis) is the first
species of frog known to science without lungs.

Frogs are known for their three-chambered heart, which they share with all tetrapods except
birds, crocodilians and mammals. In the three-chambered heart, oxygenated blood from the lungs
and de-oxygenated blood from the respiring tissues enter by separate atria, and are directed via a
spiral valve to the appropriate vessel—aorta for oxygenated blood and pulmonary artery for
deoxygenated blood. This special structure is essential to keeping the mixing of the two types of
blood to a minimum, which enables frogs to have higher metabolic rates, and to be more active
than otherwise.

Some species of frog have remarkable adaptations that allow them to survive in oxygen deficient
water. The lake titicaca frog (Telmatobius culeus) is one such species and to survive in the
poorly oxygenated waters of Lake Titicaca it has incredibly wrinkly skin that increases its
surface area to enhance gas exchange. This frog will also do 'push-ups' on the lake bed to
increase the flow of water around its body.[26]
Digestion and excretion

The frog's digestive system begins with the mouth. Frogs have teeth along their upper jaw called
the maxillary teeth, which are used to grind food before swallowing. These teeth are very weak,
and cannot be used to catch or harm agile prey. Instead, the frog uses its sticky tongue to catch
food (such as flies or other insects). The food then moves through the esophagus into the
stomach. The food then proceeds to the small intestine (duodenum and ileum) where most
digestion occurs. Frogs carry pancreatic juice from the pancreas, and bile (produced by the liver)
through the gallbladder from the liver to the small intestine, where the fluids digest the food and
extract the nutrients. When the food passes into the large intestine, the water is reabsorbed and
wastes are routed to the cloaca. All wastes exit the body through the cloaca and the cloacal vent.

Nervous system

The frog has a highly developed nervous system which consists of a brain, spinal cord and
nerves. Many parts of the frog's brain correspond with those of humans. The medulla oblongata
regulates respiration, digestion, and other automatic functions. Muscular coordination and
posture are controlled by the cerebellum. The relative size of the cerebrum of a frog is much
smaller than that of a human. Frogs have ten cranial nerves (nerves which pass information from
the outside directly to the brain) and ten pairs of spinal nerves (nerves which pass information
from extremities to the brain through the spinal cord). By contrast, all amniotes (mammals, birds
and reptiles) have twelve cranial nerves. Frogs do not have external ears; the eardrums (tympanic
membranes) are directly exposed. As in all animals, the ear contains semicircular canals which
help control balance and orientation. Due to their short cochlea, frogs use electrical tuning to
expand their range of audible frequencies.

Natural history
The life cycle of frogs, like that of other amphibians, consists of four main stages: egg, tadpole,
metamorphosis and adult. The reliance of frogs on an aquatic environment for the egg and
tadpole stages gives rise to a variety of breeding behaviours that include the well-known mating
calls used by the males of most species to attract females to the bodies of water that they have
chosen for breeding. Some frogs also look after their eggs—and in some cases even the tadpoles
—for some time after laying.

Life cycle
Frogspawn

Frogspawn development

Tadpole of Haswell's Froglet (Paracrinia haswelli

The life cycle of a frog starts with an egg. A female generally lays gelatinous egg masses
containing thousands of eggs, in water. Each anuran species lays eggs in a distinctive,
identifiable manner. An example are the long strings of eggs laid by the common American toad.
The eggs are highly vulnerable to predation, so frogs have evolved many techniques to ensure
the survival of the next generation. In colder areas the embryo is black to absorb more heat from
the sun, which speeds up the development. Most commonly, this involves synchronous
reproduction. Many individuals will breed at the same time, overwhelming the actions of
predators; the majority of the offspring will still die due to predation, but there is a greater
chance some will survive. Another way in which some species avoid the predators and pathogens
eggs are exposed to in ponds is to lay eggs on leaves above the pond, with a gelatinous coating
designed to retain moisture. In these species the tadpoles drop into the water upon hatching. The
eggs of some species laid out of water can detect vibrations of nearby predatory wasps or snakes,
and will hatch early to avoid being eaten.[27] Some species, such as the Cane Toad (Bufo
marinus), lay poisonous eggs to minimise predation. While the length of the egg stage depends
on the species and environmental conditions, aquatic eggs generally hatch within one week.
Other species go through their whole larval phase inside the eggs or the mother, or they have
direct development. Unlike salamanders and newts, frogs and toads never become sexually
mature while still in their larval stage.

Eggs hatch and continue life as tadpoles (occasionally known as polliwogs), which typically
have oval bodies and long, vertically flattened tails. At least one species (Nannophrys
ceylonensis) has tadpoles that are semi-terrestrial and live among wet rocks,[28][29] but as a general
rule, free living larvae are fully aquatic. They lack eyelids and have a cartilaginous skeleton, a
lateral line system, gills for respiration (external gills at first, internal gills later) and tails with
dorsal and ventral folds of skin for swimming.[30] From pretty early onward they develop a gill
pouch that covers the gills and the front legs and also the lungs are developed in an early stage as
an accessory breathing organ. Some species which go through the metamorphosis inside the egg
and hatch to small frogs never develop gills, instead there are specialised areas of skin that takes
care of the respiration. Tadpoles also lack true teeth, but the jaws in most species usually have
two elongate, parallel rows of small keratinized structures called keradonts in the upper jaw
while the lower jaw has three rows of keradonts, surrounded by a horny beak, but the number of
rows can be lower or absent, or much higher.[31] Tadpoles are typically herbivorous, feeding
mostly on algae, including diatoms filtered from the water through the gills. Some species are
carnivorous at the tadpole stage, eating insects, smaller tadpoles, and fish. Cannibalism has been
observed among tadpoles. Early developers who gain legs may be eaten by the others, so the late
bloomers survive longer. This has been observed in England in the species Rana temporaria
(common frog).[32]

Tadpoles are highly vulnerable to predation by fish, newts, predatory diving beetles and birds
such as kingfishers. Poisonous tadpoles are present in many species, such as Cane Toads. The
tadpole stage may be as short as a week, or tadpoles may overwinter and metamorphose the
following year in some species, such as the midwife toad (Alytes obstetricans) and the common
spadefoot (Pelobates fuscus). In the Pipidae, with the exception for Hymenochirus, the tadpoles
have paired anterior barbels which make them resemble small catfish.[33]

With the exception of the base of the tail, where a few vertebral structures develop to give rise to
the urostyle later in life, the tail lacks the completely solid, segmental, skeletal elements of
cartilage or bony tissue that are so typical for other vertebrates, although it does contain a
notochord

At the end of the tadpole stage, frogs undergo metamorphosis, in which they undergo a transition
into the adult form. This [metamorphosis]] last typically only 24 hours and consists of:
Larva of the common frog Rana temporaria a day before metamorphisis

Common frog - Metamorphosis stage. Notice the deformed jaws, large eyes and the remains of
the gill pouch.

Young frog with tail remains after metamorphosis

Adult leopard frog

 The disappearance of the gill pouch, making the front legs visible.
 The transformation of the jaws into the big jaws of predatory frogs (most tadpoles are
scraping of algae or are filter feeders)
 The transformation of the digestive system: the long spiral gut of the larva is being
replaced by the typical short gut of a predator.
 An adaptation of the nervous system for stereoscopic vision, locomotion and feeding
 A quick growth and movement of the eyes to higher up the skull and the formation of
eyelids.
 Formation of skin glands, thickening of the skin and loss of the lateral line system
  An eardrum is developed to lock the middle ear.

The disappearance of the tail is somewhat later (occurs at higher thyroxin levels) and after the
tail has been resorbed the animals are ready to leave the water. The material of the tail is being
used for a quick growth of the legs. The disappearing of the larval structures is a regulated
process called apoptosis. tail.

Incident of frog cannibalism

After metamorphosis, young adults may leave the water and disperse into terrestrial habitats, or
continue to live in the aquatic habitat as adults. Almost all species of frogs are carnivorous as
adults, eating invertebrates such as arthropods, annelids and gastropods. A few of the larger
species may eat prey such as small mammals, fish and smaller frogs. Some frogs use their sticky
tongues to catch fast-moving prey, while others capture their prey and force it into their mouths
with their hands. However, there are a very few species of frogs that primarily eat plants.[34]
Adult frogs are themselves preyed upon by birds, large fish, snakes, otters, foxes, badgers, coatis,
and other animals. Frogs are also eaten by people (see section on uses in agriculture and
research, below).

Frogs and toads can live for many years; though little is known about their life span in the wild,
captive frogs and toads are recorded living up to 40 years.[35]

Frogs from temperate climates hibernate through the winter, and 4 species are known to freeze
during this time, most notably Rana sylvatica.[36]

Reproduction of frogs

Once adult frogs reach maturity, they will assemble at a water source such as a pond or stream to
breed. Many frogs return to the bodies of water where they were born, often resulting in annual
migrations involving thousands of frogs. In continental Europe, a large proportion of migrating
frogs used to die on roads, before special fences and tunnels were built for them.
Male and female Common toad (Bufo bufo) in amplexus

A Male and Female common toad in amplexus. The black strands are eggs released into open
water minutes after birth.

Once at the breeding ground, male frogs call to attract a mate, collectively becoming a chorus of
frogs. The call is unique to the species, and will attract females of that species. Some species
have satellite males who do not call, but intercept females that are approaching a calling male.

The male and female frogs then undergo amplexus. This involves the male mounting the female
and gripping her (sometimes with special nuptial pads) tightly. Fertilization is external: the egg
and sperm meet outside of the body. The female releases her eggs, which the male frog covers
with a sperm solution. The eggs then swell and develop a protective coating. The eggs are
typically brown or black, with a clear, gelatin-like covering.

Most temperate species of frogs reproduce between late autumn and early spring. In the UK,
most common frog populations produce frogspawn in February, although there is wide variation
in timing. Water temperatures at this time of year are relatively low, typically between four and
10 degrees Celsius. Reproducing in these conditions helps the developing tadpoles because
dissolved oxygen concentrations in the water are highest at cold temperatures. More importantly,
reproducing early in the season ensures that appropriate food is available to the developing frogs
at the right time.

Parental care
Colour plate from Ernst Haeckel's 1904 Kunstformen der Natur, depicting frog species that
include two examples of parental care.

Although care of offspring is poorly understood in frogs, it is estimated that up to 20% of

amphibian species may care for their young in one way or another, and there is a great diversity
of parental behaviours.[37] Some species of poison dart frog lay eggs on the forest floor and
protect them, guarding the eggs from predation and keeping them moist. The frog will urinate on
them if they become too dry. After hatching, a parent (the sex depends upon the species) will
move them, on its back, to a water-holding bromeliad. The parent then feeds them by laying
unfertilized eggs in the bromeliad until the young have metamorphosed. Other frogs carry the
eggs and tadpoles on their hind legs or back (e.g. the midwife toads, Alytes spp.). Some frogs
even protect their offspring inside their own bodies. The male Australian Pouched Frog (Assa
darlingtoni) has pouches along its side in which the tadpoles reside until metamorphosis. The
female Gastric-brooding Frogs (genus Rheobatrachus) from Australia, now probably extinct,
swallows its tadpoles, which then develop in the stomach. To do this, the Gastric-brooding Frog
must stop secreting stomach acid and suppress peristalsis (contractions of the stomach). Darwin's
Frog (Rhinoderma darwinii) from Chile puts the tadpoles in its vocal sac (see next section) for
development. Some species of frog will leave a 'babysitter' to watch over the frogspawn until it
hatches.

The evolution of parental care in frogs is driven primarily by the size of the water body in which
they breed. There is an inverse relationship between the level of parental care in a frog species
and the degree of parental care they exhibit—frogs that breed in smaller water bodies tend to
have more complex parental care behaviors.[38] Water body size shows this strong relationship
with parental care because it encompasses several important variables that interact to select for
parental care: predation, desiccation, competition, and resource limitation. Because predation of
eggs and larvae is high in large water bodies, a number of frog species evolved terrestrial
oviposition. Once eggs are deposited on land, the desiccating terrestrial environment demands
uniparental care in the form of egg hydration to ensure egg survival.[39] The subsequent need to
transport hatched tadpoles to a water source requires an even more intense form of uniparental
care. In small water bodies where predators are mostly absent, such as phytotelmata (water-filled
leaf axils or small woody cavities), inter-tadpole competition becomes the variable that
constrains tadpole survival. Certain frogs species avoid this competition by evolving the use of
smaller phytotelmata as tadpole deposition sites.[40] However, while these smaller tadpole rearing
sites are free of competition, they also lack nutrients. Because they do not have sufficient
nutrients to support a tadpole without parental provisioning behavior, frog species that
transitioned from the use of larger to smaller phytotelmata have evolved trophic (unfertilized)
egg laying. In this complex form of biparental care, the female provides her offspring with
nutritive eggs. While each of these variables select for different behaviors, they correlate with the
size of a species' tadpole-rearing site and influence the degree of parental care displayed by a
species.

Call

A male Dendropsophus microcephalus displaying its vocal sac during its call.

Some frog calls are so loud, they can be heard up to a mile away.[41] The call of a frog is unique
to its species. Frogs call by passing air through the larynx in the throat. In most calling frogs, the
sound is amplified by one or more vocal sacs, membranes of skin under the throat or on the
corner of the mouth that distend during the amplification of the call. The field of neuroethology
studies the neurocircuitry that underlies frog audition.

Some frogs lack vocal sacs, such as those from the genera Heleioporus and Neobatrachus, but
these species can still produce a loud call. Their buccal cavity is enlarged and dome-shaped,
acting as a resonance chamber that amplifies their call. Species of frog without vocal sacs and
that do not have a loud call tend to inhabit areas close to flowing water. The noise of flowing
water overpowers any call, so they must communicate by other means.

The main reason for calling is to allow males to attract a mate. Males call either individually or
in a group called a chorus. Females of many frog species, for example Polypedates leucomystax,
produce calls reciprocal to the males', which act as the catalyst for the enhancement of
reproductive activity in a breeding colony.[42] A male frog emits a release call when mounted by
another male. Tropical species also have a rain call that they make on the basis of humidity cues
prior to a rain shower. Many species also have a territorial call that is used to chase away other
males. All of these calls are emitted with the mouth of the frog closed.
A distress call, emitted by some frogs when they are in danger, is produced with the mouth open,
resulting in a higher-pitched call. The effectiveness of the call is unknown; however, it is
suspected the call intrigues the predator until another animal is attracted, distracting them enough
for its escape.

Many species of frog have deep calls, or croaks. The English onomatopoeic spelling is "ribbit".
The croak of the American bullfrog (Rana catesbiana) is sometimes spelt "jug o' rum".[43] Other
examples are Ancient Greek brekekekex koax koax for probably Rana ridibunda, and the
description in Rigveda 7:103.6 gómāyur éko ajámāyur ékaħ = "one has a voice like a cow's, one
has a voice like a goat's".

Distribution and conservation status

The Red-eyed Tree Frog (Litoria chloris) is a species of tree frog native to eastern Australia.
See also: Decline in amphibian populations

The habitat of frogs extends almost worldwide, but they do not occur in Antarctica and are not
present on many oceanic islands.[44][45] The greatest diversity of frogs occurs in the tropical areas
of the world, where water is readily available, suiting frogs' requirements due to their skin. Some
frogs inhabit arid areas such as deserts, where water may not be easily accessible, and rely on
specific adaptations to survive. The Australian genus Cyclorana and the American genus
Pternohyla will bury themselves underground, create a water-impervious cocoon and hibernate
during dry periods. Once it rains, they emerge, find a temporary pond and breed. Egg and tadpole
development is very fast in comparison to most other frogs so that breeding is complete before
the pond dries up. Some frog species are adapted to a cold environment; for instance the wood
frog, whose habitat extends north of the Arctic Circle, buries itself in the ground during winter
when much of its body freezes.
Golden toad (Bufo periglenes) – last seen in 1989

Frog populations have declined dramatically since the 1950s: more than one third of species are
believed to be threatened with extinction and more than 120 species are suspected to be extinct
since the 1980s.[46] Among these species are the golden toad of Costa Rica and the Gastric-
brooding frogs of Australia. Habitat loss is a significant cause of frog population decline, as are
pollutants, climate change, the introduction of non-indigenous predators/competitors, and
emerging infectious diseases including chytridiomycosis. Many environmental scientists believe
that amphibians, including frogs, are excellent biological indicators of broader ecosystem health
because of their intermediate position in food webs, permeable skins, and typically biphasic life
(aquatic larvae and terrestrial adults).[47] It appears that it is the species with both aquatic eggs
and aquatic larvae that are most affected by the decline, while those with direct development are
the most resistant.[48]

A Canadian study conducted in 2006, suggested heavy traffic near frog habitats as a large threat
to frog populations.[49] In a few cases, captive breeding programs have been attempted to
alleviate the pressure on frog populations, and these have proved successful.[50][51] In 2007, it was
reported the application of certain probiotic bacteria could protect amphibians from
chytridiomycosis.[52] One current project, The Panama Amphibian Rescue and Conservation
Project, has subsequently been developed in order to rescue species at risk of chytridiomycosis in
eastern Panama, and to develop field applications of this probiotic cure.[53]

Zoos and aquariums around the world named 2008 the Year of the Frog, to draw attention to the
conservation issues.[54]

Evolution
A fossilized frog from the Czech Republic, possibly Palaeobatrachus gigas

Until the discovery of the Early Permian Gerobatrachus hottoni in 2008, a stem-batrachian with
many salamander-like characteristics, the earliest known proto-frog was Triadobatrachus
massinoti, from the 250 million year old early Triassic of Madagascar.[55] The skull is frog-like,
being broad with large eye sockets, but the fossil has features diverging from modern amphibia.
These include a different ilium, a longer body with more vertebrae, and separate vertebrae in its
tail (whereas in modern frogs, the tail vertebrae are fused, and known as the urostyle or coccyx).
The tibia and fibula bones are unfused and separate, making it probable Triadobatrachus was not
an efficient leaper.

Another fossil frog, Prosalirus bitis, was discovered in 1995. The remains were recovered from
Arizona's Kayenta Formation, which dates back to the Early Jurassic epoch,[56] somewhat
younger than Triadobatrachus. Like Triadobatrachus, Prosalirus did not have greatly enlarged
legs, but had the typical three-pronged pelvic structure. Unlike Triadobatrachus, Prosalirus had
already lost nearly all of its tail[citation needed] and was well adapted for jumping.[57]

The earliest known "true frog" is Vieraella herbsti, from the early Jurassic (188–213 million
years ago). It is known only from the dorsal and ventral impressions of a single animal and was
estimated to be 33 mm (1.3 in) from snout to vent. Notobatrachus degiustoi from the middle
Jurassic is slightly younger, about 155–170 million years old. It is likely the evolution of modern
Anura was completed by the Jurassic period. The main evolutionary changes involved the
shortening of the body and the loss of the tail.

The earliest full fossil record of a modern frog is of sanyanlichan, which lived 125 million years
ago[58] and had all modern frog features, but bore 9 presacral vertebrae instead of the 8 of modern
frogs.[59]

Frog fossils have been found on all continents except Antarctica, but biogeographic evidence
suggests they inhabited Antarctica when it was warmer.[60]

Uses in agriculture and research

For more details on this topic, see Animal testing on frogs.

Frogs are raised commercially for several purposes. Frogs are used as a food source; frog legs
are a delicacy in China, France, the Philippines, the north of Greece and in many parts of the
American South, especially Louisiana. Dead frogs are sometimes used for dissections in high
school and university anatomy classes, often after being injected with coloured plastics to
enhance the contrast between the organs. This practice has declined in recent years with the
increasing concerns about animal welfare.

Frogs have served as important model organisms throughout the history of science. Eighteenth-
century biologist Luigi Galvani discovered the link between electricity and the nervous system
through studying frogs. The African clawed frog or platanna (Xenopus laevis) was first widely
used in laboratories in pregnancy assays in the first half of the 20th century. When human
chorionic gonadotropin, a hormone found in substantial quantities in the urine of pregnant
women, is injected into a female X. laevis, it induces them to lay eggs. In 1952, Robert Briggs
and Thomas J. King cloned a frog by somatic cell nuclear transfer, the same technique later used
to create Dolly the Sheep, their experiment was the first time successful nuclear transplantation
had been accomplished in metazoans.[61]

Frogs are used in cloning research and other branches of embryology because frogs are among
the closest living relatives of man to lack egg shells characteristic of most other vertebrates, and
therefore facilitate observations of early development. Although alternative pregnancy assays
have been developed, biologists continue to use Xenopus as a model organism in developmental
biology because it is easy to raise in captivity and has a large and easily manipulatable embryo.
Recently, X. laevis is increasingly being displaced by its smaller relative X. tropicalis, which
reaches its reproductive age in five months rather than one to two years (as in X. laevis),[62]
facilitating faster studies across generations. The genome sequence of X. tropicalis will probably
be completed by 2015 at the latest.[63]

Cultural beliefs
For more details on this topic, see Frogs in popular culture.

Moche Frog 200 A.D. Larco Museum Collection Lima, Peru

Frogs feature prominently in folklore, fairy tales and popular culture. They tend to be portrayed
as benign, ugly, clumsy, but with hidden talents. Examples include Michigan J. Frog, The Frog
Prince, and Kermit the Frog. Michigan J. Frog, featured in the Warner Brothers cartoon One
Froggy Evening, only performs his singing and dancing routine for his owner. Once another
person looks at him, he will return to a frog-like pose. "The Frog Prince" is a fairy tale of a frog
who turns into a handsome prince once kissed. Kermit the Frog, on the other hand, is a
conscientious and disciplined character of The Muppet Show and Sesame Street; while openly
friendly and greatly talented, he is often portrayed as cringing at the fanciful behavior of more
flamboyant characters.

The Moche people of ancient Peru worshipped animals and often depicted frogs in their art.[64] In
Panama local legend promised luck to anyone who spotted a golden frog in the wild and some
believed that when Panamanian Golden Frogs died, they would turn into a gold talisman, known
as a huaca. Today, despite being extinct in the wild, Panamanian Golden Frogs remain an
important cultural symbol and can be found on decorative cloth molas made by the Kuna
Indians, on T-shirts, as inlaid design on a new overpass in Panama City and even on lottery
tickets.[65]

See also
 See-through frog
 Stannius ligature

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 Estes, R., and O. A. Reig. (1973). "The early fossil record of frogs: a review of the
evidence." pp. 11–63 In J. L. Vial (Ed.), Evolutionary Biology of the Anurans:
Contemporary Research on Major Problems. University of Missouri Press, Columbia.
 Gissi, Carmela; Diego San Mauro, Graziano Pesole and Rafael Zardoya (February 2006).
"Mitochondrial phylogeny of Anura (Amphibia): A case study of congruent phylogenetic
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doi:10.1016/j.gene.2005.07.034. PMID 16307849.
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(May 2005). "Initial diversification of living amphibians predated the breakup of
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External links

Wikimedia Commons has media related to: Frog

 Wikibooks Cookbook has a recipe/module on
Frog
The Wikibook School Science has a page on the topic of
Frog dissection

Wikiquote has a collection of quotations related to: Frogs

Wikispecies has information related to: Anura

 AmphibiaWeb
 The Whole Frog Project - Virtual frog dissection and anatomy
 Disappearance of toads, frogs has some scientists worried San Francisco Chronicle, 20
April 1992
 Recording UK frogspawn sightings - Springwatch 2006
 Amphibian photo gallery by scientific name - Features many unusual frogs
 Scientific American: Researchers Pinpoint Source of Poison Frogs' Deadly Defenses
 Frogwatch USA - Volunteer frog and toad monitoring program by National Wildlife
Federation and USGS, includes links to frog calls of the United States

Media

 Time-lapse video showing the egg's development until hatching

 Frog calls - Short video clips of calling frogs and interviews with scientists about frog
issues
 Frog vocalisations from around the world - From the British Library Sound Archive
 Frog calls - From Manitoba, Canada
 Frog calls - From eastern and central North America