Amphibians

Amphibians are ectothermic, anamniotic, four-limbed vertebrate animals that constitute the class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all tetrapods excluding the amniotes (tetrapods with an amniotic membrane, such as modern reptiles, birds and mammals). All extant (living) amphibians belong to the monophyletic subclass Lissamphibia, with three living orders: Anura (frogs and toads), Urodela (salamanders), and Gymnophiona (caecilians). Evolved to be mostly semiaquatic, amphibians have adapted to inhabit a wide variety of habitats, with most species living in freshwater, wetland or terrestrial ecosystems (such as riparian woodland, fossorial and even arboreal habitats). Their life cycle typically starts out as aquatic larvae with gills known as tadpoles, but some species have developed behavioural adaptations to bypass this.

Young amphibians generally undergo metamorphosis from an aquatic larval form with gills to an air-breathing adult form with lungs. Amphibians use their skin as a secondary respiratory interface and some small terrestrial salamanders and frogs lack lungs and rely entirely on their skin. They are superficially similar to reptiles like lizards, but unlike reptiles and other amniotes, require access to water bodies to breed. With their complex reproductive needs and permeable skins, amphibians are often ecological indicators to habitat conditions; in recent decades there has been a dramatic decline in amphibian populations for many species around the globe.

The number of known amphibian species is approximately 8,000, of which nearly 90% are frogs. The smallest amphibian (and vertebrate) in the world is a frog from New Guinea (Paedophryne amauensis) with a length of just 7.7 mm. The largest living amphibian is the 1.8 m South China giant salamander (Andrias sligoi), but this is dwarfed by prehistoric temnospondyls such as Mastodonsaurus which could reach up to 6 m in length.

The superclass Tetrapoda is divided into four classes of vertebrate animals with four limbs. Reptiles, birds and mammals are amniotes, the eggs of which are either laid or carried by the female and are surrounded by several membranes, some of which are impervious. Lacking these membranes, amphibians require water bodies for reproduction, although some species have developed various strategies for protecting or bypassing the vulnerable aquatic larval stage. They are not found in the sea with the exception of one or two frogs that live in brackish water in mangrove swamps; the Anderson’s salamander meanwhile occurs in brackish or salt water lakes. On land, amphibians are restricted to moist habitats because of the need to keep their skin damp.

Amphibians are ectothermic (cold-blooded) vertebrates that do not maintain their body temperature through internal physiological processes. Their metabolic rate is low and as a result, their food and energy requirements are limited. In the adult state, they have tear ducts and movable eyelids, and most species have ears that can detect airborne or ground vibrations. They have muscular tongues, which in many species can be protruded. Modern amphibians have fully ossified vertebrae with articular processes. Their ribs are usually short and may be fused to the vertebrae. Their skulls are mostly broad and short, and are often incompletely ossified. Their skin contains little keratin and lacks scales, apart from a few fish-like scales in certain caecilians. The skin contains many mucous glands and in some species, poison glands (a type of granular gland). The hearts of amphibians have three chambers, two atria and one ventricle. They have a urinary bladder and nitrogenous waste products are excreted primarily as urea. Most amphibians lay their eggs in water and have aquatic larvae that undergo metamorphosis to become terrestrial adults. Amphibians breathe by means of a pump action in which air is first drawn into the buccopharyngeal region through the nostrils. These are then closed and the air is forced into the lungs by contraction of the throat. They supplement this with gas exchange through the skin.

Reproduction

For the purpose of reproduction, most amphibians require fresh water although some lay their eggs on land and have developed various means of keeping them moist. A few (e.g. Fejervarya raja) can inhabit brackish water, but there are no true marine amphibians. There are reports, however, of particular amphibian populations unexpectedly invading marine waters.

Several hundred frog species in adaptive radiations (e.g., Eleutherodactylus, the Pacific Platymantis, the Australo-Papuan microhylids, and many other tropical frogs), however, do not need any water for breeding in the wild. They reproduce via direct development, an ecological and evolutionary adaptation that has allowed them to be completely independent from free-standing water. Almost all of these frogs live in wet tropical rainforests and their eggs hatch directly into miniature versions of the adult, passing through the tadpole stage within the egg. Reproductive success of many amphibians is dependent not only on the quantity of rainfall, but the seasonal timing.

In the tropics, many amphibians breed continuously or at any time of year. In temperate regions, breeding is mostly seasonal, usually in the spring, and is triggered by increasing day length, rising temperatures or rainfall. Experiments have shown the importance of temperature, but the trigger event, especially in arid regions, is often a storm. In anurans, males usually arrive at the breeding sites before females and the vocal chorus they produce may stimulate ovulation in females and the endocrine activity of males that are not yet reproductively active.

In caecilians, fertilisation is internal, the male extruding an intromittent organ, the phallodeum, and inserting it into the female cloaca. The paired Müllerian glands inside the male cloaca secrete a fluid which resembles that produced by mammalian prostate glands and which may transport and nourish the sperm. Fertilisation probably takes place in the oviduct.

The majority of salamanders also engage in internal fertilisation. In most of these, the male deposits a spermatophore, a small packet of sperm on top of a gelatinous cone, on the substrate either on land or in the water. The female takes up the sperm packet by grasping it with the lips of the cloaca and pushing it into the vent. The spermatozoa move to the spermatheca in the roof of the cloaca where they remain until ovulation which may be many months later. Courtship rituals and methods of transfer of the spermatophore vary between species. In some, the spermatophore may be placed directly into the female cloaca while in others, the female may be guided to the spermatophore or restrained with an embrace called amplexus. Certain primitive salamanders in the families Sirenidae, Hynobiidae and Cryptobranchidae practice external fertilisation in a similar manner to frogs, with the female laying the eggs in water and the male releasing sperm onto the egg mass.

With a few exceptions, frogs use external fertilisation. The male grasps the female tightly with his forelimbs either behind the arms or in front of the back legs, or in the case of Epipedobates tricolor, around the neck. They remain in amplexus with their cloacae positioned close together while the female lays the eggs and the male covers them with sperm. Roughened nuptial pads on the male’s hands aid in retaining grip. Often the male collects and retains the egg mass, forming a sort of basket with the hind feet. An exception is the granular poison frog (Oophaga granulifera) where the male and female place their cloacae in close proximity while facing in opposite directions and then release eggs and sperm simultaneously. The tailed frog (Ascaphus truei) exhibits internal fertilisation. The “tail” is only possessed by the male and is an extension of the cloaca and used to inseminate the female. This frog lives in fast-flowing streams and internal fertilisation prevents the sperm from being washed away before fertilisation occurs. The sperm may be retained in storage tubes attached to the oviduct until the following spring.

Most frogs can be classified as either prolonged or explosive breeders. Typically, prolonged breeders congregate at a breeding site, the males usually arriving first, calling and setting up territories. Other satellite males remain quietly nearby, waiting for their opportunity to take over a territory. The females arrive sporadically, mate selection takes place and eggs are laid. The females depart and territories may change hands. More females appear and in due course, the breeding season comes to an end. Explosive breeders on the other hand are found where temporary pools appear in dry regions after rainfall. These frogs are typically fossorial species that emerge after heavy rains and congregate at a breeding site. They are attracted there by the calling of the first male to find a suitable place, perhaps a pool that forms in the same place each rainy season. The assembled frogs may call in unison and frenzied activity ensues, the males scrambling to mate with the usually smaller number of females.

There is a direct competition between males to win the attention of the females in salamanders and newts, with elaborate courtship displays to keep the female’s attention long enough to get her interested in choosing him to mate with. Some species store sperm through long breeding seasons, as the extra time may allow for interactions with rival sperm.

Life cycle

Most amphibians go through metamorphosis, a process of significant morphological change after birth. In typical amphibian development, eggs are laid in water and larvae are adapted to an aquatic lifestyle. Frogs, toads and salamanders all hatch from the egg as larvae with external gills. Metamorphosis in amphibians is regulated by thyroxine concentration in the blood, which stimulates metamorphosis, and prolactin, which counteracts thyroxine’s effect. Specific events are dependent on threshold values for different tissues. Because most embryonic development is outside the parental body, it is subject to many adaptations due to specific environmental circumstances. For this reason tadpoles can have horny ridges instead of teeth, whisker-like skin extensions or fins. They also make use of a sensory lateral line organ similar to that of fish. After metamorphosis, these organs become redundant and will be reabsorbed by controlled cell death, called apoptosis. The variety of adaptations to specific environmental circumstances among amphibians is wide, with many discoveries still being made.

In the egg, the embryo is suspended in perivitelline fluid and surrounded by semi-permeable gelatinous capsules, with the yolk mass providing nutrients. As the larvae hatch, the capsules are dissolved by enzymes secreted from gland at the tip of the snout. The eggs of some salamanders and frogs contain unicellular green algae. These penetrate the jelly envelope after the eggs are laid and may increase the supply of oxygen to the embryo through photosynthesis. They seem to both speed up the development of the larvae and reduce mortality. In the wood frog (Rana sylvatica), the interior of the globular egg cluster has been found to be up to 6 °C warmer than its surroundings, which is an advantage in its cool northern habitat.

The eggs may be deposited singly, in cluster or in long strands. Sites for laying eggs include water, mud, burrows, debris and on plants or under logs or stones. The greenhouse frog (Eleutherodactylus planirostris) lays eggs in small groups in the soil where they develop in about two weeks directly into juvenile frogs without an intervening larval stage. The tungara frog (Physalaemus pustulosus) builds a floating nest from foam to protect its eggs. First a raft is built, then eggs are laid in the centre, and finally a foam cap is overlaid. The foam has anti-microbial properties. It contains no detergents but is created by whipping up proteins and lectins secreted by the female.

The eggs of amphibians are typically laid in water and hatch into free-living larvae that complete their development in water and later transform into either aquatic or terrestrial adults. In many species of frog and in most lungless salamanders (Plethodontidae), direct development takes place, the larvae growing within the eggs and emerging as miniature adults. Many caecilians and some other amphibians lay their eggs on land, and the newly hatched larvae wriggle or are transported to water bodies. Some caecilians, the alpine salamander (Salamandra atra) and some of the African live-bearing toads (Nectophrynoides spp.) are viviparous. Their larvae feed on glandular secretions and develop within the female’s oviduct, often for long periods. Other amphibians, but not caecilians, are ovoviviparous. The eggs are retained in or on the parent’s body, but the larvae subsist on the yolks of their eggs and receive no nourishment from the adult. The larvae emerge at varying stages of their growth, either before or after metamorphosis, according to their species. The toad genus Nectophrynoides exhibits all of these developmental patterns among its dozen or so members. Amphibian larvae are known as tadpoles. They have thick, rounded bodies with powerful muscular tails.

Frogs

Unlike in other amphibians, frog tadpoles do not resemble adults. The free-living larvae are normally fully aquatic, but the tadpoles of some species (such as Nannophrys ceylonensis) are semi-terrestrial and live among wet rocks. Tadpoles have cartilaginous skeletons, gills for respiration (external gills at first, internal gills later), lateral line systems and large tails that they use for swimming. Newly hatched tadpoles soon develop gill pouches that cover the gills. These internal gills and operculum are not homologous with those of fish, and are only found in tadpoles as both salamanders and caecilians have external gills only. Combined with buccal pumping the internal gills has allowed tadpoles to adopt a filter feeding lifestyle, even if several species have since evolved other types of feeding strategies.The lungs develop early and are used as accessory breathing organs, the tadpoles rising to the water surface to gulp air. Some species complete their development inside the egg and hatch directly into small frogs. These larvae do not have gills but instead have specialised areas of skin through which respiration takes place. While tadpoles do not have true teeth, in most species, the jaws have long, parallel rows of small keratinized structures called keradonts surrounded by a horny beak. Front legs are formed under the gill sac and hind legs become visible a few days later.

In fact, tadpoles developing in ponds and streams are typically herbivorous. Pond tadpoles tend to have deep bodies, large caudal fins and small mouths; they swim in the quiet waters feeding on growing or loose fragments of vegetation. Stream dwellers mostly have larger mouths, shallow bodies and caudal fins; they attach themselves to plants and stones and feed on the surface films of algae and bacteria. They also feed on diatoms, filtered from the water through the gills, and stir up the sediment at bottom of the pond, ingesting edible fragments. They have a relatively long, spiral-shaped gut to enable them to digest this diet. Some species are carnivorous at the tadpole stage, eating insects, smaller tadpoles and fish. Young of the Cuban tree frog (Osteopilus septentrionalis) can occasionally be cannibalistic, the younger tadpoles attacking a larger, more developed tadpole when it is undergoing metamorphosis.

At metamorphosis, rapid changes in the body take place as the lifestyle of the frog changes completely. The spiral-shaped mouth with horny tooth ridges is reabsorbed together with the spiral gut. The animal develops a large jaw, and its gills disappear along with its gill sac. Eyes and legs grow quickly, and a tongue is formed. There are associated changes in the neural networks such as development of stereoscopic vision and loss of the lateral line system. All this can happen in about a day. A few days later, the tail is reabsorbed, due to the higher thyroxine concentration required for this to take place.

Salamanders

At hatching, a typical salamander larva has eyes without lids, teeth in both upper and lower jaws, three pairs of feathery external gills, and a long tail with dorsal and ventral fins. The forelimbs may be partially developed and the hind limbs are rudimentary in pond-living species but may be rather more developed in species that reproduce in moving water. Pond-type larvae often have a pair of balancers, rod-like structures on either side of the head that may prevent the gills from becoming clogged up with sediment. Both of these are able to breed. Some have larvae that never fully develop into the adult form, a condition known as neoteny. Neoteny occurs when the animal’s growth rate is very low and is usually linked to adverse conditions such as low water temperatures that may change the response of the tissues to the hormone thyroxine, as well as lack of food. There are fifteen species of obligate neotenic salamanders, including species of Necturus, Proteus and Amphiuma, and many examples of facultative ones, such as the northwestern salamander (Ambystoma gracile) and the tiger salamander (A. tigrinum) that adopt this strategy under appropriate environmental circumstances.

Lungless salamanders in the family Plethodontidae are terrestrial and lay a small number of unpigmented eggs in a cluster among damp leaf litter. Each egg has a large yolk sac and the larva feeds on this while it develops inside the egg, emerging fully formed as a juvenile salamander. The female salamander often broods the eggs. In the genus Ensatinas, the female has been observed to coil around them and press her throat area against them, effectively massaging them with a mucous secretion.

In newts and salamanders, metamorphosis is less dramatic than in frogs. This is because the larvae are already carnivorous and continue to feed as predators when they are adults so few changes are needed to their digestive systems. Their lungs are functional early, but the larvae do not make as much use of them as do tadpoles. Their gills are never covered by gill sacs and are reabsorbed just before the animals leave the water. Other changes include the reduction in size or loss of tail fins, the closure of gill slits, thickening of the skin, the development of eyelids, and certain changes in dentition and tongue structure. Salamanders are at their most vulnerable at metamorphosis as swimming speeds are reduced and transforming tails are encumbrances on land. Adult salamanders often have an aquatic phase in spring and summer, and a land phase in winter. For adaptation to a water phase, prolactin is the required hormone, and for adaptation to the land phase, thyroxine. External gills do not return in subsequent aquatic phases because these are completely absorbed upon leaving the water for the first time.

Caecilians

Most terrestrial caecilians that lay eggs do so in burrows or moist places on land near bodies of water. The development of the young of Ichthyophis glutinosus, a species from Sri Lanka, has been much studied. The eel-like larvae hatch out of the eggs and make their way to water. They have three pairs of external red feathery gills, a blunt head with two rudimentary eyes, a lateral line system and a short tail with fins. They swim by undulating their body from side to side. They are mostly active at night, soon lose their gills and make sorties onto land. Metamorphosis is gradual. By the age of about ten months they have developed a pointed head with sensory tentacles near the mouth and lost their eyes, lateral line systems and tails. The skin thickens, embedded scales develop and the body divides into segments. By this time, the caecilian has constructed a burrow and is living on land.

In the majority of species of caecilians, the young are produced by viviparity. Typhlonectes compressicauda, a species from South America, is typical of these. Up to nine larvae can develop in the oviduct at any one time. They are elongated and have paired sac-like gills, small eyes and specialised scraping teeth. At first, they feed on the yolks of the eggs, but as this source of nourishment declines they begin to rasp at the ciliated epithelial cells that line the oviduct. This stimulates the secretion of fluids rich in lipids and mucoproteins on which they feed along with scrapings from the oviduct wall. They may increase their length sixfold and be two-fifths as long as their mother before being born. By this time they have undergone metamorphosis, lost their eyes and gills, developed a thicker skin and mouth tentacles, and reabsorbed their teeth. A permanent set of teeth grow through soon after birth.

Gills are only necessarily during embryonic development, and in species that give birth the offspring is born after gill degeneration. In egg laying caecilians the gills are either reabsorbed before hatching, or, in species that hatch with gill remnants still present, short lived and only leaves behind a gill slit. For species with scales under their skin, the scales does not form before during metamorphosis.

The ringed caecilian (Siphonops annulatus) has developed a unique adaptation for the purposes of reproduction. The progeny feed on a skin layer that is specially developed by the adult in a phenomenon known as maternal dermatophagy. The brood feed as a batch for about seven minutes at intervals of approximately three days which gives the skin an opportunity to regenerate. Meanwhile, they have been observed to ingest fluid exuded from the maternal cloaca.

Feeding and diet

With a few exceptions, adult amphibians are predators, feeding on virtually anything that moves that they can swallow. The diet mostly consists of small prey that do not move too fast such as beetles, caterpillars, earthworms and spiders. The sirens (Siren spp.) often ingest aquatic plant material with the invertebrates on which they feed and a Brazilian tree frog (Xenohyla truncata) includes a large quantity of fruit in its diet. The Mexican burrowing toad (Rhinophrynus dorsalis) has a specially adapted tongue for picking up ants and termites. It projects it with the tip foremost whereas other frogs flick out the rear part first, their tongues being hinged at the front.

Food is mostly selected by sight, even in conditions of dim light. Movement of the prey triggers a feeding response. Frogs have been caught on fish hooks baited with red flannel and green frogs (Rana clamitans) have been found with stomachs full of elm seeds that they had seen floating past. Toads, salamanders and caecilians also use smell to detect prey. This response is mostly secondary because salamanders have been observed to remain stationary near odoriferous prey but only feed if it moves. Cave-dwelling amphibians normally hunt by smell. Some salamanders seem to have learned to recognize immobile prey when it has no smell, even in complete darkness.

Amphibians usually swallow food whole but may chew it lightly first to subdue it. They typically have small hinged pedicellate teeth, a feature unique to amphibians. The base and crown of these are composed of dentine separated by an uncalcified layer and they are replaced at intervals. Salamanders, caecilians and some frogs have one or two rows of teeth in both jaws, but some frogs (Rana spp.) lack teeth in the lower jaw, and toads (Bufo spp.) have no teeth. In many amphibians there are also vomerine teeth attached to a facial bone in the roof of the mouth.

The tiger salamander (Ambystoma tigrinum) is typical of the frogs and salamanders that hide under cover ready to ambush unwary invertebrates. Other amphibians, such as the Bufo spp. toads, actively search for prey, while the Argentine horned frog (Ceratophrys ornata) lures inquisitive prey closer by raising its hind feet over its back and vibrating its yellow toes. Among leaf litter frogs in Panama, frogs that actively hunt prey have narrow mouths and are slim, often brightly coloured and toxic, while ambushers have wide mouths and are broad and well-camouflaged. Caecilians do not flick their tongues, but catch their prey by grabbing it with their slightly backward-pointing teeth. The struggles of the prey and further jaw movements work it inwards and the caecilian usually retreats into its burrow. The subdued prey is gulped down whole.

When they are newly hatched, frog larvae feed on the yolk of the egg. When this is exhausted some move on to feed on bacteria, algal crusts, detritus and raspings from submerged plants. Water is drawn in through their mouths, which are usually at the bottom of their heads, and passes through branchial food traps between their mouths and their gills where fine particles are trapped in mucus and filtered out. Others have specialised mouthparts consisting of a horny beak edged by several rows of labial teeth. They scrape and bite food of many kinds as well as stirring up the bottom sediment, filtering out larger particles with the papillae around their mouths. Some, such as the spadefoot toads, have strong biting jaws and are carnivorous or even cannibalistic.

Vocalization

The calls made by caecilians and salamanders are limited to occasional soft squeaks, grunts or hisses and have not been much studied. A clicking sound sometimes produced by caecilians may be a means of orientation, as in bats, or a form of communication. Most salamanders are considered voiceless, but the California giant salamander (Dicamptodon ensatus) has vocal cords and can produce a rattling or barking sound. Some species of salamander emit a quiet squeak or yelp if attacked.

Frogs are much more vocal, especially during the breeding season when they use their voices to attract mates. The presence of a particular species in an area may be more easily discerned by its characteristic call than by a fleeting glimpse of the animal itself. In most species, the sound is produced by expelling air from the lungs over the vocal cords into one or more air sacs in the throat or at the corner of the mouth. This may distend like a balloon and acts as a resonator, helping to transfer the sound to the atmosphere, or the water at times when the animal is submerged. The main vocalisation is the male’s loud advertisement call which seeks to both encourage a female to approach and discourage other males from intruding on its territory. This call is modified to a quieter courtship call on the approach of a female or to a more aggressive version if a male intruder draws near. Calling carries the risk of attracting predators and involves the expenditure of much energy. Other calls include those given by a female in response to the advertisement call and a release call given by a male or female during unwanted attempts at amplexus. When a frog is attacked, a distress or fright call is emitted, often resembling a scream. The usually nocturnal Cuban tree frog (Osteopilus septentrionalis) produces a rain call when there is rainfall during daylight hours.

Territorial behaviour

Little is known of the territorial behaviour of caecilians, but some frogs and salamanders defend home ranges. These are usually feeding, breeding or sheltering sites. Males normally exhibit such behaviour though in some species, females and even juveniles are also involved. Although in many frog species, females are larger than males, this is not the case in most species where males are actively involved in territorial defence. Some of these have specific adaptations such as enlarged teeth for biting or spines on the chest, arms or thumbs.

In salamanders, defence of a territory involves adopting an aggressive posture and if necessary attacking the intruder. This may involve snapping, chasing and sometimes biting, occasionally causing the loss of a tail. The behaviour of red back salamanders (Plethodon cinereus) has been much studied. 91% of marked individuals that were later recaptured were within a metre of their original daytime retreat under a log or rock. A similar proportion, when moved experimentally a distance of 30 m, found their way back to their home base. The salamanders left odour marks around their territories which averaged 0.16 to 0.33 sq m in size and were sometimes inhabited by a male and female pair. These deterred the intrusion of others and delineated the boundaries between neighbouring areas. Much of their behaviour seemed stereotyped and did not involve any actual contact between individuals. An aggressive posture involved raising the body off the ground and glaring at the opponent who often turned away submissively. If the intruder persisted, a biting lunge was usually launched at either the tail region or the naso-labial grooves. Damage to either of these areas can reduce the fitness of the rival, either because of the need to regenerate tissue or because it impairs its ability to detect food.

In frogs, male territorial behaviour is often observed at breeding locations; calling is both an announcement of ownership of part of this resource and an advertisement call to potential mates. In general, a deeper voice represents a heavier and more powerful individual, and this may be sufficient to prevent intrusion by smaller males. Much energy is used in the vocalization and it takes a toll on the territory holder who may be displaced by a fitter rival if he tires. There is a tendency for males to tolerate the holders of neighbouring territories while vigorously attacking unknown intruders. Holders of territories have a “home advantage” and usually come off better in an encounter between two similar-sized frogs. If threats are insufficient, chest to chest tussles may take place. Fighting methods include pushing and shoving, deflating the opponent’s vocal sac, seizing him by the head, jumping on his back, biting, chasing, splashing, and ducking him under the water.

Defence mechanisms

Amphibians have soft bodies with thin skins, and lack claws, defensive armour, or spines. Nevertheless, they have evolved various defence mechanisms to keep themselves alive. The first line of defence in salamanders and frogs is the mucous secretion that they produce. This keeps their skin moist and makes them slippery and difficult to grip. The secretion is often sticky and distasteful or toxic. Snakes have been observed yawning and gaping when trying to swallow African clawed frogs (Xenopus laevis), which gives the frogs an opportunity to escape. Caecilians have been little studied in this respect, but the Cayenne caecilian (Typhlonectes compressicauda) produces toxic mucus that has killed predatory fish in a feeding experiment in Brazil. In some salamanders, the skin is poisonous. The rough-skinned newt (Taricha granulosa) from North America and other members of its genus contain the neurotoxin tetrodotoxin (TTX), the most toxic non-protein substance known and almost identical to that produced by pufferfish. Handling the newts does not cause harm, but ingestion of even the most minute amounts of the skin is deadly. In feeding trials, fish, frogs, reptiles, birds and mammals were all found to be susceptible. The only predators with some tolerance to the poison are certain populations of common garter snake (Thamnophis sirtalis). In locations where both snake and salamander co-exist, the snakes have developed immunity through genetic changes and they feed on the amphibians with impunity. Coevolution occurs with the newt increasing its toxic capabilities at the same rate as the snake further develops its immunity. Some frogs and toads are toxic, the main poison glands being at the side of the neck and under the warts on the back. These regions are presented to the attacking animal and their secretions may be foul-tasting or cause various physical or neurological symptoms. Altogether, over 200 toxins have been isolated from the limited number of amphibian species that have been investigated.

Poisonous species often use bright colouring to warn potential predators of their toxicity. These warning colours tend to be red or yellow combined with black, with the fire salamander (Salamandra salamandra) being an example. Once a predator has sampled one of these, it is likely to remember the colouration next time it encounters a similar animal. In some species, such as the fire-bellied toad (Bombina spp.), the warning colouration is on the belly and these animals adopt a defensive pose when attacked, exhibiting their bright colours to the predator. The frog Allobates zaparo is not poisonous, but mimics the appearance of other toxic species in its locality, a strategy that may deceive predators.

Many amphibians are nocturnal and hide during the day, thereby avoiding diurnal predators that hunt by sight. Other amphibians use camouflage to avoid being detected. They have various colourings such as mottled browns, greys and olives to blend into the background. Some salamanders adopt defensive poses when faced by a potential predator such as the North American northern short-tailed shrew (Blarina brevicauda). Their bodies writhe and they raise and lash their tails which makes it difficult for the predator to avoid contact with their poison-producing granular glands. A few salamanders will autotomise their tails when attacked, sacrificing this part of their anatomy to enable them to escape. The tail may have a constriction at its base to allow it to be easily detached. The tail is regenerated later, but the energy cost to the animal of replacing it is significant. Some frogs and toads inflate themselves to make themselves look large and fierce, and some spadefoot toads (Pelobates spp) scream and leap towards the attacker. Giant salamanders of the genus Andrias, as well as Ceratophrine and Pyxicephalus frogs possess sharp teeth and are capable of drawing blood with a defensive bite. The blackbelly salamander (Desmognathus quadramaculatus) can bite an attacking common garter snake (Thamnophis sirtalis) two or three times its size on the head and often manages to escape.

Source: Wikipedia