Frog

Frogs are the most numerous and widespread of all living amphibians. They are usually regarded as the symbol of amphibians, and are certainly the most distinctive. Most frogs are characterized by long hind legs, a short body, webbed digits (fingers or toes), protruding eyes and the absence of a tail. Frogs are widely known as exceptional jumpers, and many of the anatomical characteristics of frogs, particularly their long, powerful legs, are adaptations to improve jumping performance. Due to their permeable skin, frogs are often semi-aquatic or inhabit humid areas, but move easily on land. They typically lay their eggs in puddles, ponds or lakes, and their larvae, called tadpoles, have gills and develop in water. Adult frogs follow a carnivorous diet, mostly of arthropods, annelids and gastropods. Frogs are most noticeable by their call, which can be widely heard during the night or day, mainly in their mating season.

Frogs can be found nearly worldwide, with the exception af Antarctica and some northern regions such as Greenland. The main habitat of frogs is the tropical rainforest; however many species can be found in wetlands.

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

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

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

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

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

Classification and taxonomy
For more details on this topic, see List of Anuran Families. The order Anura contains 4,810 species in 33 families, of which the Leptodactylidae (1100 spp.), Hylidae (800 spp.) and Ranidae (750 spp.) are the richest in species. About 88% of amphibian species are frogs. The use of the common names "frog" and "toad" has no taxonomic justification. From a taxonomic perspective, all members of the order Anura are frogs, but only members of the suborder Archaeobatrachia are toads. The use of the term "frog" in common names usually refers to species that are aquatic or semi-aquatic with smooth and/or moist skins, and the term "toad" generally refers to species that tend to be terrestrial with dry, warty skin. An exception is the fire-bellied toad (Bombina bombina): while its skin is slightly warty, it prefers a watery habitat.

Frogs and toads are broadly classified into three suborders: Archaeobatrachia, which includes four families of toads; Mesobatrachia, by far the largest group, which contains 24 families of "standard" pond-living frogs and chorus frogs. Mesobatrachia is further divided into the 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. Future studies of molecular genetics should soon provide further insights to the evolutionary relationships among Anuran families. Finally, Neobatrachia contains the four familise of "Poison frogs", grouped into the superfamilies Dendrobatoidea and Mantelloidea.

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

Anatomy and physiology
Although frogs are usually viewed as the symbol of amphibians in popular culture, their morphology is unique in that they lack tails and primarily move about via jumping. Like all amphibians, frogs have a moist, highly permeable skin that can absorb oxygen. As a direct consequence, frogs must keep their skin moist with very few exceptions. At least 90% of frog species must stay in or near water.

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

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

Feet and legs
The structure of the feet and legs varies greatly among frog species, depending on whether they live primarily on the ground, in water, in trees, or in burrows. Frogs must be able to move quickly through their environment to catch prey and escape predators, and numerous adaptations help them do so. Many frogs, especially those that live in water, have webbed toes. The degree to which the toes are webbed is directly proportional to the amount of time the species lives in the water. For example, the completely aquatic African dwarf frog (Hymenochirus sp.) has fully webbed toes, whereas the toes of White's tree frog (Litoria caerulea), an arboreal species, are only a half or a quarter webbed. Arboreal frogs have "toe pads" to help grip vertical surfaces. These pads, located on the ends of the toes, do not work by suction. Rather, the surface of the pad consists of interlocking cells, with a small gap between adjacent cells. When the frog applies pressure to the toe pads, the interlocking cells grip irregularities on the substrate. The small gaps between the cells drain away all but a thin layer of moisture on the pad, and maintain a grip through capillarity. This allows the frog to grip smooth surfaces, and does not function when the pads are excessively wet.

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

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

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

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

Some frogs obtain poisons from the ants and other arthropods they eat; others, such as the Australian Corroboree Frogs (Pseudophryne corroboree and Pseudophryne pengilleyi), can manufacture an alkaloid not derived from their diet. Poisonous frogs tend to advertise their toxicity with bright colours, an adaptive strategy known as aposematism. There are at least two non-poisonous species of frogs in tropical America (Eleutherodactylus gaigei and Lithodytes lineatus) that mimic the colouration of dart poison frogs' coloration for self-protection (Batesian mimicry).

Toads in the Bufonidae family, like the Fire-bellied Toad and Cane toad contain hallucinogen poison that they secrete only when threatened, and then only from their parotoid glands. By contrast, the frogs of Neobatrachia secrete toxins through their skin constantly; these toxins tend to be much more potent. Frogs in the family Mantellidae secrete a hemotoxin that causes uncontrollable internal bleeding and destroys muscles. This hemotoxin functions like acid, seeping through the skin of would-be predators. Like the mantle frogs, dart poison frogs secrete a toxin through their skin; in their case a highly potent neurotoxin called batrachotoxin.

Calling
Some frog calls are so loud, they can be heard up to a mile away. The call of a frog is unique to its species. Frogs call by passing air through the larynx in the throat. In most calling frogs, the sound is amplified by one or more vocal sacs, membranes of skin under the throat or on the corner of the mouth that distend during the amplification of the call. The field of neuroethology studies the neurocircuitry that underlies frog audition. Some frogs lack vocal sacs, such as those from the genera Heleioporus and Neobatrachus, but these species can still produce a loud call. Their buccal cavity is enlarged and dome-shaped, acting as a resonance chamber that amplifies their call. Species of frog without vocal sacs and that do not have a loud call tend to inhabit areas close to flowing water. The noise of flowing water overpowers any call, so they must communicate by other means.

The main reason for calling is to allow males to attract a mate. Males call either individually or in a group called a chorus. Females of many frog species, for example Polypedates leucomystax, produce calls reciprocal to the males', which act as the catalyst for the enhancement of reproductive activity in a breeding colony. A male frog emits a release call when mounted by another male. Tropical species also have a rain call that they make on the basis of humidity cues prior to a rain shower. Many species also have a territorial call that is used to chase away other males. All of these calls are emitted with the mouth of the frog closed.

A distress call, emitted by some frogs when they are in danger, is produced with the mouth open, resulting in a higher-pitched call. In most frog species, it is suspected the call intrigues the predator until another animal is attracted, distracting them enough for its escape. With some species of frogs in the Mantellidae family, certain calls seem to signal other nearby frogs to poison their attacker, thus either killing or injuring the predator enough that it cannot continue an attack.

Reproduction and parental care
Once adult frogs reach maturity, they will assemble at a water source such as a pond or stream to breed. Many frogs return to the bodies of water where they were born, often resulting in annual migrations involving thousands of frogs. In continental Europe, a large proportion of migrating frogs used to die on roads, before special fences and tunnels were built for them. Once at the breeding ground, male frogs call to attract a mate, collectively becoming a chorus of frogs. The call is unique to the species, and will attract females of that species. Some species have satellite males who do not call, but intercept females that are approaching a calling male.

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

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