|The frog is an amphibian in the order Anura (meaning "tail-less", from Greek an-, without + oura, tail), formerly referred to as Salientia (Latin saltare, to jump). The name frog derives from Old English frogga, (compare Old Norse frauki, German Frosch, older Dutch spelling kikvorsch), cognate with Sanskrit plava (frog), probably deriving from Proto-Indo-European praw = "to jump".
Adult frogs are characterized by long hind legs, a short body, webbed digits, protruding eyes and the absence of a tail. Most frogs have a semi-aquatic lifestyle, but move easily on land by jumping or climbing. They typically lay their eggs in puddles, ponds or lakes, and their larvae, called tadpoles, have gills and develop in water. Adult frogs follow a carnivorous diet, mostly of arthropods, annelids and gastropods. Frogs are most noticeable by their call, which can be widely heard during the night or day, mainly in their mating season.
The distribution of frogs ranges from tropic to sub arctic regions, but most species are found in tropical rainforests. Consisting of more than 5,000 species described, they are among the most diverse groups of vertebrates. However, populations of certain frog species are significantly declining.
A distinction is often made between frogs and toads on the basis of their appearance, caused by the convergent adaptation among so-called toads to dry environments; however, this distinction has no taxonomic basis. The only family exclusively given the common name "toad" is Bufonidae, but many species from other families are also called "toads," and the species within the toad genus Atelopus are referred to as "harlequin frogs."
The use of the common names "frog" and "toad" has no taxonomic justification. From a taxonomic perspective, all members of the order Anura are frogs, but only members of the family Bufonidae are considered "true toads". The use of the term "frog" in common names usually refers to species that are aquatic or semi-aquatic with smooth and/or moist skins, and the term "toad" generally refers to species that tend to be terrestrial with dry, warty skin. An exception is the fire-bellied toad (Bombina bombina): while its skin is slightly warty, it prefers a watery habitat.
Frogs and toads are broadly classified into three suborders: Archaeobatrachia, which includes four families of primitive frogs; Mesobatrachia, which includes five families of more evolutionary intermediate frogs; and Neobatrachia, by far the largest group, which contains the remaining 24 families of "modern" frogs, including most common species throughout the world. Neobatrachia is further divided into the Hyloidea and Ranoidea. 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 frog families.
Some species of anurans hybridise readily. For instance, the edible frog (Rana esculenta) is a hybrid of the pool frog (R. lessonae) and the marsh frog (R. ridibunda). Bombina bombina and Bombina variegata similarly form hybrids, although these are less fertile, giving rise to a hybrid zone.
Morphology and physiology
Many characteristics are not shared by all of the approximately 5,250 described frog species. However, some general characteristics distinguish them from other amphibians. Frogs are usually well suited to jumping, with long hind legs and elongated ankle bones. They have a short vertebral column, with no more than ten free vertebrae, followed by a fused tailbone (urostyle or coccyx), typically resulting in a tailless phenotype.
Frogs range in size from 10 mm (Brachycephalus didactylus of Brazil and Eleutherodactylus iberia of Cuba) to 300 mm (goliath frog, Conraua goliath, of Cameroon). The skin hangs loosely on the body because of the lack of loose connective tissue. Skin texture varies: it can be smooth, warty or folded. Frogs have three eyelid membranes: one is transparent to protect the eyes underwater, and two vary from translucent to opaque. Frogs have a tympanum on each side of the head, which is involved in hearing and, in some species, is covered by skin. Most frogs do in fact have teeth of a sort. They have a ridge of very small cone teeth around the upper edge of the jaw. These are called maxillary teeth. Frogs often also have what are called vomerine teeth on the roof of their mouth. They do not have anything that could be called teeth on their lower jaw, so they usually swallow their food whole. The so-called "teeth" are mainly used to hold the prey and keep it in place till they can get a good grip on it and squash their eyeballs down to swallow their meal. Toads, however, do not have any teeth.
Feet and legs
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.
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 color, but this is usually restricted to shades of one or two colors. 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.
Certain frogs change color between night and day, as light and moisture stimulate the pigment cells and cause them to expand or contract.
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. Some native people of South America extract poison from the poison dart frogs and apply it to their darts for hunting, although few species are toxic enough to be used for this purpose. It was previously a misconception the poison was placed on arrows rather than darts. The common name of these frogs was thus changed from "poison arrow frog" to "poison dart frog" in the early 1980s. Poisonous frogs tend to advertise their toxicity with bright colours, an adaptive strategy known as aposematism. There are at least two non-poisonous species of frogs in tropical America (Eleutherodactylus gaigei and Lithodytes lineatus) that mimic the colouration of dart poison frogs' coloration for self-protection (Batesian mimicry).
Because frog toxins are extraordinarily diverse, they have raised the interest of biochemists as a "natural pharmacy". The alkaloid epibatidine, a painkiller 200 times more potent than morphine, is found in some species of poison dart frogs. Other chemicals isolated from the skin of frogs may offer resistance to HIV infection. Arrow and dart poisons are under active investigation for their potential as therapeutic drugs.
The skin secretions of some toads, such as the Colorado River toad and cane toad, contain bufotoxins, some of which, such as bufotenin, are psychoactive, and have therefore been used as recreational drugs. Typically, the skin secretions are dried and smoked. Skin licking is especially dangerous, and appears to constitute an urban myth. See psychoactive toad.
Respiration and circulation
Frogs are known for their three-chambered heart, which they share with all tetrapods except birds and mammals. In the three-chambered heart, oxygenated blood from the lungs and de-oxygenated blood from the respiring tissues enter by separate atria, and are directed via a spiral valve to the appropriate vessel aorta for oxygenated blood and pulmonary vein for deoxygenated blood. This special structure is essential to keeping the mixing of the two types of blood to a minimum, which enables frogs to have higher metabolic rates, and to be more active than otherwise.
Eggs hatch and continue life as tadpoles (occasionally known as polliwogs). Tadpoles are aquatic, lack front and hind legs, and have gills for respiration and tails with fins for swimming. Tadpoles are typically herbivorous, feeding mostly on algae, including diatoms filtered from the water through the gills. Some species are carnivorous at the tadpole stage, eating insects, smaller tadpoles, and fish. Tadpoles are highly vulnerable to predation by fish, newts, predatory diving beetles and birds such as kingfishers. Cannibalism has been observed among tadpoles. Poisonous tadpoles are present in many species, such as Cane Toads. The tadpole stage may be as short as a week, or tadpoles may over winter and metamorphose the following year in some species, such as the midwife toad (Alytes obstetricans) and the common spadefoot (Pelobates fuscus).
At the end of the tadpole stage, frogs undergo metamorphosis, in which they transition into adult form. Metamorphosis involves a dramatic transformation of morphology and physiology, as tadpoles develop hind legs, then front legs, lose their gills and develop lungs. Their intestines shorten as they shift from an herbivorous to a carnivorous diet. Eyes migrate rostrally and dorsally, allowing for binocular vision exhibited by the adult frog. This shift in eye position mirrors the shift from prey to predator, as the tadpole develops and depends less upon a larger and wider field of vision and more upon depth perception. The final stage of development from froglet to adult frog involves apoptosis (programmed cell death) and resorption of the tail.
After metamorphosis, young adults may leave the water and disperse into terrestrial habitats, or continue to live in the aquatic habitat as adults. Almost all species of frogs are carnivorous as adults, eating invertebrates such as arthropods, annelids and gastropods. A few of the larger species may eat prey such as small mammals, fish and smaller frogs. Some frogs use their sticky tongues to catch fast-moving prey, while others capture their prey and force it into their mouths with their hands. However, there are a very few species of frogs that primarily eat plants. Adult frogs are themselves preyed upon by birds, large fish, snakes, otters, foxes, badgers, coatis, and other animals. Frogs are also eaten by people (see section on uses in agriculture and research, below).
Although it is not common knowledge, some species of frog in their tadpole stage are known to be carnivorous. Early developers who gain legs may be eaten by the others, so the late bloomers survive longer. This has been observed in England in the species Rana temporaria (common frog).
Reproduction of frogs
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 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.
Other frogs carry the eggs and tadpoles on their hind legs or back (e.g., the midwife toads). Some frogs even protect their offspring inside their own bodies. The male Australian pouched frog (Assa darlingtoni) has pouches along its side in which the tadpoles reside until metamorphosis. The female gastric-brooding frogs (genus Rheobatrachus) from Australia, now probably extinct, swallows its tadpoles, which then develop in the stomach. To do this, the gastric-brooding frog must stop secreting stomach acid and suppress peristalsis (contractions of the stomach). Darwin's frog (Rhinoderma darwinii) from Chile puts the tadpoles in its vocal sac for development. Some species of frog will leave a 'babysitter' to watch over the frogspawn until it hatches.
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. The effectiveness of the call is unknown; however, it is suspected the call intrigues the predator until another animal is attracted, distracting them enough for its escape.
Many species of frog have deep calls, or croaks. The onomatopoeic spelling is "ribbit". The croak of the American bullfrog (Rana catesbiana) is sometimes spelt "jug o' rum". Other examples are Ancient Greek brekekekex koax koax for probably Rana ridibunda, and the description in Rigveda 7:103.6 góm āyur éko ajám āyur ékaħ = "one has a voice like a cow's, one has a voice like a goat's".
Distribution and conservation status
Frog populations have declined dramatically since the 1950s: more than one third of species are believed to be threatened with extinction and more than 120 species are suspected to be extinct since the 1980s. Among these species are the golden toad of Costa Rica and the Gastric-brooding frogs of Australia. Habitat loss is a significant cause of frog population decline, as are pollutants, climate change, the introduction of non-indigenous predators/competitors, and emerging infectious diseases including chytridiomycosis. Many environmental scientists believe that amphibians, including frogs, are excellent biological indicators of broader ecosystem health because of their intermediate position in food webs, permeable skins, and typically biphasic life (aquatic larvae and terrestrial adults).
A Canadian study conducted in 2006 proposed heavy traffic near frog habitats as a large threat to frog populations.
In a few cases, captive breeding programs have been attempted to alleviate the pressure on frog populations, and these have proved successful. In May 2007, it was reported the application of certain probiotic bacteria could protect amphibians from chytridiomycosis.
Another fossil frog, discovered in Arizona and called Prosalirus bitis, was uncovered in 1985, and dates from roughly the same time as Triadobatrachus. Like Triadobatrachus, Prosalirus did not have greatly enlarged legs, but had the typical three-pronged pelvic structure. Unlike Triadobatrachus, Prosalirus had already lost nearly all of its tail.
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 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, apparently still being a transitional species.
Uses in agriculture and research
Frogs have served as important model organisms throughout the history of science. Eighteenth-century biologist Luigi Galvani discovered the link between electricity and the nervous system through studying frogs. The African clawed frog or platanna (Xenopus laevis) was first widely used in laboratories in pregnancy assays in the first half of the 20th century. When human chorionic gonadotropin, a hormone found in substantial quantities in the urine of pregnant women, is injected into a female X. laevis, it induces them to lay eggs. In 1952, Robert Briggs and Thomas J. King cloned a frog by somatic cell nuclear transfer, the same technique later used to create Dolly the Sheep, their experiment was the first time successful nuclear transplantation had been accomplished in metazoans.
Frogs are used in cloning research and other branches of embryology because frogs are among the closest living relatives of man to lack egg shells characteristic of most other vertebrates, and therefore facilitate observations of early development. Although alternative pregnancy assays have been developed, biologists continue to use Xenopus as a model organism in developmental biology because it is easy to raise in captivity and has a large and easily manipulatable embryo. Recently, X. laevis is increasingly being displaced by its smaller relative X. tropicalis, which reaches its reproductive age in five months rather than one to two years (as in X. laevis), facilitating faster studies across generations. The genome sequence of X. tropicalis will probably be completed by 2015 at the latest.
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