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). Like other amphibians, oxygen 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. The ribs are poorly developed, so the lungs are filled by buccal pumping and a frog deprived of its lungs can maintain its body functions without them. 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.

Frogs range in size from the recently discovered 7.7-mm-long Paedophryne amauensis of Papua New Guinea  to the 300-mm goliath frog (Conraua goliath) of Cameroon. The skin hangs loosely on the body because of the lack of loose connective tissue. Frogs have three eyelid membranes: one is transparent to protect the eyes underwater, and two vary from translucent to opaque. They have a tympanum 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 in which the crown is separated from the root by fibrous tissue. These are on the edge of the upper jaw and vomerine teeth 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. The African bullfrog (Pyxicephalus), 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.

A bullfrog skeleton, showing elongated limb bones and extra joints. Red marks indicate bones which have been substantially elongated in frogs and joints which have become mobile. Blue indicates joints and bones which have not been modified or only somewhat elongated.
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 morphology modified for this purpose. The tibia, fibula, and tarsals have been fused into a single, strong bone, as have the radius and ulna in the fore limbs (which must absorb the impact on landing). The metatarsals 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 has elongated and formed a mobile joint with the sacrum 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.

Tyler's tree frog (Litoria tyleri) has large toe pads and webbed feet.
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.

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

Arboreal frogs 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. This allows the frog to climb on smooth surfaces, but the system does not function efficiently when the pads are excessively wet.

In many arboreal frogs, a small "intercalary structure" on each toe increases the surface area touching the substrate. 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.

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 (Scaphiopus couchii) have a flap-like toe extension on the hind feet, a keratinised tubercle often referred to as a spade, that helps them to burrow.

Sometimes during the tadpole stage, one of the developing rear legs is eaten by a predator such as a dragonfly nymph. 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 (Ribeiroia ondatrae) digs into the rear of a tadpole, causing a rearrangement of the limb bud cells and the frog develops an extra leg or two.

Evolution

The origins and evolutionary relationships between the three main groups of amphibians are hotly debated. A molecular phylogeny based on rDNA analysis dating from 2005 suggests salamanders and caecilians are more closely related to each other than they are to frogs and the divergence of the three groups took place in the Paleozoic or early Mesozoic before the breakup of the supercontinent Pangaea and soon after their divergence from the lobe-finned fishes. This would help account for the relative scarcity of amphibian fossils from the period before the groups split. Another molecular phylogenetic analysis conducted about the same time concluded the lissamphibians first appeared about 330 million years ago and that the temnospondyl-origin hypothesis is more credible than other theories. The neobatrachians seemed to have originated in Africa/India, the salamanders in East Asia and the caecilians in tropical Pangaea. 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, rather less than 300 million years ago, a date in better agreement with the palaeontological data. 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 and that it should be nested within Lepospondyli rather than within Temnospondyli. The study postulated the Lissamphibia originated no earlier than the late Carboniferous, some 290 to 305 million years ago. The split between Anura and Caudata 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.

A fossilized frog from the Czech Republic, possibly Palaeobatrachus gigas
In 2008, Gerobatrachus hottoni, a temnospondyl with many frog- and salamander-like characteristics, was discovered in Texas. It dated back 290 million years and was hailed as a missing link, a stem batrachian close to the common ancestor of frogs and salamanders, consistent with the widely accepted hypothesis that frogs and salamanders are more closely related to each other (forming a clade called Batrachia) than they are to caecilians. The earliest known salientians (see below), closer to the extant frogs than to the extant salamanders, are Triadobatrachus massinoti, from the Early Triassic of Madagascar (about 250 million years ago), and the fragmentary Czatkobatrachus polonicus from the Early Triassic of Poland (about the same age as Triadobatrachus). 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. 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.

Salientia (Latin salere (salio), "to jump") is a stem group including modern frogs in the order Anura and their close fossil relatives the "proto-frogs" (e.g., Triadobatrachus and Czatkobatrachus). The common features possessed by the "proto-frogs" in the Salientia group include 14 presacral vertebrae (modern frogs have eight or 9), a long and forward-sloping ilium in the pelvis, the presence of a frontoparietal bone, and a lower jaw without teeth. The earliest frog fossil that falls into the anuran lineage proper, Prosalirus bitis, lived in the early Jurassic. It was discovered in 1995 in the Kayenta Formation of Arizona and dates back to the Early Jurassic epoch (199.6 to 175 million years ago), making Prosalirus somewhat more recent than Triadobatrachus. Like the latter, Prosalirus did not have greatly enlarged legs, but had the typical three-pronged pelvic structure of modern frogs. Unlike Triadobatrachus, Prosalirus had already lost nearly all of its tail and was well adapted for jumping.

The earliest known "true frog" is Vieraella herbsti, from the Early Jurassic. It is known only from the dorsal and ventral impressions of a single animal and was estimated to be 33 mm (1.3 in) from snout to vent. Notobatrachus degiustoi from the middle Jurassic is slightly younger, about 155–170 million years old. 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 have taken place about 20 times faster in mammals than in frogs, which means speciation is occurring more rapidly in mammals.

Frog fossils have been found on all continents except Antarctica, but biogeographic evidence suggests they also inhabited Antarctica in an earlier era when the climate was warmer.

Etymology and taxonomy

The name frog derives from Old English frogga, abbreviated to frox, forsc, and frosc, probably deriving from Proto-Indo-European preu = "to jump". About 88% of amphibian species are classified in the order Anura. These include around 4,810 species in 33 families, of which the Leptodactylidae (1,100 spp.), Hylidae (800 spp.) and Ranidae (750 spp.) are the richest in species.
The use of the common names "frog" and "toad" has no taxonomic justification. From a classification perspective, all members of the order Anura are frogs, but only members of the family Bufonidae are considered "true toads". The use of the term "frog" in common names usually refers to species that are aquatic or semi-aquatic and have smooth, moist skins; the term "toad" generally refers to species that are terrestrial with dry, warty skins. There are numerous exceptions to this rule. The European fire-bellied toad (Bombina bombina) has a slightly warty skin and prefers a watery habitat whereas the Panamanian golden frog (Atelopus zeteki) is in the toad family Bufonidae and has a smooth skin.

The Anura include all modern frogs and any fossil 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, no tail, shorter fore limbs than hind limbs, radius and ulna fused, tibia and fibula fused, elongated ankle bones, absence of a prefrontal bone, presence of a hyoid plate, a lower jaw without teeth (with the exception of Gastrotheca guentheri) consisting of three pairs of bones (angulosplenial, dentary, and mentomeckelian, with the last pair being absent in Pipoidea), an unsupported tongue, lymph spaces underneath the skin, and a muscle, the protractor lentis, attached to the lens of the eye. The anuran larva or tadpole has a single central respiratory spiracle and mouthparts consisting of keratinous beaks and denticles.

Panamanian golden frog (Atelopus zeteki)
Frogs and toads are broadly classified into three suborders: Archaeobatrachia, which includes four families of primitive frogs; Mesobatrachia, which includes five families of more evolutionary intermediate frogs; and Neobatrachia, by far the largest group, which contains the remaining 24 families of modern frogs, including most common species found throughout the world. The Neobatrachia suborder is further divided into the two superfamilies Hyloidea and Ranoidea. This classification is based on such morphological features as the number of vertebrae, the structure of the pectoral girdle, and the morphology of tadpoles. While this classification is largely accepted, relationships among families of frogs are still debated.

Some species of anurans hybridize readily. For instance, the edible frog (Pelophylax esculentus) is a hybrid between the pool frog (P. lessonae) and the marsh frog (P. ridibundus). The fire-bellied toads Bombina bombina and B. variegata are similar in forming hybrids. These are less fertile than their parents, giving rise to a hybrid zone where the hybrids are prevalent.

Frogs

Frogs are a diverse and largely carnivorous group of short-bodied, tailless amphibians composing the order Anura (Ancient Greek an-, without + oura, tail). The oldest fossil "proto-frog" appeared in the early Triassic of Madagascar, but molecular clock dating suggests their origins may extend further back to the Permian, 265 million years ago. Frogs are widely distributed, ranging from the tropics to subarctic regions, but the greatest concentration of species diversity is found in tropical rainforests. 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 orders.

The body plan of an adult frog is generally characterized by a stout body, protruding eyes, cleft tongue, limbs folded underneath and the absence of a tail. 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, with secretions ranging from distasteful to toxic. Warty species of frog tend to be called toads but the distinction between frogs and toads is based on informal naming conventions concentrating on the warts rather than taxonomy or evolutionary history; some toads are more closely related to frogs than to other toads. Frogs' skins vary in colour from well-camouflaged dappled brown, grey and green to vivid patterns of bright red or yellow and black to advertise toxicity and warn off predators.

Frogs typically lay their eggs in water. The eggs hatch into aquatic larvae called tadpoles that have tails and internal gills. They have highly specialized rasping mouth parts suitable for herbivorous, omnivorous or planktivorous diets. The life cycle is completed when they metamorphose 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. Frogs are a keystone group in the food web dynamics of many of the world's ecosystems. The skin is semi-permeable, 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, particularly in their breeding season, and exhibit many different kinds of complex behaviours 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 with extinction and over one hundred and twenty are believed to have become extinct since the 1980s. The number of malformations among frogs is on the rise and an emerging fungal disease, chytridiomycosis, has spread around the world. Conservation biologists 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.