Overview
A frog is any member of a diverse and largely carnivorous group of short-bodied, tailless amphibians composing the order Anura (literally without tail in Ancient Greek). 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 in tropical rainforests. There are over 6,300 recorded species, accounting for around 88% of extant amphibian species. They are also one of the five most diverse vertebrate orders. Warty frog species tend to be called toads, but the distinction between frogs and toads is informal, not from taxonomy or evolutionary history.
An adult frog has a stout body, protruding eyes, anteriorly-attached tongue, limbs folded underneath, and no tail (except in tailed frogs). Frogs have glandular skin, with secretions ranging from distasteful to toxic. Their skin varies in color from well-camouflaged dappled brown, grey and green to vivid patterns of bright red or yellow and black to show toxicity and ward off predators. Adult frogs live in fresh water and on dry land; some species are adapted for living underground or in trees.
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 plant matter. Frog skin has a rich microbiome which is important to their health. Frogs are extremely efficient at converting what they eat into body mass. They are an important food source for predators and part of 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 behaviors to attract mates, to fend off predators and to generally survive.
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 that 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 that 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 should be placed 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 that 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. However, others have suggested that Gerobatrachus hottoni was only a dissorophoid temnospondyl unrelated to extant amphibians.
Salientia (Latin salere (salio), "to jump") is the name of the total group that includes modern frogs in the order Anura as well as their close fossil relatives, the "proto-frogs" or "stem-frogs". The common features possessed by these proto-frogs 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 known amphibians that were more closely related to frogs than to salamanders are Triadobatrachus massinoti, from the early Triassic period of Madagascar (about 250 million years ago), and 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 in modern frogs. The tibia and fibula bones are also separate, making it probable that Triadobatrachus was not an efficient leaper.
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 frogs 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.
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 frogs.
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. Exceptions include flying frogs in the Hylidae and Rhacophoridae, which also have fully webbed toes used in gliding.