(Anura)
Frogs and Toads
Безхвості земноводні
A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order Anura. Frog species with rough skin texture due to wart-like parotoid glands tend to be called toads, but the distinction between frogs and toads is informal and purely cosmetic, not from taxonomy or evolutionary history.
Frogs are widely distributed, ranging from the tropics to subarctic regions, but the greatest concentration of species diversity is in tropical rainforest and associated wetlands. They account for around 88% of extant amphibian species, and are one of the five most diverse vertebrate orders.
Adult frogs have a stout body, protruding eyes, anteriorly-attached tongue, limbs folded underneath, and no tail (the “tail” of tailed frogs is an extension of the male cloaca). Frogs have glandular skin, with secretions ranging from distasteful to toxic. Their skin varies in colour 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 both fresh water and on dry land; some species are adapted for living underground or in trees. As their skin is semi-permeable, making them susceptible to dehydration, they either live in moist niches or have special adaptations to deal with drier habitats. Frogs produce a wide range of vocalisations, particularly in their breeding season, and exhibit many different kinds of complex behaviors to attract mates, to fend off predators and to generally survive.
Being oviparous anamniotes, frogs typically spawn their eggs in bodies of water. The eggs then hatch into fully aquatic larvae called tadpoles, which have tails and internal gills. A few species lay eggs on land or bypass the tadpole stage altogether. Tadpoles have highly specialised rasping mouth parts suitable for herbivorous, omnivorous or planktivorous diets. The life cycle is completed when they metamorphose into semiaquatic adults capable of terrestrial locomotion and hybrid respiration using both lungs aided by buccal pumping and gas exchange across the skin, and the larval tail regresses into an internal urostyle. Adult frogs generally have a carnivorous diet consisting of small invertebrates, especially insects, but omnivorous species exist and a few feed on plant matter. Frogs generally seize and ingest food by protruding their adhesive tongue and then swallow the item whole, often using their eyeballs and extraocular muscles to help pushing down the throat, and their digestive system is extremely efficient at converting what they eat into body mass. Being low-level consumers, both tadpoles and adult frogs are an important food source for other predators and a vital part of the food web dynamics of many of the world’s ecosystems.
Feet and legs
A frog’s foot and leg structure is related to its habitat. Across species, these structures vary based on whether the species lives primarily on the ground, in water, in trees, or in burrows. Adult anurans have four fingers on the hands and five toes on the feet, but the smallest species often have hands and feet where some of the digits are vestigial. 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 jumpers or descend 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, allowing frogs to push against the ground for a longer period on take-off. The ilium 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 frogs to transfer force 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.
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 adhesion. 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, 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. 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 one or more extra legs.
Skin
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 (granular glands). The secretion is often sticky and helps keep the skin moist, protects against the entry of moulds and bacteria, and makes the animal slippery and more able to escape from predators.[60] 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.
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. 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 (Chiromantis xerampelina) is even able to turn white to minimise the chance of overheating.
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 specialised to produce sticky secretions to aid in amplexus. 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. Other frogs have adopted behaviours to conserve water, including becoming nocturnal 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. Woodhouse’s toad (Bufo woodhousii), if given access to water after confinement in a dry location, sits in the shallows to rehydrate. The male hairy frog (Trichobatrachus robustus) has dermal papillae projecting from its lower back and thighs, giving it a bristly appearance. These contain blood vessels and are thought to increase the area of the skin available for respiration.
Some species have bony plates embedded in the skin, a trait that appears to have evolved independently several times. 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).
Camouflage is a common defensive mechanism in frogs. Features such as warts and skin folds are usually on ground-dwelling frogs, for whom 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. Some are even able to control their skin texture. The Pacific tree frog (Pseudacris regilla) has green and brown morphs, plain or spotted, and changes colour depending on the time of year and general background colour. The Wood frog (Lithobates sylvaticus) uses disruptive coloration including black eye markings similar to voids between leaves, bands of the dorsal skin (dorsolateral dermal plica) similar to a leaf midrib as well as stains, spots and leg stripes similar to fallen leaf features.
Respiration and circulation
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. Ribs are generally absent, so the lungs are filled by buccal pumping and a frog deprived of its lungs can maintain its body functions without them. The fully aquatic Bornean flat-headed frog (Barbourula kalimantanensis) is the first frog known to lack lungs entirely.
Frogs have three-chambered hearts, a feature they share with lizards. Oxygenated blood from the lungs and de-oxygenated blood from the respiring tissues enter the heart through separate atria. When these chambers contract, the two blood streams pass into a common ventricle before being pumped via a spiral valve to the appropriate vessel, the aorta for oxygenated blood and pulmonary artery for deoxygenated blood.
Some species of frog have adaptations that allow them to survive in oxygen deficient water. The Titicaca water frog (Telmatobius culeus) 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.
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 insects 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. In amphibians there are salivary glands on the tongue, which in frogs produce what is called a two-phase viscoelastic fluid. When exposed to pressure, like when the tongue is wrapping around a prey, it becomes runny and covers the prey’s body. As the pressure drops, it returns to a thick and elastic state, which gives the tongue an extra grip. Some frogs have no tongue and just stuff food into their mouths with their hands. 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. 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.
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.
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 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. The nitrogen is excreted as ammonia by tadpoles and aquatic frogs but mainly as urea, 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. The urine passes along paired ureters to the urinary bladder from which it is vented periodically into the cloaca. All bodily wastes exit the body through the cloaca which terminates in a cloacal vent.
Reproductive system
In the male frog, the two testes are attached to the kidneys and semen passes into the kidneys through fine tubes called efferent ducts. 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.
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 and may be held for several days. The male frog has certain hormone-dependent secondary sexual characteristics. These include the development of special pads on his thumbs in the breeding season, to give him a firm hold. The grip of the male frog during amplexus stimulates the female to release eggs, usually wrapped in jelly, as spawn. 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 to amplify the sound.
Nervous system
Frogs have a highly developed nervous system that consists of a brain, spinal cord and nerves. Many parts of frog brains 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, and the medulla oblongata regulates respiration, digestion and other automatic functions. The relative size of the cerebrum in frogs is much smaller than it is in humans. Frogs have ten pairs of cranial nerves which pass information from the outside directly to the brain, and ten pairs of spinal nerves which pass information from the extremities to the brain through the spinal cord. By contrast, all amniotes (mammals, birds and reptiles) have twelve pairs of cranial nerves.
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 over a field of 100° to the front and a total visual field of almost 360°. 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 which provides further protection, especially when the frog is swimming. Members of the aquatic family Pipidae have the eyes located at the top of the head, a position better suited for detecting prey in the water above. The irises come in a range of colours and the pupils in a range of shapes. The common toad (Bufo bufo) has golden irises and horizontal slit-like pupils, the red-eyed tree frog (Agalychnis callidryas) has vertical slit pupils, the poison dart frog has dark irises, the fire-bellied toad (Bombina spp.) has triangular pupils and the tomato frog (Dyscophus spp.) has circular ones. The irises of the southern toad (Anaxyrus terrestris) are patterned so as to blend in with the surrounding camouflaged skin.
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. 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. Although it was formerly debated, more recent research has shown that frogs can see in colour, even in very low light.
Hearing
Frogs can hear both in the air and below water. They do not have external ears; the eardrums (tympanic membranes) 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. 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. Because the cochlea is short, frogs use electrical tuning to extend their range of audible frequencies and help discriminate different sounds. This arrangement enables detection of the territorial and breeding calls of their conspecifics. In some species that inhabit arid regions, the sound of thunder or heavy rain may arouse them from a dormant state. 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.
Call
The call or croak of a frog is unique to its species. Frogs create this sound 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. Some frog calls are so loud that they can be heard up to a mile (1.6 km) away. Additionally, some species have been found to use man-made structures such as drain pipes for artificial amplification of their call. The coastal tailed frog (Ascaphus truei) lives in mountain streams in North America and does not vocalise.
The main function of calling is for male frogs to attract mates. Males may call individually or there may be a chorus of sound where numerous males have converged on breeding sites. In many frog species, such as the common tree frog (Polypedates leucomystax), females reply to males’ calls, which acts to reinforce reproductive activity in a breeding colony. 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.
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. Tree frogs and some non-aquatic species have a rain call that they make on the basis of humidity cues prior to a shower. 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. 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 disorient the attacker so that it releases the frog.
Many species of frog have deep calls. The croak of the American bullfrog (Rana catesbiana) is sometimes written as “jug o’ rum”. The Pacific tree frog (Pseudacris regilla) produces the onomatopoeic “ribbit” often heard in films. Other renderings of frog calls into speech include “brekekekex koax koax”, the call of the marsh frog (Pelophylax ridibundus) in The Frogs, an Ancient Greek comic drama by Aristophanes. The calls of the Concave-eared torrent frog (Amolops tormotus) are unusual in many aspects. The males are notable for their varieties of calls where upward and downward frequency modulations take place. When they communicate, they produce calls that fall in the ultrasound frequency range. The last aspect that makes this species of frog’s calls unusual is that nonlinear acoustic phenomena are important components in their acoustic signals.
Torpor
During extreme conditions, some frogs enter a state of torpor and remain inactive for months. In colder regions, many species of frog hibernate in winter. Those that live on land such as the American toad (Bufo americanus) dig a burrow and make a hibernaculum in which to lie dormant. Others, less proficient at digging, find a crevice or bury themselves in dead leaves. Aquatic species such as the American bullfrog (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 such as the wood frog, moor frog, or spring peeper 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 its heartbeat can restart when conditions warm up.
At the other extreme, the striped burrowing frog (Cyclorana alboguttata) regularly aestivates 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 formed by its shed skin. Researchers at the University of Queensland have found that during aestivation, the metabolism of the frog is altered and the operational efficiency of the mitochondria 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. Other research showed that, to provide these energy requirements, muscles atrophy, but hind limb muscles are preferentially unaffected. Frogs have been found to have upper critical temperatures of around 41° C.
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.
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. 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.
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 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. Some female New Mexico spadefoot toads (Spea multiplicata) only spawn half of the available eggs at a time, perhaps retaining some in case a better reproductive opportunity arises later.
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; fertilisation is external. In many species such as the Great Plains toad (Bufo cognatus), the male restrains the eggs with his back feet, holding them in place for about three minutes. Members of the West African genus Nimbaphrynoides are unique among frogs in that they are viviparous; Limnonectes larvaepartus, Eleutherodactylus jasperi and members of the Tanzanian genus Nectophrynoides are the only frogs known to be ovoviviparous. In these species, fertilisation is internal and females give birth to fully developed juvenile frogs, except L. larvaepartus, which give birth to tadpoles.
Life cycle
Frogs may lay their eggs as clumps, surface films, strings, or individually. Around half of species deposit eggs in water, others lay eggs in vegetation, on the ground or in excavations. The tiny yellow-striped pygmy eleuth (Eleutherodactylus limbatus) lays eggs singly, burying them in moist soil. The smoky jungle frog (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. 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.
In certain species, such as the wood frog (Rana sylvatica), symbiotic 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. The interior of globular egg clusters of the wood frog has also been found to be up to 6 °C warmer than the surrounding water and this speeds up the development of the larvae. The larvae developing in the eggs can detect vibrations caused by nearby predatory wasps or snakes, and will hatch early to avoid being eaten. 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 released by the developing larvae.
Direct development, where eggs hatch into juveniles like small adults, is also known in many frogs, for example, Ischnocnema henselii, Eleutherodactylus coqui, and Raorchestes ochlandrae and Raorchestes chalazodes.
The larvae that emerge from the eggs are known as tadpoles (or occasionally polliwogs). Tadpoles lack eyelids and limbs, and have cartilaginous skeletons, gills for respiration (external gills at first, internal gills later), and tails they use for swimming. As a general rule, free-living larvae are fully aquatic, but at least one species (Nannophrys ceylonensis) has semiterrestrial tadpoles which live among wet rocks.
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 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 provide a means for species identification. In the Pipidae, with the exception of Hymenochirus, the tadpoles have paired anterior barbels, which make them resemble small catfish. Their tails are stiffened by a notochord, 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 try to grasp them by the tail.
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. The Cuban tree frog (Osteopilus septentrionalis) is one of a number of species in which the tadpoles can be cannibalistic. Tadpoles that develop legs early may be eaten by the others, so late developers may have better long-term survival prospects.
Tadpoles are highly vulnerable to being eaten by fish, newts, predatory diving beetles, and birds, particularly water birds, such as storks and herons and domestic ducks. Some tadpoles, including those of the cane toad (Rhinella marina), 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.
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 thyroxine. 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. Homeostatic feedback control of food intake is largely absent, making tadpoles eat constantly when food is present. But shortly before and during metamorphosis the sensation of hunger is suppressed, and they stop eating while their gut and internal organs are reorganised and prepared for a different diet. Also the gut microbiota changes, from being similar to that of fish to resembling that of amniotes. Exceptions are carnivorous tadpoles like Lepidobatrachus laevis, which has a gut already adapted to a diet similar to that of adults. These continue to eat during metamorphosis. The nervous system becomes adapted for hearing and stereoscopic vision, and for new methods of locomotion and feeding. 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. 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. 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.
Adult frogs may live in or near water, but few are fully aquatic. Almost all frog species are carnivorous as adults, preying on invertebrates, including insects, crabs, spiders, mites, worms, snails, and slugs. A few of the larger ones may eat other frogs, small mammals and reptiles, and fish. A few species also eat plant matter; the tree frog Xenohyla truncata is partly herbivorous, its diet including a large proportion of fruit, floral structures and nectar. Leptodactylus mystaceus has been found to eat plants, and folivory occurs in Euphlyctis hexadactylus, with plants constituting 79.5% of its diet by volume. Many frogs use their sticky tongues to catch prey, while others simply grab them with their mouths. Adult frogs are themselves attacked by many predators. The northern leopard frog (Rana pipiens) is eaten by herons, hawks, fish, large salamanders, snakes, raccoons, skunks, mink, bullfrogs, and other animals.
Frogs are primary predators and an important part of the food web. Being cold-blooded, they make efficient use of the food they eat with little energy being used for metabolic processes, while the rest is transformed into biomass. 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.
Little is known about the longevity of frogs and toads in the wild, but some can live for many years. Skeletochronology is a method of examining bones to determine age. Using this method, the ages of mountain yellow-legged frogs (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. 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 (Rhinella marina) has been known to survive 24 years in captivity, and the American bullfrog (Rana catesbeiana) 14 years. 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).
Defence
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.
The skin of many frogs contains mild toxic substances called bufotoxins to make them unpalatable to potential predators. Most toads and some frogs have large poison glands, the parotoid glands, 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. Poisonous frogs tend to advertise their toxicity with bright colours, an adaptive strategy known as aposematism. 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 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. Other species, such as the European fire-bellied toad (Bombina bombina), have their warning colour underneath. They “flash” this when attacked, adopting a pose that exposes the vivid colouring on their bellies.
Some frogs, such as the poison dart frogs, are especially toxic. The native peoples of South America extract poison from these frogs to apply to their weapons for hunting, although few species are toxic enough to be used for this purpose. At least two non-poisonous frog species in tropical America (Eleutherodactylus gaigei and Lithodytes lineatus) mimic the colouration of dart poison frogs for self-protection. 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 synthesize the alkaloids themselves. The chemicals involved may be irritants, hallucinogens, convulsants, nerve poisons or vasoconstrictors. 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.
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. 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. Another tactic used by some frogs is to “scream”, the sudden loud noise tending to startle the predator. The grey tree frog (Hyla versicolor) makes an explosive sound that sometimes repels the shrew Blarina brevicauda. Although toads are avoided by many predators, the common garter snake (Thamnophis sirtalis) regularly feeds on them. The strategy employed by juvenile American toads (Bufo americanus) 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.
Distribution
Frogs live on every continent except Antarctica, but they are not present on certain islands, especially those far away from continental land masses. 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 human-made barriers. Usually, a greater diversity of frogs occurs in tropical areas than in temperate regions, such as Europe. Some frogs inhabit arid areas, such as deserts, and rely on specific adaptations to survive. Members of the Australian genus Cyclorana bury themselves underground where they create a water-impervious cocoon in which to aestivate during dry periods. Once it rains, they emerge, find a temporary pool, and breed. Egg and tadpole development is very fast compared with those of most other frogs, so breeding can be completed before the pond dries up. Some frog species are adapted to a cold environment. The wood frog (Rana sylvatica), whose habitat extends into the Arctic Circle, 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.
