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(Redirected from Frogs[1])
For other uses, see Frog (disambiguation)[2].
Frogs are a diverse and largely carnivorous[3] group of short-bodied, tailless amphibians[4] composing the order[5] Anura
(Ancient Greek[6] an-, without + oura, tail). The oldest fossil "proto-frog"[7] appeared in the early Triassic[8] of Madagascar[9],
but molecular clock dating[10] suggests their origins may extend further back to the Permian[11], 265 million years ago. Frogs
are widely distributed, ranging from the tropics[12] to subarctic[13] regions, but the greatest concentration of species
diversity[14] is found in tropical rainforests[15]. There are approximately 4,800 recorded species, accounting for over 85% of
extant amphibian species. They are also one of the five most diverse vertebrate[16] orders.
The body plan[17] of an adult frog is generally characterized by a stout body, protruding eyes[18], cleft tongue[19], limbs folded
underneath and the absence of a tail[20] in adults. Besides living in fresh water and on dry land, the adults of some species are
adapted for living underground or in trees. The skin of the frog is glandular[21], with secretions ranging from distasteful to
toxic. Warty species of frog tend to be called toads[22] but the distinction between frogs and toads is based on informal naming
conventions concentrating on the warts rather than taxonomy[23] or evolutionary history; some toads are more closely related
to frogs than to other toads. Frogs' skins vary in colour from well-camouflaged[24] dappled brown, grey and green to vivid
patterns of bright red or yellow and black to advertise toxicity and warn off predators[25].
Frogs typically lay their eggs[26] in water. The eggs hatch into aquatic larvae[27] called tadpoles[28] that have tails and internal
gills[29]. They have highly specialized rasping mouth parts suitable for herbivorous[30], omnivorous[31] or planktivorous[32]
diets. The life cycle[33] is completed when they metamorphose[34] into adults. A few species deposit eggs on land or bypass the
tadpole stage. Adult frogs generally have a carnivorous diet consisting of small invertebrates, but omnivorous species exist and
a few feed on fruit. Frogs are extremely efficient at converting what they eat into body mass, which makes them an important
food source for predators[35]. Frogs are a keystone group[36] in the food web[37] dynamics of many of the world's
ecosystems[38]. The skin is semi-permeable[39], making them susceptible to dehydration, so they either live in moist places or
have special adaptations to deal with dry habitats. Frogs produce a wide range of vocalizations[40], particularly in their
breeding season[41], and exhibit many different kinds of complex behaviours[42] to attract mates, to fend off predators and to
generally survive.
Frog populations have declined significantly since the 1950s. More than one third of species are considered to be threatened[43]
with extinction[44] and over one hundred and twenty are believed to have become extinct since the 1980s.1 The number of
malformations among frogs is on the rise and an emerging fungal[45] disease, chytridiomycosis[46], has spread around the
world. Conservation biologists[47] are working to understand the causes of these problems and to resolve them. Frogs are
valued as food by humans and also have many cultural roles in literature, symbolism and religion.

Etymology and taxonomy

The name frog derives from Old English[48] frogga, abbreviated to frox, forsc, and frosc, probably deriving from Proto-IndoEuropean[49] preu = "to jump".2 About 88% of amphibian[50] species are classified[51] in the order[52] Anura.3 These include
around 4,810 species in 33 families[53], of which the Leptodactylidae[54] (1,100 spp.), Hylidae[55] (800 spp.) and Ranidae[56]
(750 spp.) are the richest in species[57].3

[58]

The use of the common names "frog" and "toad[59]" has no taxonomic justification. From a classification perspective, all
members of the order Anura are frogs, but only members of the family Bufonidae[60] are considered "true toads". The use of the
term "frog" in common names usually refers to species that are aquatic or semi-aquatic and have smooth, moist skins; the term
"toad" generally refers to species that are terrestrial with dry, warty skins.45 There are numerous exceptions to this rule. The
European fire-bellied toad[61] (Bombina bombina) has a slightly warty skin and prefers a watery habitat6 whereas the
Panamanian golden frog[62] (Atelopus zeteki) is in the toad family Bufonidae and has a smooth skin.7
The Anura include all modern frogs and any fossil[63] species that fit within the anuran definition. The characteristics of
anuran adults include: 9 or fewer presacral vertebrae, a long and forward-sloping ilium, the presence of a urostyle[64], no tail,
shorter fore limbs than hind limbs, radius[65] and ulna[66] fused, tibia[67] and fibula[68] fused, elongated ankle bones[69],
absence of a prefrontal bone, presence of a hyoid plate[70], a lower jaw[71] without teeth (with the exception of Gastrotheca
guentheri[72]) consisting of three pairs of bones (angulosplenial, dentary, and mentomeckelian, with the last pair being absent
in Pipoidea[73]),8 an unsupported tongue, lymph[74] spaces underneath the skin, and a muscle, the protractor lentis, attached
to the lens of the eye[75].9 The anuran larva or tadpole has a single central respiratory spiracle[76] and mouthparts consisting of
keratinous[77] beaks[78] and denticles[79].9

[80]

Frogs and toads are broadly classified into three suborders: Archaeobatrachia[81], which includes four families of primitive
frogs; Mesobatrachia[82], which includes five families of more evolutionary intermediate frogs; and Neobatrachia[83], by far the
largest group, which contains the remaining 24 families of modern frogs, including most common species found throughout
the world. The Neobatrachia suborder is further divided into the two superfamilies Hyloidea[84] and Ranoidea[85].10 This
classification is based on such morphological[86] features as the number of vertebrae, the structure of the pectoral girdle[87],
and the morphology of tadpoles. While this classification is largely accepted, relationships among families of frogs are still
debated.11
Some species of anurans hybridize[88] readily. For instance, the edible frog[89] (Pelophylax esculentus) is a hybrid between the
pool frog[90] (P. lessonae) and the marsh frog[91] (P. ridibundus).12 The fire-bellied toads Bombina bombina and B. variegata[92]
are similar in forming hybrids. These are less fertile than their parents, giving rise to a hybrid zone[93] where the hybrids are
prevalent.13

Evolution
The origins and evolutionary relationships between the three main groups of amphibians are hotly debated. A molecular
phylogeny[94] based on rDNA[95] analysis dating from 2005 suggests salamanders[96] and caecilians[97] are more closely related
to each other than they are to frogs and the divergence[98] of the three groups took place in the Paleozoic[99] or early
Mesozoic[100] before the breakup of the supercontinent Pangaea[101] and soon after their divergence from the lobe-finned
fishes[102]. This would help account for the relative scarcity of amphibian fossils from the period before the groups split.14
Another molecular phylogenetic analysis conducted about the same time concluded the lissamphibians[103] first appeared
about 330 million years ago and that the temnospondyl-origin[104] hypothesis is more credible than other theories. The
neobatrachians[105] seemed to have originated in Africa/India, the salamanders in East Asia and the caecilians in tropical
Pangaea.15 Other researchers, while agreeing with the main thrust of this study, questioned the choice of calibration points
used to synchronise the data. They proposed that the date of lissamphibian diversification be put in the Permian[106], rather
less than 300 million years ago, a date in better agreement with the palaeontological data.16 A further study in 2011 using both
extinct and living taxa sampled for morphological, as well as molecular data, came to the conclusion that Lissamphibia is
monophyletic[107] and that it should be nested within Lepospondyli[108] rather than within Temnospondyli[109]. The study
postulated the Lissamphibia originated no earlier than the late Carboniferous[110], some 290 to 305 million years ago. The split
between Anura and Caudata[111] was estimated as taking place 292 million years ago, rather later than most molecular studies
suggest, with the caecilians splitting off 239 million years ago.17
In 2008, Gerobatrachus hottoni[112], a temnospondyl[113] with many frog- and salamander-like characteristics, was discovered
in Texas[114]. It dated back 290 million years and was hailed as a missing link[115], a stem[116] batrachian close to the common
ancestor[117] of frogs and salamanders, consistent with the widely accepted hypothesis that frogs and salamanders are more
closely related to each other (forming a clade[118] called Batrachia) than they are to caecilians.1819 However, others have
suggested that Gerobatrachus hottoni[119] was only a dissorophoid[120] temnospondyl unrelated to extant amphibians.20 The
earliest known salientians (see below), closer to the extant frogs than to the extant salamanders, are Triadobatrachus[121]
massinoti, from the Early Triassic[122] of Madagascar[123] (about 250 million years ago), and the fragmentary
Czatkobatrachus[124] polonicus from the Early Triassic of Poland[125] (about the same age as Triadobatrachus).21 The skull of
Triadobatrachus is frog-like, being broad with large eye sockets, but the fossil has features diverging from modern frogs. These
include a longer body with more vertebrae[126]. The tail has separate vertebrae unlike the fused urostyle or coccyx found in
modern frogs. The tibia and fibula bones are also separate, making it probable that Triadobatrachus was not an efficient
leaper.21
Salientia (Latin salere (salio), "to jump") is a stem group[127] including modern frogs in the order Anura and their close fossil
relatives the "proto-frogs" (e.g., Triadobatrachus[128] and Czatkobatrachus[129]). The common features possessed by the "protofrogs" in the Salientia[130] group include 14 presacral vertebrae[131] (modern frogs have eight or 9), a long and forward-sloping
ilium[132] in the pelvis[133], the presence of a frontoparietal bone[134], and a lower jaw[135] without teeth. The earliest frog
fossil that falls into the anuran lineage proper, Prosalirus bitis[136], lived in the early Jurassic[137].422 It was discovered in
1995[138] in the Kayenta Formation[139] of Arizona[140] and dates back to the Early Jurassic[141] epoch (199.6 to 175 million
years ago), making Prosalirus somewhat more recent than Triadobatrachus.23 Like the latter, Prosalirus did not have greatly
enlarged legs, but had the typical three-pronged pelvic[142] structure of modern frogs. Unlike Triadobatrachus, Prosalirus had
already lost nearly all of its tail24 and was well adapted for jumping.25
The earliest known "true frog" is Vieraella herbsti[143], from the Early Jurassic. It is known only from the dorsal[144] and
ventral[145] impressions of a single animal and was estimated to be 33 mm (1.3 in) from snout to vent. Notobatrachus
degiustoi[146] from the middle Jurassic is slightly younger, about 155170 million years old. The main evolutionary changes in
this species involved the shortening of the body and the loss of the tail. The evolution of modern Anura likely was complete by
the Jurassic period. Since then, evolutionary changes in chromosome numbers[147] have taken place about 20 times faster in

mammals than in frogs, which means speciation[148] is occurring more rapidly in mammals.26
Frog fossils have been found on all continents except Antarctica[149], but biogeographic[150] evidence suggests they also
inhabited Antarctica in an earlier era when the climate was warmer.27
[show]Frog classification
A cladogram[151] showing the relationships of the different families[152] of frogs in the clade Anura can be seen in the table
above. This diagram, in the form of a tree[153], shows how each frog family is related to other families, with each node
representing a point of common ancestry. It is based on Frost et al. (2006)28 and Heinicke et al. (2009).29 and Pyron and Wiens
(2011).30

Morphology and physiology

Frogs have no tail, except as larvae, and most have long hind legs, elongated ankle bones, webbed toes, no claws, large eyes,
and a smooth or warty skin. They have short vertebral columns, with no more than 10 free vertebrae and fused tailbones
(urostyle or coccyx).31 Like other amphibians, oxygen[154] can pass through their highly permeable skins. This unique feature
allows them to remain in places without access to the air, respiring through their skins.32 The ribs are poorly developed, so the
lungs are filled by buccal pumping[155] and a frog deprived of its lungs can maintain its body functions without them.32 For the
skin to serve as a respiratory organ, it must remain moist. This makes frogs susceptible to various substances they may
encounter in the environment, some of which may be toxic and can dissolve in the water film and be passed into their
bloodstream. This may be one of the causes of the worldwide decline in frog populations[156].33343536
Frogs range in size from the recently discovered 7.7-mm-long Paedophryne amauensis[157] of Papua New Guinea[158] 37 to the
300-mm goliath frog[159] (Conraua goliath) of Cameroon[160]. The skin hangs loosely on the body because of the lack of loose
connective tissue[161]. Frogs have three eyelid membranes: one is transparent to protect the eyes underwater, and two vary
from translucent to opaque. They have a tympanum[162] on each side of their heads which is involved in hearing and, in some
species, is covered by skin. True toads completely lack teeth, but most frogs have them, specifically pedicellate teeth[163] in
which the crown is separated from the root by fibrous tissue. These are on the edge of the upper jaw and vomerine teeth[164]
are also on the roof of their mouths. No teeth are in the lower jaw and frogs usually swallow their food whole. The teeth are
mainly used to grip the prey and keep it in place till swallowed, a process assisted by retracting the eyes into the head.38 The
African bullfrog (Pyxicephalus[165]), which preys on relatively large animals such as mice and other frogs, has cone shaped bony
projections called odontoid processes at the front of the lower jaw which function like teeth.3

Feet and legs

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 to do so. Most frogs are either proficient at jumping or are descended
from ancestors that were, with much of the musculoskeletal[166] morphology[167] modified for this purpose. The tibia, fibula,
and tarsals[168] have been fused into a single, strong bone[169], as have the radius and ulna in the fore limbs (which must
absorb the impact on landing). The metatarsals[170] have become elongated to add to the leg length and allow the frog to push
against the ground for a longer period on take-off. The illium[171] has elongated and formed a mobile joint with the sacrum[172]
which, in specialist jumpers such as ranids and hylids, functions as an additional limb joint to further power the leaps. The tail
vertebrae have fused into a urostyle which is retracted inside the pelvis. This enables the force to be transferred from the legs
to the body during a leap.31

[173]

Webbed hind foot of common frog

(Rana temporaria)
The muscular system has been similarly modified. The hind limbs of ancestral 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 main leg muscles accounting for over 17% of the total mass of the frog.39
Many frogs have webbed feet and the degree of webbing is directly proportional to the amount of time the species spends in the
water.40 The completely aquatic African dwarf frog[174] (Hymenochirus sp.) has fully webbed toes, whereas those of White's tree
frog[175] (Litoria caerulea), an arboreal species, are only a quarter or half webbed.41
Arboreal frogs[176] have pads located on the ends of their toes to help grip vertical surfaces. These are not suction pads, the
surface consisting instead of columnar cells with flat tops with small gaps between them lubricated by mucous glands. When
the frog applies pressure, the cells adhere to irregularities on the surface and the grip is maintained through surface
tension[177]. This allows the frog to climb on smooth surfaces, but the system does not function efficiently when the pads are
excessively wet.42
In many arboreal frogs, a small "intercalary structure" on each toe increases the surface area touching the substrate[178].
Furthermore, since hopping through trees can be dangerous, many arboreal frogs have hip joints to allow both hopping and
walking. Some frogs that live high in trees even possess an elaborate degree of webbing between their toes. This allows the
frogs to "parachute" or make a controlled glide from one position in the canopy to another.43
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 such as Couch's spadefoot[179] (Scaphiopus couchii) have a flap-like toe extension on the
hind feet, a keratinised[180] tubercle[181] often referred to as a spade, that helps them to burrow.44
Sometimes during the tadpole stage, one of the developing rear legs is eaten by a predator such as a dragonfly nymph[182]. In
some cases, the full leg still grows, but in others it does not, although the frog may still live out its normal lifespan with only
three limbs. Occasionally, a parasitic flatworm[183] (Ribeiroia ondatrae[184]) digs into the rear of a tadpole, causing a
rearrangement of the limb bud cells and the frog develops an extra leg or two.45
A frog's skin is protective, has a respiratory function, can absorb water and helps control body temperature. It has many glands,
particularly on the head and back, which often exude distasteful and toxic substances. The secretion is often sticky and helps
keep the skin moist, protects against the entry of moulds and bacteria, and make the animal slippery and more able to escape
from predators.46 The skin is shed every few weeks. It usually splits down the middle of the back and across the belly, and the
frog pulls its arms and legs free. The sloughed skin is then worked towards the head where it is quickly eaten.47
Being cold-blooded, frogs have to adopt suitable behaviour patterns to regulate their temperature. To warm up, they can move
into the sun or onto a warm surface; if they overheat, they can move into the shade or adopt a stance that exposes the
minimum area of skin to the air. This posture is also used to prevent water loss and involves the frog squatting close to the
substrate with its hands and feet tucked under its chin and body.48 The colour of a frog's skin is used for thermoregulation. In

cool damp conditions, the colour will be darker than on a hot dry day. The grey foam-nest tree frog[185] (Chiromantis
xerampelina) is even able to turn white to minimize the chance of overheating.49
Many frogs are able to absorb water and oxygen directly through the skin, especially around the pelvic area, but the
permeability of a frog's skin can also result in water loss. Glands located all over the body exude mucus which helps keep the
skin moist and reduces evaporation. Some glands on the hands and chest of males are specialized to produce sticky secretions
to aid in amplexus[186]. Similar glands in tree frogs produce a glue-like substance on the adhesive discs of the feet. Some
arboreal frogs reduce water loss by having a waterproof layer of skin, and several South American species coat their skin with a
waxy secretion. Others frogs have adopted behaviours to conserve water, including becoming nocturnal[187] and resting in a
water-conserving position. Some frogs may also rest in large groups with each frog pressed against its neighbours. This reduces
the amount of skin exposed to the air or a dry surface, and thus reduces water loss.48 Woodhouse's toad[188] (Bufo woodhousii),
if given access to water after confinement in a dry location, sits in the shallows to rehydrate.50 The male hairy frog[189]
(Trichobatrachus robustus) has dermal papillae[190] projecting from its lower back and thighs, giving it a bristly appearance.
They contain blood vessels and are thought to increase the area of the skin available for respiration.51
Some species have bony plates[191] embedded in their skin, a trait that appears to have evolved independently several times.52
In certain other species, the skin at the top of the head is compacted and the connective tissue of the dermis is co-ossified with
the bones of the skull (exostosis[192]).5354
Camouflage[193] is a common defensive mechanism in frogs. Most camouflaged frogs are nocturnal; during the day, they seek
out a position where they can blend into the background and remain undetected. Some frogs have the ability to change
colour[194], but this is usually restricted to a small range of colours. For example, White's tree frog (Litoria caerulea) varies
between pale green and dull brown according to the temperature, and the Pacific tree frog[195] (Pseudacris regilla) has green
and brown morphs, plain or spotted, and changes colour depending on the time of year and general background colour.55
Features such as warts and skin folds[196] are usually found on ground-dwelling frogs, where a smooth skin would not provide
such effective camouflage. Certain frogs change colour between night and day, as light and moisture stimulate the pigment
cells and cause them to expand or contract.32

[197]

Pouched frog[198] (Assa darlingtoni) camouflaged against leaf litter

Respiration and circulation

The skin of a frog is permeable to oxygen[199] and carbon dioxide[200], as well as to water. There are blood vessels near the
surface of the skin and when a frog is underwater, oxygen diffuses directly into the blood. When not submerged, a frog breathes
by a process known as buccal pumping[201]. Its lungs are similar to those of humans, but the chest muscles are not involved in
respiration, and no ribs[202] or diaphragm[203] exist to help move air in and out. Instead, it puffs out its throat and draws air in
through the nostrils, which in many species can then be closed by valves. When the floor of the mouth is compressed, air is

forced into the lungs.56 The Borneo flat-headed frog[204] (Barbourula kalimantanensis) was first discovered in a remote part of
Indonesia in 2007. It is entirely aquatic and is the first species of frog known to science that has no lungs.57
Frogs have three-chambered hearts[205], a feature they share with lizards[206].58 Oxygenated blood from the lungs and deoxygenated blood from the respiring[207] tissues enter the heart through separate atria[208]. When these chambers contract,
the two blood streams pass into a common ventricle[209] before being pumped via a spiral valve to the appropriate vessel, the
aorta[210] for oxygenated blood and pulmonary artery[211] for deoxygenated blood. The ventricle is partially divided into
narrow cavities which minimizes the mixing of the two types of blood. These features enable frogs to have a higher metabolic
rate and be more active than would otherwise be possible.58
Some species of frog have adaptations that allow them to survive in oxygen deficient water. The Lake Titicaca[212] frog
(Telmatobius culeus[213]) is one such species and has wrinkly skin that increases its surface area to enhance gas exchange. It
normally makes no use of its rudimentary lungs but will sometimes raise and lower its body rhythmically while on the lake bed
to increase the flow of water around it.59
[214]

Dissected frog:1 Right atrium, 2 Lungs, 3 Aorta, 4 Egg mass, 5 Colon, 6 Left atrium,
7 Ventricle, 8 Stomach, 9 Liver, 10 Gallbladder, 11 Small intestine, 12 Cloaca

Digestion and excretion

Frogs have maxillary teeth along their upper jaw which are used to hold food before it is
swallowed. These teeth are very weak, and cannot be used to chew or catch and harm agile
prey. Instead, the frog uses its sticky, cleft tongue to catch flies and other small moving prey.
The tongue normally lies coiled in the mouth, free at the back and attached to the mandible at
the front. It can be shot out and retracted at great speed.40 Some frogs have no tongue and
just stuff food into their mouths with their hands.40 The eyes assist in the swallowing of food
as they can be retracted through holes in the skull and help push food down the throat.40 The food then moves through the
oesophagus into the stomach where digestive enzymes are added and it is churned up. It then proceeds to the small intestine
(duodenum and ileum) where most digestion occurs. Pancreatic juice from the pancreas, and bile, produced by the liver and
stored in the gallbladder, are secreted into the small intestine, where the fluids digest the food and the nutrients are absorbed.
The food residue passes into the large intestine where excess water is removed and the wastes are passed out through the
cloaca[215].60
Although adapted to terrestrial life, frogs resemble freshwater fish in their inability to conserve body water effectively. When
they are on land, much water is lost by evaporation from the skin. The excretory system is similar to that of mammals and
there are two kidneys[216] that remove nitrogenous products from the blood. Frogs produce large quantities of dilute urine in
order to flush out toxic products from the kidney tubules.61 The nitrogen is excreted as ammonia[217] by tadpoles and aquatic
frogs but mainly as urea[218], a less toxic product, by most terrestrial adults. A few species of tree frog with little access to
water excrete the even less toxic uric acid[219].61 The urine passes along paired ureters[220] to the urinary bladder[221] from
which it is vented periodically into the cloaca. All bodily wastes exit the body through the cloaca which terminates in a cloacal
vent.62

Reproductive system
In the male frog, the two testes[222] are attached to the kidneys and semen[223] passes into the kidneys through fine tubes
called efferent ducts[224]. It then travels on through the ureters, which are consequently known as urinogenital ducts. There is
no penis, and sperm is ejected from the cloaca directly onto the eggs as the female lays them. The ovaries of the female frog are
beside the kidneys and the eggs pass down a pair of oviducts and through the cloaca to the exterior.62

When frogs mate, the male climbs on the back of the female and wraps his fore limbs round her body, either behind the front
legs or just in front of the hind legs. This position is called amplexus[225] and may be held for several days.63 The male frog has
certain hormone-dependent secondary sexual characteristics[226]. These include the development of special pads on his
thumbs in the breeding season, to give him a firm hold.64 The grip of the male frog during amplexus stimulates the female to
release eggs, usually wrapped in jelly, as spawn.62 In many species the male is smaller and slimmer than the female. Males
have vocal cords and make a range of croaks, particularly in the breeding season, and in some species they also have vocal
sacs[227] to amplify the sound.62

Nervous system
The frog has a highly developed nervous system that consists of a brain, spinal cord and nerves. Many parts of the frog's brain
correspond with those of humans. It consists of two olfactory lobes, two cerebral hemispheres, a pineal body, two optic lobes, a
cerebellum and a medulla oblongata. Muscular coordination and posture are controlled by the cerebellum[228], and the medulla
oblongata[229] regulates respiration, digestion and other automatic functions.62 The relative size of the cerebrum[230] in frogs
is much smaller than it is in humans. Frogs have ten pairs of cranial nerves[231] which pass information from the outside
directly to the brain, and ten pairs of spinal nerves[232] which pass information from the extremities to the brain through the
spinal cord.62 By contrast, all amniotes[233] (mammals, birds and reptiles) have twelve pairs of cranial nerves.65

[234]

Close-up of frog's head showing eye, nostril, mouth and tympanum.

Sight
The eyes of most frogs are located on either side of the head near the top and project outwards as hemispherical bulges. They
provide binocular vision[235] over a field of 100 to the front and a total visual field of almost 360.66 They may be the only part
of an otherwise submerged frog to protrude from the water. Each eye has closable upper and lower lids and a nictitating
membrane[236] which provides further protection, especially when the frog is swimming.67 Members of the aquatic family
Pipidae[237] have the eyes located at the top of the head, a position better suited for detecting prey in the water above.66 The
irises[238] come in a range of colours and the pupils in a range of shapes. The common toad[239] (Bufo bufo) has golden irises
and horizontal slit-like pupils, the red-eyed tree frog[240] (Agalychnis callidryas) has vertical slit pupils, the poison dart frog[241]
has dark irises, the fire-bellied toad[242] (Bombina spp.) has triangular pupils and the tomato frog[243] (Dyscophus spp.) has
circular ones. The irises of the southern toad[244] (Anaxyrus terrestris) are patterned so as to blend in with the surrounding
camouflaged skin.67
The distant vision of a frog is better than its near vision. Calling frogs will quickly become silent when they see an intruder or
even a moving shadow but the closer an object is, the less well it is seen.67 When a frog shoots out its tongue to catch an insect
it is reacting to a small moving object that it cannot see well and must line it up precisely beforehand because it shuts its eyes
as the tongue is extended.40 Whether a frog sees in colour is debatable but it has been shown that it responds positively to blue
light, perhaps because that colour is associated with bodies of water that can provide refuge when the frog feels threatened.68

Hearing
Frogs can hear both in the air and below water. They do not have external ears; the eardrums (tympanic membranes[245]) are

directly exposed or may be covered by a layer of skin and are visible as a circular area just behind the eye. The size and distance
apart of the eardrums is related to the frequency and wavelength at which the frog calls. In some species such as the bullfrog,
the size of the tympanum indicates the sex of the frog; males have tympani that are larger than their eyes while in females, the
eyes and tympani are much the same size.69 A noise causes the tympanum to vibrate and the sound is transmitted to the
middle and inner ear. The middle ear contains semicircular canals which help control balance and orientation. In the inner ear,
the auditory hair cells are arranged in two areas of the cochlea, the basilar papilla and the amphibian papilla. The former
detects high frequencies and the latter low frequencies.70 Because the cochlea is short, frogs use electrical tuning[246] to
extend their range of audible frequencies and help discriminate different sounds.71 This arrangement enables detection of the
territorial and breeding calls of their conspecifics[247]. In some species that inhabit arid regions, the sound of thunder or heavy
rain may arouse them from a dormant state.70 A frog may be startled by an unexpected noise but it will not usually take any
action until it has located the source of the sound by sight.69
See also: Sexual selection in frogs[248]
The call or croak of a frog is unique to its species. Frogs create this sound by passing air through the larynx[249] 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. Some frog calls are so loud that they can be heard up to a mile
away.72
Frogs in the genera Heleioporus[250] and Neobatrachus[251] lack vocal sacs but can still produce a loud call. Their buccal
cavity[252] is enlarged and dome-shaped, acting as a resonance chamber[253] that amplifies the sound. Species of frog that lack
vocal sacs and that do not have a loud call tend to inhabit areas close to constantly noisy, flowing water. They need to use an
alternative means to communicate. The coastal tailed frog[254] (Ascaphus truei) lives in mountain streams in North America
and does not vocalize.73
The main reason for calling is to allow male frogs to attract a mate. Males may call individually or there may be a chorus of
sound where numerous males have converged on breeding sites. Females of many frog species, such as the common tree
frog[255] (Polypedates leucomystax), reply to the male calls, which acts to reinforce reproductive activity in a breeding colony.74
Female frogs prefer males that produce sounds of greater intensity and lower frequency, attributes that stand out in a crowd.
The rationale for this is thought to be that by demonstrating his prowess, the male shows his fitness to produce superior
offspring.75
A different call is emitted by a male frog or unreceptive female when mounted by another male. This is a distinct chirruping
sound and is accompanied by a vibration of the body.76 Tree frogs and some non-aquatic species have a rain call that they
make on the basis of humidity cues prior to a shower.76 Many species also have a territorial call that is used to drive away other
males. All of these calls are emitted with the mouth of the frog closed.76 A distress call, emitted by some frogs when they are in
danger, is produced with the mouth open resulting in a higher-pitched call. It is typically used when the frog has been grabbed
by a predator and may serve to distract or disorientate the attacker so that it releases the frog.76
Many species of frog have deep calls. The croak of the American bullfrog[256] (Rana catesbiana) is sometimes written as "jug o'
rum".77 The Pacific tree frog[257] (Pseudacris regilla) produces the onomatopoeic[258] "ribbit" often heard in films.78 Other
renderings of frog calls into speech include "brekekekex koax koax", the call of the marsh frog (Pelophylax ridibundus[259]) in
The Frogs, an Ancient Greek comic drama by Aristophanes[260].79

Torpor
During extreme conditions, some frogs enter a state of torpor[261] and remain inactive for months. In colder regions, many
species of frog hibernate[262] in winter. Those that live on land such as the American toad[263] (Bufo americanus) dig a burrow
and make a hibernaculum[264] in which to lie dormant[265]. Others, less proficient at digging, find a crevice or bury themselves

in dead leaves. Aquatic species such as the American bullfrog[266] (Rana catesbeiana) normally sink to the bottom of the pond
where they lie, semi-immersed in mud but still able to access the oxygen dissolved in the water. Their metabolism slows down
and they live on their energy reserves. Some frogs can even survive being frozen. Ice crystals form under the skin and in the
body cavity but the essential organs are protected from freezing by a high concentration of glucose. An apparently lifeless,
frozen frog can resume respiration and the heart beat can restart when conditions warm up.80
At the other extreme, the striped burrowing frog[267] (Cyclorana alboguttata) regularly aestivates[268] during the hot, dry season
in Australia, surviving in a dormant state without access to food and water for nine or ten months of the year. It burrows
underground and curls up inside a protective cocoon[269] formed by its shed skin. Researchers at the University of
Queensland[270] have found that during aestivation, the metabolism[271] of the frog is altered and the operational efficiency of
the mitochondria[272] is increased. This means that the limited amount of energy available to the comatose frog is used in a
more efficient manner. This survival mechanism is only useful to animals that remain completely unconscious for an extended
period of time and whose energy requirements are low because they are cold-blooded and have no need to generate heat.81
Other research showed that, to provide these energy requirements, muscles atrophy, but hind limb muscles are preferentially
unaffected.82

Locomotion
Different species of frog use a number of methods of moving around including jumping[273], running[274], walking[275],
swimming[276], burrowing[277], climbing[278] and gliding[279].

[280]

Jumping
Frogs are generally recognized as exceptional jumpers and, relative to their size, the best jumpers of all vertebrates.83 The
Australian rocket frog, Litoria nasuta[281], can leap over 2 metres (6 ft 7 in), a distance that is more than fifty times its body
length of 5.5 centimetres (2.2 in).84 There are tremendous differences between species in jumping capability. Within a species,
jump distance increases with increasing size, but relative jumping distance (body-lengths jumped) decreases. The Indian
skipper frog[282] (Euphlyctis cyanophlyctis) has the ability to leap out of the water from a position floating on the surface.85 The
tiny northern cricket frog[283] (Acris crepitans) can "skitter" across the surface of a pond with a series of short rapid jumps.86
Slow-motion photography shows that the muscles have passive flexibility. They are first stretched while the frog is still in the
crouched position, then they are contracted before being stretched again to launch the frog into the air. The fore legs are
folded against the chest and the hind legs remain in the extended, streamlined position for the duration of the jump.39 In
some extremely capable jumpers, such as the Cuban tree frog[284] (Osteopilus septentrionalis) and the northern leopard frog[285]
(Rana pipiens), the peak power exerted during a jump can exceed that which the muscle is theoretically capable of producing.
When the muscles contract, the energy is first transferred into the stretched tendon which is wrapped around the ankle bone.
Then the muscles stretch again at the same time as the tendon releases its energy like a catapult[286] to produce a powerful
acceleration beyond the limits of muscle-powered acceleration.87 A similar mechanism has been documented in locusts[287]
and grasshoppers[288].88
Walking and running

[289]

Frogs in the families Bufonidae, Rhinophrynidae[290], and Microhylidae[291] have short back legs and tend to walk rather than
jump.89 When they try to move rapidly, they speed up the rate of movement of their limbs or resort to an ungainly hopping
gait. The western narrow-mouthed toad[292] (Gastrophryne olivacea) has been described as having a gait that is "a combination
of running and short hops that are usually only an inch or two in length".90 In an experiment, Fowler's toad[293] (Bufo fowleri)
was placed on a treadmill which was turned at varying speeds. By measuring the toad's uptake of oxygen it was found that
hopping was an inefficient use of resources during sustained locomotion but was a useful strategy during short bursts of highintensity activity.91
The red-legged running frog[294] (Kassina maculata) has short, slim hind limbs unsuited to jumping. It can move fast by using a
running gait in which the two hind legs are used alternately. Slow-motion photography shows, unlike a horse that can trot or
gallop, the frog's gait remained similar at slow, medium, and fast speeds.92 This species can also climb trees, and shrubs and
does so at night to catch insects.93 The Indian skipper frog (Euphlyctis cyanophlyctis) has broad feet and can run across the
surface of the water for several metres (yards).86
Swimming

[295]

Common toad[296] (Bufo bufo) swimming.

Frogs that live in or visit water have adaptations that improve their swimming abilities. The hind limbs are heavily muscled and
strong. The webbing between the toes of the hind feet increases the area of the foot and helps propel the frog powerfully
through the water. Members of the family Pipidae[297] are wholly aquatic and show the most marked specialization. They have
inflexible vertebral columns, flattened, streamlined bodies, lateral line[298] systems, and powerful hind limbs with large
webbed feet.94 Tadpoles mostly have large tail fins which provide thrust when the tail is moved from side to side.95
Burrowing
Some frogs have become adapted for burrowing and a life underground. They tend to have rounded bodies, short limbs, small
heads with bulging eyes, and hind feet adapted for excavation. An extreme example of this is the purple frog[299]
(Nasikabatrachus sahyadrensis) from southern India which feeds on termites[300] and spends almost its whole life underground.
It emerges briefly during the monsoon[301] to mate and breed in temporary pools. It has a tiny head with a pointed snout and a
plump, rounded body. Because of this fossorial[302] existence, it was first described[303] in 2003, being new to the scientific
community at that time, although previously known to local people.96

[304]

Purple frog (Nasikabatrachus sahyadrensis).

The spadefoot toads of North America are also adapted to underground life. The plains spadefoot toad[305] (Spea bombifrons) is
typical and has a flap of keratinised bone attached to one of the metatarsals[306] of the hind feet which it uses to dig itself
backwards into the ground. As it digs, the toad wriggles its hips from side to side to sink into the loose soil. It has a shallow
burrow in the summer from which it emerges at night to forage. In winter, it digs much deeper and has been recorded at a
depth of 4.5 m (15 ft).97 The tunnel is filled with soil and the toad hibernates in a small chamber at the end. During this time,
urea accumulates in its tissues and water is drawn in from the surrounding damp soil by osmosis[307] to supply the toad's
needs.97 Spadefoot toads are "explosive breeders", all emerging from their burrows at the same time and converging on
temporary pools, attracted to one of these by the calling of the first male to find a suitable breeding location.98
The burrowing frogs of Australia have a rather different lifestyle. The western spotted frog[308] (Heleioporus albopunctatus) digs
a burrow beside a river or in the bed of an ephemeral stream and regularly emerges to forage. Mating takes place and eggs are
laid in a foam nest inside the burrow. The eggs partially develop there, but do not hatch until they are submerged following
heavy rainfall. The tadpoles then swim out into the open water and rapidly complete their development.99 Madagascan
burrowing frogs are less fossorial and mostly bury themselves in leaf litter. One of these, the green burrowing frog[309]
(Scaphiophryne marmorata), has a flattened head with a short snout and well-developed metatarsal tubercles on its hind feet to
help with excavation. It also has greatly enlarged terminal discs on its fore feet that help it to clamber around in bushes.100 It
breeds in temporary pools that form after rains.101
Climbing

[310]

Tree frogs are found high in the canopy[311], where they scramble around on the branches, twigs, and leaves, sometimes never
coming down to earth. The "true" tree frogs belong to the family Hylidae, but members of other frog families have
independently adopted an arboreal habit, a case of convergent evolution[312]. These include the glass frogs[313]
(Centrolenidae), the bush frogs[314] (Hyperoliidae), some of the narrow-mouthed frogs (Microhylidae), and the shrub frogs[315]
(Rhacophoridae).89 Most tree frogs are under 10 cm (4 in) in length, with long legs and long toes with adhesive pads on the
tips. The surface of the toe pads is formed from a closely packed layer of flat-topped, hexagonal epidermal[316] cells separated
by grooves into which glands secrete mucus[317]. These toe pads, moistened by the mucus, provide the grip on any wet or dry
surface, including glass. The forces involved include boundary friction[318] of the toe pad epidermis on the surface and also
surface tension[319] and viscosity[320].102 Tree frogs are very acrobatic and can catch insects while hanging by one toe from a
twig or clutching onto the blade of a windswept reed.103 Some members of the subfamily Phyllomedusinae[321] have
opposable toes on their feet. The reticulated leaf frog[322] (Phyllomedusa ayeaye) has a single opposed digit[323] on each fore
foot and two opposed digits on its hind feet. This allows it to grasp the stems of bushes as it clambers around in its riverside
habitat.104
Gliding

During the evolutionary history of the frog, several different groups have independently taken to the air.105 Some frogs in the
tropical rainforest are specially adapted for gliding from tree to tree or parachuting to the forest floor. Typical of them is
Wallace's flying frog[324] (Rhacophorus nigropalmatus) from Malaysia and Borneo. It has large feet with the fingertips expanded
into flat adhesive discs and the digits fully webbed.Flaps of skin occur on the lateral margins of the limbs and across the tail
region. With the digits splayed, the limbs outstretched, and these flaps spread, it can glide considerable distances, but is unable
to undertake powered flight.106 It can alter its direction of travel and navigate distances of up to 15 m (49 ft) between trees.107

Life history
Like other amphibians, the life cycle of a frog normally starts in water with an egg that hatches into a limbless larva with gills,
commonly known as a tadpole[325]. After further growth, during which it develops limbs and lungs, the tadpole undergoes
metamorphosis[326] in which its appearance and internal organs are rearranged. After this it is able to leave the water as a
miniature, air-breathing frog.

Reproduction
Two main types of reproduction occur in frogs, prolonged breeding and explosive breeding. In the former, adopted by the
majority of species, adult frogs at certain times of year assemble at a pond, lake or stream to breed. Many frogs return to the
bodies of water in which they developed as larvae. This often results in annual migrations involving thousands of individuals.
In explosive breeders, mature adult frogs arrive at breeding sites in response to certain trigger factors such as rainfall occurring
in an arid area. In these frogs, mating and spawning take place promptly and the speed of larval growth is rapid in order to
make use of the ephemeral pools before they dry up.108
Among prolonged breeders, males usually arrive at the breeding site first and remain there for some time whereas females tend
to arrive later and depart soon after they have spawned. This means that males outnumber females at the water's edge and
defend territories from which they expel other males. They advertise their presence by calling, often alternating their croaks
with neighbouring frogs. Larger, stronger males tend to have deeper calls and maintain higher quality territories. Females
select their mates at least partly on the basis of the depth of their voice.109 In some species there are satellite males who have
no territory and do not call. They may intercept females that are approaching a calling male or take over a vacated territory.
Calling is an energy-sapping activity. Sometimes the two roles are reversed and a calling male gives up its territory and
becomes a satellite.108
In explosive breeders, the first male that finds a suitable breeding location such as a temporary pool, calls loudly and other
frogs of both sexes converge on the pool. Explosive breeders tend to call in unison creating a chorus that can be heard from far
away. The spadefoot toads (Scaphiopus[327] spp.) of North America fall into this category. Mate selection and courtship is not as
important as speed in reproduction. In some years, suitable conditions may not occur and the frogs may go for two or more
years without breeding.108 Some female New Mexico spadefoot toads[328] (Spea multiplicata) only spawn half of the available
eggs at a time, perhaps retaining some in case a better reproductive opportunity arises later.110
At the breeding site, the male mounts the female and grips her tightly round the body. Typically, amplexus takes place in the
water, the female releases her eggs and the male covers them with sperm; fertilization is external[329]. In many species such as
the Great Plains toad[330] (Bufo cognatus), the male restrains the eggs with his back feet, holding them in place for about three
minutes.108 Members of the West African genus Nimbaphrynoides[331] are unique among frogs in that they are viviparous[332];
members of the Tanzanian[333] genus Nectophrynoides[334] are the only frogs known to be ovoviviparous[335]. In both cases,
fertilization is internal[336] and females give birth to fully developed juvenile frogs.111112

Life cycle

Eggs / frogspawn

[337]

Frogspawn
Frogs' embryos are typically surrounded by several layers of gelatinous material. When several eggs are clumped together, they
are collectively known as frogspawn[338]. The jelly provides support and protection while allowing the passage of oxygen,
carbon dioxide[339] and ammonia. It absorbs moisture and swells on contact with water. After fertilization, the innermost
portion liquifies to allow free movement of the developing embryo. In certain species, such as the Northern red-legged
frog[340] (Rana aurora) and the wood frog[341] (Rana sylvatica), symbiotic[342] unicellular green algae are present in the
gelatinous material. It is thought that these may benefit the developing larvae by providing them with extra oxygen through
photosynthesis[343].113 Most eggs are black or dark brown and this has the advantage of absorbing warmth from the sun which
the insulating capsule retains. The interior of globular egg clusters of the wood frog[344] (Rana sylvatica) has been found to be
up to 6 C (11 F) warmer than the surrounding water and this speeds up the development of the larvae.114
The shape and size of the egg mass is characteristic of the species. Ranids tend to produce globular clusters containing large
numbers of eggs whereas bufonids produce long, cylindrical strings. The tiny yellow-striped pygmy eleuth[345]
(Eleutherodactylus limbatus) lays eggs singly, burying them in moist soil.115 The smoky jungle frog[346] (Leptodactylus
pentadactylus) makes a nest of foam in a hollow. The eggs hatch when the nest is flooded, or the tadpoles may complete their
development in the foam if flooding does not occur.116 The red-eyed treefrog (Agalychnis callidryas) deposits its eggs on a leaf
above a pool and when they hatch, the larvae fall into the water below.117 The larvae developing in the eggs can detect
vibrations caused by nearby predatory wasps or snakes, and will hatch early to avoid being eaten.118 In general, the length of
the egg stage depends on the species and the environmental conditions. Aquatic eggs normally hatch within one week when
the capsule splits as a result of enzymes[347] released by the developing larvae.119

Tadpoles

[348]

Frogspawn development.
The larvae that emerge from the eggs, known as tadpoles (or occasionally polliwogs), typically have oval bodies and long,
vertically flattened tails. As a general rule, free-living larvae are fully aquatic, but at least one species (Nannophrys
ceylonensis[349]) has semiterrestrial tadpoles which live among wet rocks.120121 Tadpoles lack eyelids and have cartilaginous
skeletons, lateral line systems[350], gills for respiration (external gills at first, internal gills later), and vertically flattened tails
they use for swimming.95
From early in its development, a gill pouch covers the tadpole's gills and front legs. The lungs soon start to develop and are

used as an accessory breathing organ. Some species go through metamorphosis while still inside the egg and hatch directly into
small frogs. Tadpoles lack true teeth, but the jaws in most species have two elongated, parallel rows of small, keratinized[351]
structures called keradonts in their upper jaws. Their lower jaws usually have three rows of keradonts surrounded by a horny
beak, but the number of rows can vary and the exact arrangements of mouth parts provides a means for species
identification.119 In the Pipidae, with the exception of Hymenochirus[352], the tadpoles have paired anterior barbels, which
make them resemble small catfish[353].94 Their tails are stiffened by a notochord[354], but does not contain any bony or
cartilaginous elements except for a few vertebrae at the base which forms the urostyle during metamorphosis. This has been
suggested as an adaptation to their lifestyles; because the transformation into frogs happens very fast, the tail is made of soft
tissue only, as bone and cartilage take a much longer time to be broken down and absorbed. The tail fin and tip is fragile and
will easily tear, which is seen as an adaptation to escape from predators which tries to grasp them by the tail.122
Tadpoles are typically herbivorous[355], feeding mostly on algae[356], including diatoms[357] filtered from the water through the
gills[358]. Some species are carnivorous at the tadpole stage, eating insects, smaller tadpoles, and fish. The Cuban tree frog
(Osteopilus septentrionalis) is one of a number of species in which the tadpoles can be cannibalistic[359]. Tadpoles that develop
legs early may be eaten by the others, so late developers may have better long-term survival prospects.123
Tadpoles are highly vulnerable to being eaten by fish, newts[360], predatory diving beetles[361], and birds, such as
kingfishers[362]. Some tadpoles, including those of the cane toad[363] (Bufo marinus), are poisonous. The tadpole stage may be
as short as a week in explosive breeders or it may last through one or more winters followed by metamorphosis in the
spring.124

Metamorphosis
At the end of the tadpole stage, a frog undergoes metamorphosis in which its body makes a sudden transition into the adult
form. This metamorphosis typically lasts only 24 hours, and is initiated by production of the hormone[364] thyroxine[365]. This
causes different tissues to develop in different ways. The principal changes that take place include the development of the
lungs and the disappearance of the gills and gill pouch, making the front legs visible. The lower jaw transforms into the big
mandible of the carnivorous adult, and the long, spiral gut of the herbivorous tadpole is replaced by the typical short gut of a
predator.119 The nervous system becomes adapted for hearing and stereoscopic vision, and for new methods of locomotion
and feeding.119 The eyes are repositioned higher up on the head and the eyelids and associated glands are formed. The
eardrum, middle ear, and inner ear are developed. The skin becomes thicker and tougher, the lateral line system is lost, and
skin glands are developed.119 The final stage is the disappearance of the tail, but this takes place rather later, the tissue being
used to produce a spurt of growth in the limbs.125 Frogs are at their most vulnerable to predators when they are undergoing
metamorphosis. At this time, the tail is being lost and locomotion by means of limbs is only just becoming established.89

[366]

Larva of the common frog Rana temporaria a day before metamorphosis

[367]

Metamorphosis stage with deforming jaws, large eyes, and remains of gill pouch

[368]

Young frog with a stumpy tail, metamorphosis nearly complete

Adults
After metamorphosis, young adults may disperse into terrestrial habitats or continue to live in water. Almost all species of
frogs are carnivorous[369] as adults, preying on invertebrates, including arthropods[370], worms[371], snails[372], and slugs[373].
A few of the larger ones may eat other frogs, small mammals[374], and fish[375]. Some frogs use their sticky tongues to catch
fast-moving prey, while others push food into their mouths with their hands. A few species also eat plant matter; The tree frog
Xenohyla truncata[376] is partly herbivorous, its diet including a large proportion of fruit,126 Leptodactylus mystaceus[377] has
been found to eat plants,127 and folivory[378] occurs in Euphlyctis hexadactylus[379], with plants constituting 79.5% of its diet
by volume.128 Adult frogs are themselves attacked by many predators. The northern leopard frog[380] (Rana pipiens) is eaten by
herons[381], hawks[382], fish, large salamanders[383], snakes[384], raccoons[385], skunks[386], mink[387], bullfrogs, and other
animals.129

[388]

A trophic pyramid showing frogs as primary predators.

Frogs are primary predators and an important part of the food web[389]. Being cold-blooded[390], they make efficient use of the
food they eat with little energy being used for metabolic processes, while the rest is transformed into biomass[391]. They are
themselves eaten by secondary predators and are the primary terrestrial consumers of invertebrates, most of which feed on
plants. By reducing herbivory, they play a part in increasing the growth of plants and are thus part of a delicately balanced
ecosystem.130
Little is known about the longevity of frogs and toads in the wild, but some can live for many years. Skeletochronology[392] is a
method of examining bones to determine age. Using this method, the ages of mountain yellow-legged frogs[393] (Rana
muscosa) were studied, the phalanges of the toes showing seasonal lines where growth slows in winter. The oldest frogs had ten
bands, so their age was believed to be 14 years, including the four-year tadpole stage.131 Captive frogs and toads have been
recorded as living for up to 40 years, an age achieved by a European common toad (Bufo bufo). The cane toad (Bufo marinus) has
been known to survive 24 years in captivity, and the American bullfrog (Rana catesbeiana) 14 years.132 Frogs from temperate

climates hibernate during the winter, and four species are known to be able to withstand freezing during this time, including
the wood frog (Rana sylvatica).133

Parental care

[394]

Pouched frog[395] (Assa darlingtoni)

Although care of offspring is poorly understood in frogs, up to an estimated 20% of amphibian species may care for their young
in some way.134 The evolution of parental care[396] in frogs is driven primarily by the size of the water body in which they
breed. Those that breed in smaller water bodies tend to have greater and more complex parental care behaviour.135 Because
predation of eggs and larvae is high in large water bodies, a number of frog species started to lay their eggs on land. Once this
happened, the desiccating terrestrial environment demands that one or both parents keep them moist to ensure their
survival.136 The subsequent need to transport hatched tadpoles to a water body required an even more intense form of
parental care.135
In small pools, predators are mostly absent and competition between tadpoles becomes the variable that constrains their
survival. Certain frog species avoid this competition by making use of smaller phytotelmata[397] (water-filled leaf axils[398] or
small woody cavities) as sites for depositing a few tadpoles.137 While these smaller rearing sites are free from competition,
they also lack sufficient nutrients to support a tadpole without parental assistance. Frog species that changed from the use of
larger to smaller phytotelmata have evolved a strategy of providing their offspring with nutritive but unfertilized eggs.135 The
female strawberry poison-dart frog[399] (Oophaga pumilio) lays her eggs on the forest floor. The male frog guards them from
predation and carries water in his cloaca to keep them moist. When they hatch, the female moves the tadpoles on her back to a
water-holding bromeliad[400] or other similar water body, depositing just one in each location. She visits them regularly and
feeds them by laying one or two unfertilized eggs in the phytotelma, continuing to do this until the young are large enough to
undergo metamorphosis.138 The granular poison frog[401] (Oophaga granulifera) looks after its tadpoles in a similar way.139
Many other diverse forms of parental care are seen in frogs. The tiny male Colostethus subpunctatus[402] stands guard over his
egg cluster, laid under a stone or log. When the eggs hatch, he transports the tadpoles on his back to a temporary pool, where
he partially immerses himself in the water and one or more tadpoles drop off. He then moves on to another pool.140 The male
common midwife toad[403] (Alytes obstetricans) carries the eggs around with him attached to his hind legs. He keeps them damp
in dry weather by immersing himself in a pond, and prevents them from getting too wet in soggy vegetation by raising his
hindquarters. After three to six weeks, he travels to a pond and the eggs hatch into tadpoles.141 The tungara frog[404]
(Physalaemus pustulosus) builds a floating nest from foam to protect its eggs from predation. The foam is made from
proteins[405] and lectins[406], and seems to have antimicrobial properties.142 Several pairs of frogs may form a colonial nest on
a previously built raft. The eggs are laid in the centre, followed by alternate layers of foam and eggs, finishing with a foam
capping.143
Some frogs protect their offspring inside their own bodies. Both male and female pouched frogs[407] (Assa darlingtoni) guard
their eggs, which are laid on the ground. When the eggs hatch, the male lubricates his body with the jelly surrounding them
and immerses himself in the egg mass. The tadpoles wriggle into skin pouches on his side, where they develop until they
metamorphose into juvenile frogs.144 The female gastric-brooding frog[408] (Rheobatrachus sp.) from Australia[409], now
probably extinct, swallows her fertilized eggs, which then develop inside her stomach. She ceases to feed and stops secreting
stomach acid[410]. The tadpoles rely on the yolks of the eggs for nourishment. After six or seven weeks, they are ready for
metamorphosis. The mother regurgitates the tiny frogs, which hop away from her mouth.145 The female Darwin's frog[411]

(Rhinoderma darwinii) from Chile[412] lays up to 40 eggs on the ground, where they are guarded by the male. When the tadpoles
are about to hatch, they are engulfed by the male, which carries them around inside his much-enlarged vocal sac. Here they are
immersed in a frothy, viscous liquid that contains some nourishment to supplement what they obtain from the yolks of the
eggs. They remain in the sac for seven to ten weeks before undergoing metamorphosis, after which they move into the male's
mouth and emerge.146

Defence

[413]

At first sight, frogs seem rather defenceless because of their small size, slow movement, thin skin, and lack of defensive
structures, such as spines, claws or teeth. Many use camouflage to avoid detection, the skin often being spotted or streaked in
neutral colours that allow a stationary frog to merge into its surroundings. Some can make prodigious leaps, often into water,
that help them to evade potential attackers, while many have other defensive adaptations and strategies.108
The skin of many frogs contains mild toxic substances called bufotoxins[414] to make them unpalatable to potential predators.
Most toads and some frogs have large poison glands, the parotoid glands[415], located on the sides of their heads behind the
eyes and other glands elsewhere on their bodies. These glands secrete mucus and a range of toxins that make frogs slippery to
hold and distasteful or poisonous. If the noxious effect is immediate, the predator may cease its action and the frog may
escape. If the effect develops more slowly, the predator may learn to avoid that species in future.147 Poisonous frogs tend to
advertise their toxicity with bright colours, an adaptive strategy known as aposematism[416]. The poison dart frogs in the
family Dendrobatidae do this. They are typically red, orange, or yellow, often with contrasting black markings on their bodies.
Allobates zaparo[417] is not poisonous, but mimics the appearance of two different toxic species with which it shares a common
range in an effort to deceive predators.148 Other species, such as the European fire-bellied toad[418] (Bombina bombina), have
their warning colour underneath. They "flash" this when attacked, adopting a pose that exposes the vivid colouring on their
bellies.6

[419]

A common toad adopting a defensive stance.

Some frogs, such as the poison dart frogs[420], are especially toxic. The native people of South America extract poison from
these frogs to apply to their weapons[421] for hunting,149 although few species are toxic enough to be used for this purpose. At
least two non-poisonous species of frogs in tropical America (Eleutherodactylus gaigei and Lithodytes lineatus) mimic[422] the
colouration of dart poison frogs for self-protection.150151 Some frogs obtain poisons from the ants and other arthropods they
eat.152 Others, such as the Australian corroboree frogs[423] (Pseudophryne corroboree and Pseudophryne pengilleyi), can
synthesize the alkaloids[424] themselves.153 The chemicals involved may be irritants, hallucinogens[425], convulsants[426],
nerve poisons[427] or vasoconstrictors[428]. Many predators of frogs have become adapted to tolerate high levels of these
poisons, but other creatures, including humans who handle the frogs, may be severely affected.154

Some frogs use bluff or deception. The European common toad (Bufo bufo) adopts a characteristic stance when attacked,
inflating its body and standing with its hindquarters raised and its head lowered.155 The bullfrog (Rana catesbeiana) crouches
down with eyes closed and head tipped forward when threatened. This places the parotoid glands in the most effective
position, the other glands on its back begin to ooze noxious secretions and the most vulnerable parts of its body are
protected.108 Another tactic used by some frogs is to "scream", the sudden loud noise tending to startle the predator. The gray
tree frog (Hyla versicolor[429]) makes an explosive sound that sometimes repels the shrew Blarina brevicauda[430].108 Although
toads are avoided by many predators, the common garter snake[431] (Thamnophis sirtalis) regularly feeds on them. The strategy
employed by juvenile American toads (Bufo americanus[432]) on being approached by a snake is to crouch down and remain
immobile. This is usually successful, with the snake passing by and the toad remaining undetected. If it is encountered by the
snake's head, however, the toad hops away before crouching defensively.156

Distribution and conservation status

See also: Decline in amphibian populations[433]
Frogs are found on all the continents except Antarctica, but they are not present on certain islands, especially those far away
from continental land masses.157158 Many species are isolated in restricted ranges by changes of climate or inhospitable
territory, such as stretches of sea, mountain ridges, deserts, forest clearance, road construction, or other man-made
barriers.159 Usually, a greater diversity of frogs occurs in tropical areas than in temperate regions, such as Europe.160 Some
frogs inhabit arid areas, such as deserts, and rely on specific adaptations to survive. Members of the Australian genus
Cyclorana[434] bury themselves underground where they create a water-impervious cocoon in which to aestivate[435] during dry
periods. Once it rains, they emerge, find a temporary pool, and breed. Egg and tadpole development is very fast in comparison
to those of most other frogs, so breeding can be completed before the pond dries up.161 Some frog species are adapted to a
cold environment. The wood frog[436] (Rana sylvatica), whose habitat extends into the Arctic Circle[437], buries itself in the
ground during winter. Although much of its body freezes during this time, it maintains a high concentration of glucose in its
vital organs, which protects them from damage.40
In 2006, of 4,035 species of amphibians that depend on water during some lifecycle stage, 1,356 (33.6%) were considered to be
threatened. This is likely to be an underestimate because it excludes 1,427 species for which evidence was insufficient to assess
their status.162 Frog populations have declined dramatically since the 1950s. More than one-third of frog species are
considered to be threatened with extinction[438], and more than 120 species are believed to have become extinct since the
1980s.1 Among these species are the gastric-brooding frogs of Australia and the golden toad[439] of Costa Rica. The latter is of
particular concern to scientists because it inhabited the pristine Monteverde Cloud Forest Reserve[440] and suffered a
population crash in 1987, along with about 20 other frog species found in the area. This could not be linked directly to human
activities, such as deforestation, and was outside the range of normal fluctuations in population size.163 Elsewhere, habitat
loss is a significant cause of frog population decline, as are pollutants, climate change, increased UVB[441] radiation, and the
introduction of non-native[442] predators and competitors.164 A Canadian study conducted in 2006 suggested heavy traffic in
their environment was a larger threat to frog populations than was habitat loss.165 Emerging infectious diseases, including
chytridiomycosis[443] and ranavirus[444], are also devastating populations.166167
Many environmental scientists believe amphibians, including frogs, are good biological indicators[445] of broader
ecosystem[446] health because of their intermediate positions in food chains, their permeable skins, and typically biphasic lives
(aquatic larvae and terrestrial adults).168 It appears that species with both aquatic eggs and larvae are most affected by the
decline, while those with direct development are the most resistant.169

[447]

Deformed frog.
Frog mutations and genetic defects[448] have increased since the 1990s. These often include missing legs or extra legs. Various
causes have been identified or hypothesized, including an increase in ultraviolet radiation[449] affecting the spawn on the
surface of ponds, chemical contamination from pesticides and fertilizers, and parasites such as the trematode[450] Ribeiroia
ondatrae[451]. Probably all these are involved in a complex way as stressors[452], environmental factors contributing to rates of
disease, and vulnerability to attack by parasites. Malformations impair mobility and the individuals may not survive to
adulthood. An increase in the number of frogs eaten by birds may actually increase the likelihood of parasitism of other frogs,
because the trematode's complex lifecycle includes the ramshorn snail[453] and several intermediate hosts such as birds.170171
In a few cases, captive breeding programs have been established and have largely been successful.172173 In 2007, the
application of certain probiotic[454] bacteria was reported to protect amphibians from chytridiomycosis.174 One current
project, the Panama Amphibian Rescue and Conservation Project, has subsequently been developed to rescue species at risk of
this disease in eastern Panama, and to develop field applications for probiotic therapy.175176 The World Association of Zoos
and Aquariums[455] named 2008 as the "Year of the Frog" in order to draw attention to the conservation issues faced by
them.177
The cane toad[456] (Bufo marinus) is a very adaptable species native to South and Central America. In the 1930s, it was
introduced into Puerto Rico, and later various other islands in the Pacific and Caribbean region, as a biological pest control[457]
agent.178 In 1935, 3000 toads were liberated in the sugar cane[458] fields of Queensland, Australia, in an attempt to control
cane beetles[459] such as Dermolepida albohirtum, the larvae of which damage and kill the canes. Initial results in many of these
countries were positive, but it later became apparent that the toads upset the ecological balance in their new environments.
They bred freely, competed with native species of frogs, ate bees and other harmless native invertebrates, had few predators in
their adopted habitats, and poisoned pets, carnivorous birds, and mammals. In many of these countries, they are now regarded
both as pests and invasive species[460], and scientists are looking for a biological method to control them.179
See also: Animal testing on frogs[461]

[462]

Cooked frog, Jianyang, Fujian[463], China

Frog legs[464] are eaten by humans in many parts of the world. Originally, they were supplied from local wild populations, but
overexploitation led to a diminution in the supply. This resulted in the development of frog farming[465] and a global trade in
frogs.180 The main importing countries are France, Belgium, Luxembourg, and the United States, while the chief exporting

nations are Indonesia and China.180 The annual global trade in the American bullfrog[466] (Rana catesbeiana), mostly farmed
in China, varies between 1200 and 2400 tonnes.181
Frogs are sometimes used for dissections[467] in high school and university anatomy classes, often first being injected with
coloured substances to enhance the contrast between the biological systems[468]. This practice is declining with increasing
concerns about animal welfare[469], and "digital frogs" are now available for virtual dissection.182
Frogs have served as experimental animals[470] throughout the history of science. Eighteenth-century biologist Luigi
Galvani[471] discovered the link between electricity[472] and the nervous system[473] through studying frogs.183 In 1852, H. F.
Stannius used a frog's heart in a procedure called a Stannius ligature[474] to demonstrate the ventricle and atria beat
independently of each other and at different rates.184 The African clawed frog[475] or platanna (Xenopus laevis) was first widely
used in laboratories in pregnancy assays in the first half of the 20th century. A sample of urine from a pregnant woman injected
into a female frog induces it to lay eggs[476], a discovery made by English zoologist Lancelot Hogben[477]. This is because a
hormone, human chorionic gonadotropin[478], is present in substantial quantities in the urine of women during pregnancy.185
In 1952, Robert Briggs[479] and Thomas J. King[480] cloned a frog by somatic cell nuclear transfer[481]. This same technique was
later used to create Dolly the sheep[482], and their experiment was the first time a successful nuclear transplantation had been
accomplished in higher animals.186
Frogs are used in cloning research and other branches of embryology[483]. Although alternative pregnancy assays have been
developed, biologists continue to use Xenopus as a model organism[484] in developmental biology[485] because their embryos
are large and easy to manipulate, they are readily obtainable, and can easily be kept in the laboratory.187 Xenopus laevis is
increasingly being displaced by its smaller relative, Xenopus tropicalis[486], which reaches its reproductive age in five months
rather than the one to two years for X. laevis,188 thus facilitating faster studies across generations. The genome[487] of X.
tropicalis is being sequenced.189
Because frog toxins are extraordinarily diverse, they have raised the interest of biochemists as a "natural pharmacy". The
alkaloid epibatidine[488], a painkiller 200 times more potent than morphine[489], is found in some species of poison dart frogs.
Other chemicals isolated from the skins of frogs may offer resistance to HIV[490] infection.190 Dart poisons are under active
investigation for their potential as therapeutic drugs.191
It has long been suspected that pre-Columbian Mesoamericans[491] used a toxic secretion produced by the cane toad as a
hallucinogen[492], but more likely they used substances secreted by the Colorado River toad[493] (Bufo alvarius). These contain
bufotenin[494] (5-MeO-DMT), a psychoactive compound[495] that has been used in modern times as a recreational drug[496].
Typically, the skin secretions are dried and then smoked.192 Illicit drug use by licking the skin of a toad has been reported in
the media, but this may be an urban myth[497].193

[498]

Golden poison frog[499] (Phyllobates terribilis).

Exudations from the skin of the golden poison frog[500] (Phyllobates terribilis) are traditionally used by native Colombians to
poison the darts they use for hunting. The tip of the projectile is rubbed over the back of the frog and the dart is launched from
a blowgun[501]. The combination of the two alkaloid toxins batrachotoxin[502] and homobatrachotoxin[503] is so powerful, one

frog contains enough poison to kill an estimated 22,000 mice.194 Two other species, the Kokoe poison dart frog[504]
(Phyllobates aurotaenia) and the black-legged dart frog[505] (Phyllobates bicolor) are also used for this purpose. These are less
toxic and less abundant than the golden poison frog. They are impaled on pointed sticks and may be heated over a fire to
maximise the quantity of poison that can be transferred to the dart.194

Cultural beliefs
For more details on this topic, see Frogs in popular culture[506].
Frogs feature prominently in folklore[507], fairy tales[508], and popular culture. They tend to be portrayed as benign, ugly, and
clumsy, but with hidden talents. Examples include Michigan J. Frog[509], "The Frog Prince"[510], and Kermit the Frog[511]. The
Warner Brothers[512] cartoon One Froggy Evening[513] features Michigan J. Frog, that will only dance and sing for the demolition
worker who opens his time capsule, but will not perform in public.195 "The Frog Prince" is a fairy tale about a frog that turns
into a handsome prince after he has rescued a princess's golden ball and she has taken him into her palace.196 Kermit the Frog
is a conscientious and disciplined character from The Muppet Show[514] and Sesame Street[515]; while openly friendly and
greatly talented, he is often portrayed as cringing at the fanciful behavior of more flamboyant characters.197
Toads have a more sinister reputation. It was believed in European folklore that they were associated with witches[516] as their
familiar spirits[517] and had magical powers. The toxic secretions from their skin was used in brewing evil potions, but was also
put to use to create magical cures for human and livestock ailments. They were associated with the devil; in John Milton[518]'s
Paradise Lost[519], Satan[520] was depicted as a toad pouring poison into Eve[521]'s ear.198
The Moche[522] people of ancient Peru[523] worshipped animals, and often depicted frogs in their art.199 In Panama[524], local
legend held that good fortune would come to anyone who spotted a Panamanian golden frog. Some believed when one of these
frogs died, it would turn into a golden talisman[525] known as a huaca[526]. Today, despite being extinct in the wild,
Panamanian golden frogs remain an important cultural symbol and can be found on decorative cloth molas[527] made by the
Kuna people[528]. They also appear as part of the inlaid design on a new overpass in Panama City[529], on T-shirts, and even on
lottery tickets[530].200

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External links
Media
[show]
Extant anuran families by suborder
[show]
Extant amphibian[1001] positions by subclass

Links
1. http://en.wikipedia.org/w/index.php?title=Frogs&redirect=no
2. http://en.wikipedia.org/wiki/Frog_(disambiguation)
3. http://en.wikipedia.org/wiki/Carnivore
4. http://en.wikipedia.org/wiki/Amphibian

5. http://en.wikipedia.org/wiki/Order_(biology)
6. http://en.wikipedia.org/wiki/Ancient_Greek
7. http://en.wikipedia.org/wiki/Triadobatrachus
8. http://en.wikipedia.org/wiki/Early_Triassic
9. http://en.wikipedia.org/wiki/Madagascar
10. http://en.wikipedia.org/wiki/Molecular_clock
11. http://en.wikipedia.org/wiki/Permian
12. http://en.wikipedia.org/wiki/Tropics
13. http://en.wikipedia.org/wiki/Subarctic
14. http://en.wikipedia.org/wiki/Species_diversity
15. http://en.wikipedia.org/wiki/Tropical_rainforest
16. http://en.wikipedia.org/wiki/Vertebrate
17. http://en.wikipedia.org/wiki/Body_plan
18. http://en.wikipedia.org/wiki/Eye
19. http://en.wikipedia.org/wiki/Tongue
20. http://en.wikipedia.org/wiki/Tail
21. http://en.wikipedia.org/wiki/Gland
22. http://en.wikipedia.org/wiki/Toad
23. http://en.wikipedia.org/wiki/Taxonomy_(biology)
24. http://en.wikipedia.org/wiki/Camouflage
25. http://en.wikipedia.org/wiki/Aposematism
26. http://en.wikipedia.org/wiki/Egg_(biology)
27. http://en.wikipedia.org/wiki/Larva
28. http://en.wikipedia.org/wiki/Tadpole
29. http://en.wikipedia.org/wiki/Gill
30. http://en.wikipedia.org/wiki/Herbivore
31. http://en.wikipedia.org/wiki/Omnivore
32. http://en.wikipedia.org/wiki/Planktivore
33. http://en.wikipedia.org/wiki/Biological_life_cycle
34. http://en.wikipedia.org/wiki/Metamorphosis
35. http://en.wikipedia.org/wiki/Predation
36. http://en.wikipedia.org/wiki/Keystone_species
37. http://en.wikipedia.org/wiki/Food_web
38. http://en.wikipedia.org/wiki/Ecosystem
39. http://en.wikipedia.org/wiki/Semi-permeable
40. http://en.wikipedia.org/wiki/Animal_communication
41. http://en.wikipedia.org/wiki/Breeding_season
42. http://en.wikipedia.org/wiki/Ethology
43. http://en.wikipedia.org/wiki/Threatened_species
44. http://en.wikipedia.org/wiki/Extinct

45. http://en.wikipedia.org/wiki/Fungus
46. http://en.wikipedia.org/wiki/Chytridiomycosis
47. http://en.wikipedia.org/wiki/Conservation_biology
48. http://en.wikipedia.org/wiki/Old_English
49. http://en.wikipedia.org/wiki/Proto-Indo-European_language
50. http://en.wikipedia.org/wiki/Amphibian
51. http://en.wikipedia.org/wiki/Biological_classification
52. http://en.wikipedia.org/wiki/Order_(biology)
53. http://en.wikipedia.org/wiki/Family_(biology)
54. http://en.wikipedia.org/wiki/Leptodactylidae
55. http://en.wikipedia.org/wiki/Hylidae
56. http://en.wikipedia.org/wiki/Ranidae
57. http://en.wikipedia.org/wiki/Species_richness
58. http://en.wikipedia.org/wiki/File:Bombina_bombina_1_(Marek_Szczepanek)_tight_crop.jpg
59. http://en.wikipedia.org/wiki/Toad
60. http://en.wikipedia.org/wiki/Bufonidae
61. http://en.wikipedia.org/wiki/European_fire-bellied_toad
62. http://en.wikipedia.org/wiki/Panamanian_golden_frog
63. http://en.wikipedia.org/wiki/Fossil
64. http://en.wikipedia.org/wiki/Coccyx
65. http://en.wikipedia.org/wiki/Radius_(bone)
66. http://en.wikipedia.org/wiki/Ulna
67. http://en.wikipedia.org/wiki/Tibia
68. http://en.wikipedia.org/wiki/Fibula
69. http://en.wikipedia.org/wiki/Ankle_bone
70. http://en.wikipedia.org/wiki/Hyoid_bone
71. http://en.wikipedia.org/wiki/Mandible
72. http://en.wikipedia.org/wiki/Gastrotheca_guentheri
73. http://en.wikipedia.org/wiki/Pipoidea
74. http://en.wikipedia.org/wiki/Lymph
75. http://en.wikipedia.org/wiki/Lens_(anatomy)
76. http://en.wikipedia.org/wiki/Spiracle
77. http://en.wikipedia.org/wiki/Keratin
78. http://en.wikipedia.org/wiki/Beak
79. http://en.wikipedia.org/wiki/Denticle
80. http://en.wikipedia.org/wiki/File:Atelopus_zeteki1.jpg
81. http://en.wikipedia.org/wiki/Archaeobatrachia
82. http://en.wikipedia.org/wiki/Mesobatrachia
83. http://en.wikipedia.org/wiki/Neobatrachia
84. http://en.wikipedia.org/wiki/Hyloidea

85. http://en.wikipedia.org/wiki/Ranoidea
86. http://en.wikipedia.org/wiki/Morphology_(biology)
87. http://en.wikipedia.org/wiki/Pectoral_girdle
88. http://en.wikipedia.org/wiki/Hybrid_(biology)
89. http://en.wikipedia.org/wiki/Edible_frog
90. http://en.wikipedia.org/wiki/Pool_frog
91. http://en.wikipedia.org/wiki/Marsh_frog
92. http://en.wikipedia.org/wiki/Yellow-bellied_toad
93. http://en.wikipedia.org/wiki/Hybrid_zone
94. http://en.wikipedia.org/wiki/Molecular_phylogeny
95. http://en.wikipedia.org/wiki/Ribosomal_DNA
96. http://en.wikipedia.org/wiki/Salamanders
97. http://en.wikipedia.org/wiki/Caecilian
98. http://en.wikipedia.org/wiki/Genetic_divergence
99. http://en.wikipedia.org/wiki/Paleozoic
100. http://en.wikipedia.org/wiki/Mesozoic
101. http://en.wikipedia.org/wiki/Pangaea
102. http://en.wikipedia.org/wiki/Sarcopterygii
103. http://en.wikipedia.org/wiki/Lissamphibian
104. http://en.wikipedia.org/wiki/Temnospondyli
105. http://en.wikipedia.org/wiki/Neobatrachia
106. http://en.wikipedia.org/wiki/Permian
107. http://en.wikipedia.org/wiki/Monophyly
108. http://en.wikipedia.org/wiki/Lepospondyli
109. http://en.wikipedia.org/wiki/Temnospondyli
110. http://en.wikipedia.org/wiki/Carboniferous
111. http://en.wikipedia.org/wiki/Caudata
112. http://en.wikipedia.org/wiki/Gerobatrachus_hottoni
113. http://en.wikipedia.org/wiki/Temnospondyli
114. http://en.wikipedia.org/wiki/Texas
115. http://en.wikipedia.org/wiki/Transitional_fossil
116. http://en.wikipedia.org/wiki/Stem_group
117. http://en.wikipedia.org/wiki/Common_ancestor
118. http://en.wikipedia.org/wiki/Clade
119. http://en.wikipedia.org/wiki/Gerobatrachus_hottoni
120. http://en.wikipedia.org/wiki/Dissorophoidea
121. http://en.wikipedia.org/wiki/Triadobatrachus
122. http://en.wikipedia.org/wiki/Triassic
123. http://en.wikipedia.org/wiki/Madagascar
124. http://en.wikipedia.org/wiki/Czatkobatrachus

125. http://en.wikipedia.org/wiki/Poland
126. http://en.wikipedia.org/wiki/Vertebrae
127. http://en.wikipedia.org/wiki/Stem_group
128. http://en.wikipedia.org/wiki/Triadobatrachus
129. http://en.wikipedia.org/wiki/Czatkobatrachus
130. http://en.wikipedia.org/wiki/Salientia
131. http://en.wikipedia.org/wiki/Vertebral_column
132. http://en.wikipedia.org/wiki/Ilium_(bone)
133. http://en.wikipedia.org/wiki/Pelvis
134. http://en.wikipedia.org/wiki/Parietal_bone
135. http://en.wikipedia.org/wiki/Mandible
136. http://en.wikipedia.org/wiki/Prosalirus_bitis
137. http://en.wikipedia.org/wiki/Early_Jurassic
138. http://en.wikipedia.org/wiki/1995_in_paleontology
139. http://en.wikipedia.org/wiki/Kayenta_Formation
140. http://en.wikipedia.org/wiki/Arizona
141. http://en.wikipedia.org/wiki/Early_Jurassic
142. http://en.wikipedia.org/wiki/Pelvic
143. http://en.wikipedia.org/wiki/Vieraella_herbsti
144. http://en.wikipedia.org/wiki/Dorsum_(biology)
145. http://en.wikipedia.org/wiki/Ventral
146. http://en.wikipedia.org/wiki/Notobatrachus_degiustoi
147. http://en.wikipedia.org/wiki/Ploidy
148. http://en.wikipedia.org/wiki/Speciation
149. http://en.wikipedia.org/wiki/Antarctica
150. http://en.wikipedia.org/wiki/Biogeography
151. http://en.wikipedia.org/wiki/Cladogram
152. http://en.wikipedia.org/wiki/Family_(biology)
153. http://en.wikipedia.org/wiki/Phylogenetic_tree
154. http://en.wikipedia.org/wiki/Oxygen
155. http://en.wikipedia.org/wiki/Buccal_pumping
156. http://en.wikipedia.org/wiki/Decline_in_amphibian_populations
157. http://en.wikipedia.org/wiki/Paedophryne_amauensis
158. http://en.wikipedia.org/wiki/Papua_New_Guinea
159. http://en.wikipedia.org/wiki/Goliath_frog
160. http://en.wikipedia.org/wiki/Cameroon
161. http://en.wikipedia.org/wiki/Loose_connective_tissue
162. http://en.wikipedia.org/wiki/Tympanum_(zoology)
163. http://en.wikipedia.org/wiki/Pedicellate_teeth
164. http://en.wikipedia.org/wiki/Vomerine_teeth

165. http://en.wikipedia.org/wiki/Pyxicephalus
166. http://en.wikipedia.org/wiki/Musculoskeletal
167. http://en.wikipedia.org/wiki/Morphology_(biology)
168. http://en.wikipedia.org/wiki/Tarsus_(skeleton)
169. http://en.wikipedia.org/wiki/Bone
170. http://en.wikipedia.org/wiki/Metatarsals
171. http://en.wikipedia.org/wiki/Illium
172. http://en.wikipedia.org/wiki/Sacrum
173. http://en.wikipedia.org/wiki/File:Rana_temporaria_04_by-dpc.jpg
174. http://en.wikipedia.org/wiki/African_dwarf_frog
175. http://en.wikipedia.org/wiki/White%27s_tree_frog
176. http://en.wikipedia.org/wiki/Tree_frog
177. http://en.wikipedia.org/wiki/Capillarity
178. http://en.wikipedia.org/wiki/Substrate_(biology)
179. http://en.wikipedia.org/wiki/Couch%27s_Spadefoot_Toad
180. http://en.wikipedia.org/wiki/Keratin
181. http://en.wikipedia.org/wiki/Tubercle
182. http://en.wikipedia.org/wiki/Dragonfly_nymph
183. http://en.wikipedia.org/wiki/Flatworm
184. http://en.wikipedia.org/wiki/Ribeiroia_ondatrae
185. http://en.wikipedia.org/wiki/Grey_Foam-nest_Treefrog
186. http://en.wikipedia.org/wiki/Amplexus
187. http://en.wikipedia.org/wiki/Nocturnal
188. http://en.wikipedia.org/wiki/Woodhouse%27s_toad
189. http://en.wikipedia.org/wiki/Hairy_Frog
190. http://en.wikipedia.org/wiki/Dermal_papillae
191. http://en.wikipedia.org/wiki/Osteoderm
192. http://en.wikipedia.org/wiki/Exostosis
193. http://en.wikipedia.org/wiki/Camouflage
194. http://en.wikipedia.org/wiki/Category:Animals_that_can_change_color
195. http://en.wikipedia.org/wiki/Pacific_tree_frog
196. http://en.wikipedia.org/wiki/Skin_fold
197. http://en.wikipedia.org/wiki/File:Hip-pocket_Frog_-_Assa_darlingtoni.jpg
198. http://en.wikipedia.org/wiki/Pouched_frog
199. http://en.wikipedia.org/wiki/Oxygen
200. http://en.wikipedia.org/wiki/Carbon_dioxide
201. http://en.wikipedia.org/wiki/Buccal_pumping
202. http://en.wikipedia.org/wiki/Rib
203. http://en.wikipedia.org/wiki/Diaphragm_(anatomy)
204. http://en.wikipedia.org/wiki/Bornean_Flat-headed_Frog

205. http://en.wikipedia.org/wiki/Heart
206. http://en.wikipedia.org/wiki/Lizard
207. http://en.wikipedia.org/wiki/Respiration_(physiology)
208. http://en.wikipedia.org/wiki/Atrium_(anatomy)
209. http://en.wikipedia.org/wiki/Ventricle_(heart)
210. http://en.wikipedia.org/wiki/Aorta
211. http://en.wikipedia.org/wiki/Pulmonary_artery
212. http://en.wikipedia.org/wiki/Lake_Titicaca
213. http://en.wikipedia.org/wiki/Telmatobius_culeus
214. http://en.wikipedia.org/wiki/File:Frog_anatomy_tags.PNG
215. http://en.wikipedia.org/wiki/Cloaca
216. http://en.wikipedia.org/wiki/Kidney
217. http://en.wikipedia.org/wiki/Ammonia
218. http://en.wikipedia.org/wiki/Urea
219. http://en.wikipedia.org/wiki/Uric_acid
220. http://en.wikipedia.org/wiki/Ureter
221. http://en.wikipedia.org/wiki/Urinary_bladder
222. http://en.wikipedia.org/wiki/Testis
223. http://en.wikipedia.org/wiki/Semen
224. http://en.wikipedia.org/wiki/Efferent_ducts
225. http://en.wikipedia.org/wiki/Amplexus
226. http://en.wikipedia.org/wiki/Secondary_sexual_characteristic
227. http://en.wikipedia.org/wiki/Vocal_sac
228. http://en.wikipedia.org/wiki/Cerebellum
229. http://en.wikipedia.org/wiki/Medulla_oblongata
230. http://en.wikipedia.org/wiki/Cerebrum
231. http://en.wikipedia.org/wiki/Cranial_nerves
232. http://en.wikipedia.org/wiki/Spinal_nerves
233. http://en.wikipedia.org/wiki/Amniotes
234. http://en.wikipedia.org/wiki/File:Groene_kikker_achter_Bekaert-draad-detail_oog.jpg
235. http://en.wikipedia.org/wiki/Binocular_vision
236. http://en.wikipedia.org/wiki/Nictitating_membrane
237. http://en.wikipedia.org/wiki/Pipidae
238. http://en.wikipedia.org/wiki/Iris_(anatomy)
239. http://en.wikipedia.org/wiki/Common_toad
240. http://en.wikipedia.org/wiki/Agalychnis_callidryas
241. http://en.wikipedia.org/wiki/Phyllobates
242. http://en.wikipedia.org/wiki/Fire-bellied_toad
243. http://en.wikipedia.org/wiki/Dyscophus_antongilii
244. http://en.wikipedia.org/wiki/Southern_toad

245. http://en.wikipedia.org/wiki/Tympanum_(anatomy)
246. http://en.wikipedia.org/wiki/Electrical_tuning
247. http://en.wikipedia.org/wiki/Conspecificity
248. http://en.wikipedia.org/wiki/Sexual_selection_in_frogs
249. http://en.wikipedia.org/wiki/Larynx
250. http://en.wikipedia.org/wiki/Heleioporus
251. http://en.wikipedia.org/wiki/Neobatrachus
252. http://en.wikipedia.org/wiki/Mouth
253. http://en.wikipedia.org/wiki/Resonance_chamber
254. http://en.wikipedia.org/wiki/Ascaphus_truei
255. http://en.wikipedia.org/wiki/Common_Tree_Frog
256. http://en.wikipedia.org/wiki/American_bullfrog
257. http://en.wikipedia.org/wiki/Pacific_Tree_Frog
258. http://en.wikipedia.org/wiki/Onomatopoeia
259. http://en.wikipedia.org/wiki/Pelophylax_ridibundus
260. http://en.wikipedia.org/wiki/Aristophanes
261. http://en.wikipedia.org/wiki/Torpor
262. http://en.wikipedia.org/wiki/Hibernation
263. http://en.wikipedia.org/wiki/American_toad
264. http://en.wikipedia.org/wiki/Hibernaculum_(zoology)
265. http://en.wikipedia.org/wiki/Dormancy
266. http://en.wikipedia.org/wiki/American_bullfrog
267. http://en.wikipedia.org/wiki/Striped_burrowing_frog
268. http://en.wikipedia.org/wiki/Aestivation
269. http://en.wikipedia.org/wiki/Cocoon_(silk)
270. http://en.wikipedia.org/wiki/University_of_Queensland
271. http://en.wikipedia.org/wiki/Metabolism
272. http://en.wikipedia.org/wiki/Mitochondria
273. http://en.wikipedia.org/wiki/Jumping
274. http://en.wikipedia.org/wiki/Running
275. http://en.wikipedia.org/wiki/Walking
276. http://en.wikipedia.org/wiki/Aquatic_locomotion
277. http://en.wikipedia.org/wiki/Burrow
278. http://en.wikipedia.org/wiki/Climbing
279. http://en.wikipedia.org/wiki/Flying_frog
280. http://en.wikipedia.org/wiki/File:Colostethus_flotator_jumping.jpg
281. http://en.wikipedia.org/wiki/Litoria_nasuta
282. http://en.wikipedia.org/wiki/Euphlyctis_cyanophlyctis
283. http://en.wikipedia.org/wiki/Acris_crepitans
284. http://en.wikipedia.org/wiki/Cuban_tree_frog

285. http://en.wikipedia.org/wiki/Northern_Leopard_Frog
286. http://en.wikipedia.org/wiki/Catapult
287. http://en.wikipedia.org/wiki/Locust
288. http://en.wikipedia.org/wiki/Grasshopper
289. http://en.wikipedia.org/wiki/File:Phrynosequence_bodyFoR2.gif
290. http://en.wikipedia.org/wiki/Rhinophrynidae
291. http://en.wikipedia.org/wiki/Microhylidae
292. http://en.wikipedia.org/wiki/Gastrophryne_olivacea
293. http://en.wikipedia.org/wiki/Bufo_fowleri
294. http://en.wikipedia.org/wiki/Kassina_maculata
295. http://en.wikipedia.org/wiki/File:Frog_%C5%BE%C3%A1ba.gif
296. http://en.wikipedia.org/wiki/Common_toad
297. http://en.wikipedia.org/wiki/Pipidae
298. http://en.wikipedia.org/wiki/Lateral_line
299. http://en.wikipedia.org/wiki/Purple_frog
300. http://en.wikipedia.org/wiki/Termite
301. http://en.wikipedia.org/wiki/Monsoon
302. http://en.wikipedia.org/wiki/Fossorial
303. http://en.wikipedia.org/wiki/Species_description
304. http://en.wikipedia.org/wiki/File:Nasikabatrachus_sahyadrensis.jpg
305. http://en.wikipedia.org/wiki/Plains_Spadefoot_Toad
306. http://en.wikipedia.org/wiki/Metatarsus
307. http://en.wikipedia.org/wiki/Osmosis
308. http://en.wikipedia.org/wiki/Western_Spotted_Frog
309. http://en.wikipedia.org/wiki/Scaphiophryne_marmorata
310. http://en.wikipedia.org/wiki/File:Phyllomedusa_burmeisteri01.jpg
311. http://en.wikipedia.org/wiki/Canopy_(biology)
312. http://en.wikipedia.org/wiki/Convergent_evolution
313. http://en.wikipedia.org/wiki/Glass_frogs
314. http://en.wikipedia.org/wiki/Hyperoliidae
315. http://en.wikipedia.org/wiki/Rhacophoridae
316. http://en.wikipedia.org/wiki/Epidermis_(skin)
317. http://en.wikipedia.org/wiki/Mucus
318. http://en.wikipedia.org/wiki/Boundary_friction
319. http://en.wikipedia.org/wiki/Surface_tension
320. http://en.wikipedia.org/wiki/Viscosity
321. http://en.wikipedia.org/wiki/Phyllomedusinae
322. http://en.wikipedia.org/wiki/Phyllomedusa_ayeaye
323. http://en.wikipedia.org/wiki/Digit_(anatomy)
324. http://en.wikipedia.org/wiki/Rhacophorus_nigropalmatus

325. http://en.wikipedia.org/wiki/Tadpole
326. http://en.wikipedia.org/wiki/Metamorphosis
327. http://en.wikipedia.org/wiki/Scaphiopus
328. http://en.wikipedia.org/wiki/New_Mexico_Spadefoot_Toad
329. http://en.wikipedia.org/wiki/External_fertilization
330. http://en.wikipedia.org/wiki/Great_Plains_toad
331. http://en.wikipedia.org/wiki/Nimbaphrynoides
332. http://en.wikipedia.org/wiki/Vivipary
333. http://en.wikipedia.org/wiki/Tanzania
334. http://en.wikipedia.org/wiki/Nectophrynoides
335. http://en.wikipedia.org/wiki/Ovoviviparity
336. http://en.wikipedia.org/wiki/Internal_fertilization
337. http://en.wikipedia.org/wiki/File:Frogspawn_closeup.jpg
338. http://en.wiktionary.org/wiki/frogspawn
339. http://en.wikipedia.org/wiki/Carbon_dioxide
340. http://en.wikipedia.org/wiki/Northern_red-legged_frog
341. http://en.wikipedia.org/wiki/Wood_frog
342. http://en.wikipedia.org/wiki/Symbiosis
343. http://en.wikipedia.org/wiki/Photosynthesis
344. http://en.wikipedia.org/wiki/Wood_Frog
345. http://en.wikipedia.org/wiki/Yellow-Striped_Pygmy_Eleuth
346. http://en.wikipedia.org/wiki/Smoky_Jungle_Frog
347. http://en.wikipedia.org/wiki/Enzyme
348. http://en.wikipedia.org/wiki/File:Frog_spawn_time-lapse.gif
349. http://en.wikipedia.org/wiki/Nannophrys_ceylonensis
350. http://en.wikipedia.org/wiki/Lateral_line_system
351. http://en.wikipedia.org/wiki/Keratin
352. http://en.wikipedia.org/wiki/Hymenochirus
353. http://en.wikipedia.org/wiki/Catfish
354. http://en.wikipedia.org/wiki/Notochord
355. http://en.wikipedia.org/wiki/Herbivore
356. http://en.wikipedia.org/wiki/Alga
357. http://en.wikipedia.org/wiki/Diatom
358. http://en.wikipedia.org/wiki/Gill
359. http://en.wikipedia.org/wiki/Cannibalism
360. http://en.wikipedia.org/wiki/Newt
361. http://en.wikipedia.org/wiki/Diving_beetle
362. http://en.wikipedia.org/wiki/Kingfisher
363. http://en.wikipedia.org/wiki/Cane_toad
364. http://en.wikipedia.org/wiki/Hormone

365. http://en.wikipedia.org/wiki/Thyroxine
366. http://en.wikipedia.org/wiki/File:RanaTemporariaLarva2.jpg
367. http://en.wikipedia.org/wiki/File:RanaTemporariaLarvaFinalStage.JPG
368. http://en.wikipedia.org/wiki/File:Juvenile_Frog_with_tail_top_view_(1).JPG
369. http://en.wikipedia.org/wiki/Carnivore
370. http://en.wikipedia.org/wiki/Arthropod
371. http://en.wikipedia.org/wiki/Annelid
372. http://en.wikipedia.org/wiki/Gastropoda
373. http://en.wikipedia.org/wiki/Slug
374. http://en.wikipedia.org/wiki/Mammal
375. http://en.wikipedia.org/wiki/Fish
376. http://en.wikipedia.org/wiki/Xenohyla_truncata
377. http://en.wikipedia.org/wiki/Leptodactylus_mystaceus
378. http://en.wikipedia.org/wiki/Folivory
379. http://en.wikipedia.org/wiki/Euphlyctis_hexadactylus
380. http://en.wikipedia.org/wiki/Northern_leopard_frog
381. http://en.wikipedia.org/wiki/Heron
382. http://en.wikipedia.org/wiki/Hawk
383. http://en.wikipedia.org/wiki/Salamanders
384. http://en.wikipedia.org/wiki/Snake
385. http://en.wikipedia.org/wiki/Raccoon
386. http://en.wikipedia.org/wiki/Skunk
387. http://en.wikipedia.org/wiki/Mink
388. http://en.wikipedia.org/wiki/File:TrophicWeb.jpg
389. http://en.wikipedia.org/wiki/Food_web
390. http://en.wikipedia.org/wiki/Ectotherm
391. http://en.wikipedia.org/wiki/Biomass_(ecology)
392. http://en.wikipedia.org/wiki/Skeletochronology
393. http://en.wikipedia.org/wiki/Mountain_yellow-legged_frog
394. http://en.wikipedia.org/wiki/File:Assa_darlingtoni.jpg
395. http://en.wikipedia.org/wiki/Pouched_frog
396. http://en.wikipedia.org/wiki/Evolution_of_biparental_care_in_tropical_frogs
397. http://en.wikipedia.org/wiki/Phytotelma
398. http://en.wiktionary.org/wiki/axil
399. http://en.wikipedia.org/wiki/Strawberry_poison-dart_frog
400. http://en.wikipedia.org/wiki/Bromeliad
401. http://en.wikipedia.org/wiki/Granular_poison_frog
402. http://en.wikipedia.org/wiki/Colostethus_subpunctatus
403. http://en.wikipedia.org/wiki/Common_midwife_toad
404. http://en.wikipedia.org/wiki/Tungara_frog

405. http://en.wikipedia.org/wiki/Protein
406. http://en.wikipedia.org/wiki/Lectin
407. http://en.wikipedia.org/wiki/Pouched_frog
408. http://en.wikipedia.org/wiki/Gastric-brooding_frog
409. http://en.wikipedia.org/wiki/Australia
410. http://en.wikipedia.org/wiki/Stomach_acid
411. http://en.wikipedia.org/wiki/Darwin%27s_frog
412. http://en.wikipedia.org/wiki/Chile
413. http://en.wikipedia.org/wiki/File:R._imitator_Chazuta.jpg
414. http://en.wikipedia.org/wiki/Bufotoxin
415. http://en.wikipedia.org/wiki/Parotoid_gland
416. http://en.wikipedia.org/wiki/Aposematism
417. http://en.wikipedia.org/wiki/Allobates_zaparo
418. http://en.wikipedia.org/wiki/European_Fire-bellied_Toad
419. http://en.wikipedia.org/wiki/File:Bufo_bufo-defensive_reaction1.JPG
420. http://en.wikipedia.org/wiki/Poison_dart_frog
421. http://en.wikipedia.org/wiki/Dart_(missile)
422. http://en.wikipedia.org/wiki/Batesian_mimicry
423. http://en.wikipedia.org/wiki/Corroboree_frog
424. http://en.wikipedia.org/wiki/Alkaloid
425. http://en.wikipedia.org/wiki/Hallucinogen
426. http://en.wikipedia.org/wiki/Seizure
427. http://en.wikipedia.org/wiki/Neurotoxin
428. http://en.wikipedia.org/wiki/Vasoconstrictor
429. http://en.wikipedia.org/wiki/Hyla_versicolor
430. http://en.wikipedia.org/wiki/Blarina_brevicauda
431. http://en.wikipedia.org/wiki/Common_Garter_Snake
432. http://en.wikipedia.org/wiki/Bufo_americanus
433. http://en.wikipedia.org/wiki/Decline_in_amphibian_populations
434. http://en.wikipedia.org/wiki/Cyclorana
435. http://en.wikipedia.org/wiki/Aestivation
436. http://en.wikipedia.org/wiki/Wood_frog
437. http://en.wikipedia.org/wiki/Arctic_Circle
438. http://en.wikipedia.org/wiki/Extinction
439. http://en.wikipedia.org/wiki/Golden_toad
440. http://en.wikipedia.org/wiki/Monteverde_Cloud_Forest_Reserve
441. http://en.wikipedia.org/wiki/Ultraviolet
442. http://en.wikipedia.org/wiki/Indigenous_(ecology)
443. http://en.wikipedia.org/wiki/Chytridiomycosis
444. http://en.wikipedia.org/wiki/Ranavirus

445. http://en.wikipedia.org/wiki/Indicator_species
446. http://en.wikipedia.org/wiki/Ecosystem
447. http://en.wikipedia.org/wiki/File:Deformed_Frog.gif
448. http://en.wikipedia.org/wiki/Frog_mutations_and_genetic_defects
449. http://en.wikipedia.org/wiki/Ultraviolet
450. http://en.wikipedia.org/wiki/Trematode
451. http://en.wikipedia.org/wiki/Ribeiroia_ondatrae
452. http://en.wikipedia.org/wiki/Stressor
453. http://en.wikipedia.org/wiki/Planorbidae
454. http://en.wikipedia.org/wiki/Probiotic
455. http://en.wikipedia.org/wiki/World_Association_of_Zoos_and_Aquariums
456. http://en.wikipedia.org/wiki/Cane_toad
457. http://en.wikipedia.org/wiki/Biological_pest_control
458. http://en.wikipedia.org/wiki/Sugar_cane
459. http://en.wikipedia.org/wiki/Cane_beetle
460. http://en.wikipedia.org/wiki/Invasive_species
461. http://en.wikipedia.org/wiki/Animal_testing_on_frogs
462. http://en.wikipedia.org/wiki/File:Cooked_Frog_2.jpg
463. http://en.wikipedia.org/wiki/Jianyang,_Fujian
464. http://en.wikipedia.org/wiki/Frog_legs
465. http://en.wikipedia.org/wiki/Aquaculture
466. http://en.wikipedia.org/wiki/American_bullfrog
467. http://en.wikipedia.org/wiki/Dissection
468. http://en.wikipedia.org/wiki/Biological_system
469. http://en.wikipedia.org/wiki/Animal_welfare
470. http://en.wikipedia.org/wiki/Animal_testing
471. http://en.wikipedia.org/wiki/Luigi_Galvani
472. http://en.wikipedia.org/wiki/Electricity
473. http://en.wikipedia.org/wiki/Nervous_system
474. http://en.wikipedia.org/wiki/Stannius_ligature
475. http://en.wikipedia.org/wiki/African_clawed_frog
476. http://en.wikipedia.org/wiki/Egg_(biology)
477. http://en.wikipedia.org/wiki/Lancelot_Hogben
478. http://en.wikipedia.org/wiki/Human_chorionic_gonadotropin
479. http://en.wikipedia.org/wiki/Robert_William_Briggs
480. http://en.wikipedia.org/wiki/Thomas_J._King
481. http://en.wikipedia.org/wiki/Somatic_cell_nuclear_transfer
482. http://en.wikipedia.org/wiki/Dolly_(sheep)
483. http://en.wikipedia.org/wiki/Embryology
484. http://en.wikipedia.org/wiki/Model_organism

485. http://en.wikipedia.org/wiki/Developmental_biology
486. http://en.wikipedia.org/wiki/Xenopus_tropicalis
487. http://en.wikipedia.org/wiki/Genome
488. http://en.wikipedia.org/wiki/Epibatidine
489. http://en.wikipedia.org/wiki/Morphine
490. http://en.wikipedia.org/wiki/HIV
491. http://en.wikipedia.org/wiki/Mesoamerica
492. http://en.wikipedia.org/wiki/Hallucinogen
493. http://en.wikipedia.org/wiki/Colorado_River_toad
494. http://en.wikipedia.org/wiki/Bufotenin
495. http://en.wikipedia.org/wiki/Psychoactive_drug
496. http://en.wikipedia.org/wiki/Recreational_drug
497. http://en.wikipedia.org/wiki/Urban_myth
498. http://en.wikipedia.org/wiki/File:Schrecklicherpfeilgiftfrosch-01.jpg
499. http://en.wikipedia.org/wiki/Golden_poison_frog
500. http://en.wikipedia.org/wiki/Golden_poison_frog
501. http://en.wikipedia.org/wiki/Blowgun
502. http://en.wikipedia.org/wiki/Batrachotoxin
503. http://en.wikipedia.org/wiki/Homobatrachotoxin
504. http://en.wikipedia.org/wiki/Kokoe_poison_dart_frog
505. http://en.wikipedia.org/wiki/Black-legged_dart_frog
506. http://en.wikipedia.org/wiki/Frogs_in_popular_culture
507. http://en.wikipedia.org/wiki/Folklore
508. http://en.wikipedia.org/wiki/Fairy_tale
509. http://en.wikipedia.org/wiki/Michigan_J._Frog
510. http://en.wikipedia.org/wiki/The_Frog_Prince_(story)
511. http://en.wikipedia.org/wiki/Kermit_the_Frog
512. http://en.wikipedia.org/wiki/Warner_Brothers
513. http://en.wikipedia.org/wiki/One_Froggy_Evening
514. http://en.wikipedia.org/wiki/The_Muppet_Show
515. http://en.wikipedia.org/wiki/Sesame_Street
516. http://en.wikipedia.org/wiki/Witch
517. http://en.wikipedia.org/wiki/Familiar_spirit
518. http://en.wikipedia.org/wiki/John_Milton
519. http://en.wikipedia.org/wiki/Paradise_Lost
520. http://en.wikipedia.org/wiki/Satan
521. http://en.wikipedia.org/wiki/Eve
522. http://en.wikipedia.org/wiki/Moche_(culture)
523. http://en.wikipedia.org/wiki/Peru
524. http://en.wikipedia.org/wiki/Panama

525. http://en.wikipedia.org/wiki/Talisman
526. http://en.wikipedia.org/wiki/Huaca
527. http://en.wikipedia.org/wiki/Mola_(art_form)
528. http://en.wikipedia.org/wiki/Kuna_people
529. http://en.wikipedia.org/wiki/Panama_City
530. http://en.wikipedia.org/wiki/Lottery_ticket
531. http://en.wikipedia.org/wiki/Digital_object_identifier
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533. http://en.wikipedia.org/wiki/PubMed_Identifier
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538. http://tolweb.org/Salientia/
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540. http://en.wikipedia.org/wiki/Special:BookSources/1-85310-740-9
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690. http://en.wikipedia.org/wiki/Special:BookSources/1-85310-740-9
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692. http://en.wikipedia.org/wiki/Special:BookSources/0-691-03281-5
693. http://www.jneurosci.org/content/18/8/2962.full
694. http://en.wikipedia.org/wiki/PubMed_Identifier
695. http://www.ncbi.nlm.nih.gov/pubmed/9526013
696. http://www.dnr.state.oh.us/Home/species_a_to_z/SpeciesGuideIndex/bullfrog/tabid/6576/Default.aspx
697. http://www.californiaherps.com/frogs/pages/a.truei.html
698. http://www.ias.ac.in/jarch/currsci/72/00000944.pdf
699. http://en.wikipedia.org/wiki/Digital_object_identifier
700. http://dx.doi.org/10.1146%2Fannurev.es.25.110194.001453
701. http://en.wikipedia.org/wiki/International_Standard_Book_Number
702. http://en.wikipedia.org/wiki/Special:BookSources/1-85310-740-9
703. http://www.beachwatchers.wsu.edu/island/essays/TreeFrogs.htm
704. http://records.viu.ca/~johnstoi/aristophanes/frogs.htm
705. http://www.scientificamerican.com/article.cfm?id=how-do-frogs-survive-wint
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707. http://dx.doi.org/10.1016%2Fj.cbpa.2009.09.001
708. http://en.wikipedia.org/wiki/American_Journal_of_Physiology
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710. http://dx.doi.org/10.1152%2Fajpregu.00380.2005
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714. http://en.wikipedia.org/wiki/Digital_object_identifier
715. http://dx.doi.org/10.1086%2F525290
716. http://en.wikipedia.org/wiki/PubMed_Identifier
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718. http://en.wikipedia.org/wiki/Digital_object_identifier
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720. http://en.wikipedia.org/wiki/International_Standard_Book_Number

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724. http://dx.doi.org/10.1098%2Frsbl.2011.0982
725. http://en.wikipedia.org/wiki/PubMed_Central
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729. http://en.wikipedia.org/wiki/Digital_object_identifier
730. http://dx.doi.org/10.1152%2Fadvan.00037.2004
731. http://en.wikipedia.org/wiki/PubMed_Identifier
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733. http://www.britannica.com/EBchecked/topic/29023/Anura
734. http://jeb.biologists.org/content/136/1/273
735. http://en.wikipedia.org/wiki/PubMed_Identifier
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745. http://en.wikipedia.org/wiki/International_Standard_Serial_Number
746. http://www.worldcat.org/issn/0011-3891
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748. http://amphibiaweb.org/lists/Scaphiopodidae.shtml
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750. http://www.amphibian.co.uk/scaphiophryne.html
751. http://www.mvences.de/p/p1/Vences_A51.pdf
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756. http://en.wikipedia.org/wiki/Special:BookSources/978-0-241-90338-4
757. http://amphibiaweb.org/cgi-bin/amphib_query?where-genus=Phyllomedusa&where-species=ayeaye
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759. http://dx.doi.org/10.2307%2F2409604

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