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; fertilization is external. In many species such as theGreat Plains toad (Bufo cognatus), the
male restrains the eggs with his back feet, holding them in place for about three minutes.
[109]
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, fertilization is internal and females give birth to fully developed juvenile frogs, except L.
larvaepartus, which give birth to tadpoles.[112][113][114]

Life cycle
Eggs / frogspawn

Frogspawn

Frogs' embryos are typically surrounded by several layers of gelatinous material. When several eggs
are clumped together, they are collectively known as frogspawn. The jelly provides support and
protection while allowing the passage of oxygen,carbon dioxide and ammonia. It absorbs moisture
and swells on contact with water. After fertilization, the innermost portion liquifies to allow free
movement of the developing embryo. In certain species, such as the Northern red-legged frog (Rana
aurora) and 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.[115] Most eggs are black or dark brown and this has the
advantage of absorbing warmth from the sun which the insulating capsule retains. The interior of
globular egg clusters of the wood frog (Rana sylvatica) has been found to be up to 6 C (11 F)
warmer than the surrounding water and this speeds up the development of the larvae. [116]
The shape and size of the egg mass is characteristic of the species. Ranids tend to produce globular
clusters containing large numbers of eggs whereas bufonids produce long, cylindrical strings. The
tiny yellow-striped pygmy eleuth (Eleutherodactylus limbatus) lays eggs singly, burying them in moist
soil.[117] 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.[118] 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.[119] The larvae developing in
the eggs can detect vibrations caused by nearby predatory wasps or snakes, and will hatch early to

avoid being eaten.[120] 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.[121]
Tadpoles

Frogspawn development.

The larvae that emerge from the eggs, known as tadpoles (or occasionally polliwogs), typically have
oval bodies and long, vertically flattened tails. 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.[122][123] Tadpoles lack eyelids and have cartilaginous skeletons, lateral line systems, gills for
respiration (external gills at first, internal gills later), and vertically flattened tails they use for
swimming.[96]
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, keratinizedstructures 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.[121] In the Pipidae, with the exception of Hymenochirus, the tadpoles
have paired anterior barbels, which make them resemble small catfish.[95] 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 tries to grasp them by the tail.[124]
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 becannibalistic. Tadpoles that develop legs early may be eaten by the others,
so late developers may have better long-term survival prospects.[125]

Tadpoles are highly vulnerable to being eaten by fish, newts, predatory diving beetles, and birds,
such as kingfishers. Some tadpoles, including those of the cane toad(Bufo marinus), 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.[126]
Metamorphosis
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.[121] The nervous system becomes adapted for hearing and stereoscopic
vision, and for new methods of locomotion and feeding.[121] 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.[121] 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.[127]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. [90]

Larva of the common frog Rana

temporaria a day before
metamorphosis

Metamorphosis stage with

deforming jaws, large eyes, and
remains of gill pouch

Young frog with a stumpy tail,

metamorphosis nearly complete

Adults

A Xenopus laevis froglet after metamorphosis.

After metamorphosis, young adults may disperse into terrestrial habitats or continue to live in water.
Almost all frog species arecarnivorous as adults, preying on invertebrates,
including arthropods, worms, snails, and slugs. A few of the larger ones may eat other frogs,
small mammals, and fish. Some frogs use their sticky tongues to catch fast-moving prey, while
others push food into their mouths with their hands. A few species also eat plant matter; the tree
frog Xenohyla truncata is partly herbivorous, its diet including a large proportion of fruit,
[128]
Leptodactylus mystaceus has been found to eat plants,[129][130] and folivory occurs in Euphlyctis
hexadactylus, with plants constituting 79.5% of its diet by volume. [131] 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.[132]

A trophic pyramid showing frogs as primary predators.

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. [133]
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. [134] 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 (Bufo marinus) has been known to survive 24 years in captivity, and the
American bullfrog (Rana catesbeiana) 14 years.[135] 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).[136]

Parental care

Male common midwife toad with eggs

(Alytes obstetricans).

Pouched frog (Assa darlingtoni)

Although care of offspring is poorly understood in frogs, up to an estimated 20% of amphibian

species may care for their young in some way.[137] The evolution of parental care in frogs is driven
primarily by the size of the water body in which they breed. Those that breed in smaller water bodies
tend to have greater and more complex parental care behaviour.[138] Because predation of eggs and
larvae is high in large water bodies, some frog species started to lay their eggs on land. Once this
happened, the desiccating terrestrial environment demands that one or both parents keep them
moist to ensure their survival.[139] The subsequent need to transport hatched tadpoles to a water body
required an even more intense form of parental care. [138]
In small pools, predators are mostly absent and competition between tadpoles becomes the variable
that constrains their survival. Certain frog species avoid this competition by making use of
smaller phytotelmata (water-filled leaf axils or small woody cavities) as sites for depositing a few
tadpoles.[140] While these smaller rearing sites are free from competition, they also lack sufficient
nutrients to support a tadpole without parental assistance. Frog species that changed from the use
of larger to smaller phytotelmata have evolved a strategy of providing their offspring with nutritive but
unfertilized eggs.[138] The femalestrawberry poison-dart frog (Oophaga pumilio) lays her eggs on the
forest floor. The male frog guards them from predation and carries water in his cloaca to keep them
moist. When they hatch, the female moves the tadpoles on her back to a water-holdingbromeliad or
other similar water body, depositing just one in each location. She visits them regularly and feeds

them by laying one or two unfertilized eggs in the phytotelma, continuing to do this until the young
are large enough to undergo metamorphosis.[141] The granular poison frog (Oophaga granulifera)
looks after its tadpoles in a similar way.[142]
Many other diverse forms of parental care are seen in frogs. The tiny male Colostethus
subpunctatus stands guard over his egg cluster, laid under a stone or log. When the eggs hatch, he
transports the tadpoles on his back to a temporary pool, where he partially immerses himself in the
water and one or more tadpoles drop off. He then moves on to another pool. [143] The malecommon
midwife toad (Alytes obstetricans) carries the eggs around with him attached to his hind legs. He
keeps them damp in dry weather by immersing himself in a pond, and prevents them from getting
too wet in soggy vegetation by raising his hindquarters. After three to six weeks, he travels to a pond
and the eggs hatch into tadpoles.[144] The tungara frog (Physalaemus pustulosus) builds a floating
nest from foam to protect its eggs from predation. The foam is made from proteins andlectins, and
seems to have antimicrobial properties.[145] Several pairs of frogs may form a colonial nest on a
previously built raft. The eggs are laid in the centre, followed by alternate layers of foam and eggs,
finishing with a foam capping.[146]
Some frogs protect their offspring inside their own bodies. Both male and female pouched
frogs (Assa darlingtoni) guard their eggs, which are laid on the ground. When the eggs hatch, the
male lubricates his body with the jelly surrounding them and immerses himself in the egg mass. The
tadpoles wriggle into skin pouches on his side, where they develop until they metamorphose into
juvenile frogs.[147] The female gastric-brooding frog (Rheobatrachus sp.) from Australia, now probably
extinct, swallows her fertilized eggs, which then develop inside her stomach. She ceases to feed and
stops secreting stomach acid. The tadpoles rely on the yolks of the eggs for nourishment. After six or
seven weeks, they are ready for metamorphosis. The mother regurgitates the tiny frogs, which hop
away from her mouth.[148] The femaleDarwin's frog (Rhinoderma darwinii) from Chile lays up to 40
eggs on the ground, where they are guarded by the male. When the tadpoles are about to hatch,
they are engulfed by the male, which carries them around inside his much-enlarged vocal sac. Here
they are immersed in a frothy, viscous liquid that contains some nourishment to supplement what
they obtain from the yolks of the eggs. They remain in the sac for seven to ten weeks before
undergoing metamorphosis, after which they move into the male's mouth and emerge. [149]

Defence

The mildly toxic Ranitomeya imitator.

Strawberry poison-dart frogcontains numerous alkaloids which deter predators.

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. [109]
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.
[150]
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.[151] 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.[6]

A common toad adopting a defensive stance.

Some frogs, such as the poison dart frogs, are especially toxic. The native people of South America
extract poison from these frogs to apply to their weapons for hunting,[152] although few species are
toxic enough to be used for this purpose. At least two non-poisonous frog species in tropical America

(Eleutherodactylus gaigei andLithodytes lineatus) mimic the colouration of dart poison frogs for selfprotection.[153][154] Some frogs obtain poisons from the ants and other arthropods they eat. [155] Others,
such as the Australian corroboree frogs (Pseudophryne corroboreeand Pseudophryne pengilleyi),
can synthesize