Amphibians Temporal range: Late Devonian–present PreЄ Є O S D C P T J K Pg N
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Superclass:	Tetrapoda
Class:	Amphibia Linnaeus, 1758
Subclasses and Orders
Subclass Labyrinthodontia – extinct Subclass Lepospondyli – extinct Subclass Lissamphibia Order Anura Order Caudata Order Gymnophiona

Amphibians are members of the class Amphibia, a group of vertebrates whose living forms include frogs, toads, salamanders, newts and caecilians. They are characterized as non-amniote, ectothermic tetrapods, meaning their eggs are not surrounded by membranes, they are cold-blooded, and they have four limbs. Most amphibians lay their eggs in water and the larvae undergo metamorphosis from a juvenile form with gills to an adult air-breathing form with lungs. Some, however, are paedomorphs that retain the juvenile water-breathing form throughout life. Mudpuppies and olms are examples of this, retaining juvenile gills into adulthood. Adult amphibians also use their skin for respiration.

The three modern orders of amphibians are placed in the subclass Lissamphibia and are the Anura (frogs and toads), Caudata (salamanders and newts) and Gymnophiona (caecilians, limbless amphibians that resemble snakes). The total number of known amphibian species is approximately 7,000.^[1] They are superficially similar to reptiles, but reptiles, along with mammals and birds, are amniotes, having impervious membranes that surround the egg. Amphibians are ecological indicators,^[2] and in recent decades there has been a dramatic decline in amphibian populations around the globe. Many species are now threatened or extinct. The study of amphibians is called batrachology.

The earliest amphibians evolved in the Devonian Period from Sarcopterygians, fish that had lungs and bony fins,^[3] features that were helpful in adapting to dry land. They diversified and became dominant during the Carboniferous and Permian periods,^[4] but were later displaced by reptiles and other vertebrates. Over time, amphibians shrunk and decreased in diversity, leaving the modern orders of Lissamphibia.

Etymology[link]

Amphibian is derived from the Ancient Greek term ἀμφίβιος (amphíbios), which means "both kinds of life", amphi meaning "of both kinds" and bio meaning "life". The term was initially used generally for animals that could live on land or in water, including seals and otters. It became restricted to what we now call amphibians around 1600 and the taxon "Amphibia" was first used in scientific classification circa 1819.^[5]

Characteristics[link]

The amphibians are a class of vertebrate animals with four limbs. They are non-amniotes which means that their eggs are not surrounded by the several membranes, some impervious, which enable mammals, reptiles and birds to reproduce on land. Amphibians typically reproduce in fresh water and are not found in the sea, with the exception of one or two frogs that live in brackish water in mangrove swamps.^[6] Most amphibians lay eggs that have a gelatinous coating which swells when it comes in contact with water. The larvae that hatch from the eggs are mostly quite dissimilar to the adult form. In the case of frogs and toads they have a large head and a dorsally flattened tail and are known as tadpoles. They are vegetarians and breathe with gills. They have no limbs at first, the back limbs thrusting through then skin at a later stage, followed by the fore limbs after which the tail is reabsorbed. After this metamorphosis the juveniles look like miniature versions of the adult. Newt and salamander larvae have long bodies and feathery gills. They are carnivorous and the front legs develop before the back ones. They do not undergo metamorphosis in the same way that frogs and toads do.^[7] The caecilians either produce live young or lay eggs in damp positions in their burrows.^[8]

Amphibians are cold blooded animals and unable to maintain their body temperature above that of their surroundings. This means that they are only able to be active when the temperature is high enough. There is great variability in the sensitivity of different species to cold. Many species hibernate in winter, going into a state of torpor either in an underground chamber or underwater. In colder climates, they may be in a state of hibernation for more than half the year. In hot weather they may aestivate underground, sometimes buried in the mud of a dried up pond, reviving when cooler weather and rain restores their habitat. An advantage of being cold blooded is that little energy is required to provide body heat. This means that they can go without food for prolonged periods without coming to harm.^[7]

Anura[link]

The order Anura includes the frogs and toads. Members of this order with smooth skins are commonly referred to as frogs while those with warted skins are known as toads. There are no taxonomic differences between these two groups but members of the family Bufonidae are known as true toads. Frogs and toads have broad heads and plump bodies with short, stout fore limbs and long hind limbs that fold underneath them. Although many species are associated with water and damp habitats, some are specialised to live in trees and in deserts. They are found worldwide.^[7]

Caudata[link]

The order Caudata includes the salamanders and one of its constituent families, Salamandridae, includes the true salamanders and the newts. Salamanders and newts have pointed heads, long cylindrical bodies, four similar sized, short legs and long tails. They may be terrestrial or aquatic but many spend part of the year in each habitat. When on land, they mostly spend the day hidden under stones or logs or in dense vegetation, emerging in the evening and night to forage for worms, insects and other invertebrates. They are found in the Holarctic region of the northern hemisphere and in Central and South America north of the Amazon Basin.^[7]

Gymnophiona[link]

The order Gymnophiona includes the caecilians. These are long, cylindrical, limbless animals that resemble snakes or worms. Their skin has circular folds which enhances their similarity to the segments of earthworms. Some are aquatic but most live underground in burrows they hollow out. Many caecilians give birth to live young, and in the animals that do not do this, the eggs may undergo metamorphosis before they hatch. Caecilians are found in tropical Africa, Asia and Central and South America.^[8]

Evolution[link]

Possible paths of Lissamphibia evolution.

The first major groups of amphibians developed in the Devonian period from lobe-finned fish similar to the modern coelacanth and lungfish,^[3] which had evolved multi-jointed leg-like fins with digits that enabled them to crawl along the sea bottom. Some fish had developed primitive lungs to help them breathe air when the stagnant pools of the Devonian swamps were lacking in oxygen. They could also use their strong fins to hoist themselves out of the water and onto dry land if circumstances required it. Eventually, their bony fins would evolve into limbs and they would become the ancestors to all tetrapods, including amphibians, reptiles, birds, and mammals. Despite being able to crawl on land, many of these prehistoric tetrapodomorph fish still spent most of their time in the water. Ichthyostega was one of these tetrapods and had four sturdy limbs, a neck, a tail with fins and a skull very similar to the lobe-finned fish, Eusthenopteron.^[3] Amphibians evolved adaptations which allowed them to stay out of the water for longer periods. However, they never developed the amniotic egg which prevented the developing embryo from drying out and which allowed early reptiles to move on to the land to reproduce. They still need to return to water or find a damp place to lay their shell-less eggs and most have a fully aquatic larval stage.^[3]

There are large gaps in the fossil record but the discovery of a batrachian from the Early Permian in Texas in 2008 provided a missing link with a lot of the characteristics of modern frogs. Molecular analysis suggests that the frog–salamander divergence took place considerably earlier than the palaeontological evidence indicates. However the date of the divergence of the caecilians deduced by molecular phylogenetics agrees with the fossil record.^[9]

The first true amphibians appeared in the Carboniferous Period, by which time they were already moving up the food chain and occupying the ecological position currently claimed by such animals as crocodiles. Amphibians were once the top land predators, sometimes reaching several meters in length, preying on the large insects on land and many types of fish in the water. During the Triassic Period, the better-adapted reptiles began to compete with amphibians, leading to the reduction of their size and importance in the biosphere. Lissamphibia, which includes all modern amphibians and is the only surviving lineage of amphibians left, could have branched off from the extinct groups Temnospondyli and/or Lepospondyli at some time between the Late Carboniferous and the Early Triassic according to the fossil record. The relative scarcity of fossil evidence does not permit an exact date,^[4] and the most recent molecular clock study based on multi-locus data suggest a Late Carboniferous–Early Permian origin of extant amphibians.^[10]

Classification[link]

Traditionally, amphibians have included all tetrapod vertebrates that are not amniotes. They are divided into three subclasses, of which two are only known as extinct subclasses:

• Subclass Labyrinthodontia† (diverse Paleozoic and early Mesozoic group)
• Subclass Lepospondyli† (small Paleozoic group, sometimes included in the Labyrinthodontia, which may actually be more closely related to amniotes than Lissamphibia)
• Subclass Lissamphibia (frogs, toads, salamanders, newts, etc.)

Of these only the last subclass includes recent species.

With the phylogenetic classification, the taxon Labyrinthodontia has been discarded as it is a paraphyletic group without unique defining features apart from shared primitive characteristics. Classification varies according to the preferred phylogeny of the author and whether they use a stem-based or a node-based classification. Traditionally, amphibians as a class are defined as all tetrapods with a larval stage, while the group that includes the common ancestors of all living amphibians (frogs, salamanders and caecilians) and all their descendants is called Lissamphibia. The phylogeny of Paleozoic amphibians is by no means satisfactory understood, and Lissamphibia may possibly include extinct groups like the temnospondyls (traditionally placed in the subclass “Labyrinthodontia”), and the Lepospondyls, and in some analysis even the amniotes. This means that advocates of phylogenetic nomenclature have removed a large number of basal Devonian and Carboniferous amphibian-type tetrapod groups that were formerly placed in Amphibia in Linnaean taxonomy and included them elsewhere under cladistic taxonomy.^[11]

All recent amphibians are included in the subclass Lissamphibia, superorder Salientia, which is usually considered a clade (which means that it is thought that they evolved from a common ancestor apart from other extinct groups), although it has also been suggested that salamanders arose separately from a Temnospondyl-like ancestor, and even that caecilians are the sister group of the advanced reptiliomorph amphibians, and thus of amniotes.^[9][12]

Authorities also disagree as to whether Salientia is a superorder that includes the order Anura, or whether Anura is a sub-order of the order Salientia. Practical considerations seem to favor using the former arrangement. The Lissamphibia, superorder Salientia, are traditionally divided into three orders, but an extinct salamander-like family, the Albanerpetontidae, is now considered part of the Lissamphibia, besides the superorder Salientia. Furthermore, Salientia includes all three recent orders plus a single Triassic proto-frog, Triadobatrachus.

Class Amphibia

• Subclass Lissamphibia
  • Family Albanerpetontidae — Jurassic to Miocene (extinct)
  • Superorder Salientia
    • Genus Triadobatrachus — Triassic (extinct) — A stem Anuran
    • Order Anura (frogs and toads): Jurassic to recent — 5,602 recent species in 48 families
    • Order Caudata or Urodela (salamanders, newts): Jurassic to recent — 571 recent species in 10 families
    • Order Gymnophiona (caecilians): Jurassic to recent — 190 recent species in 10 families

The actual number of species partly also depends on the taxonomic classification followed, the two most common classifications being the classification of the website AmphibiaWeb, University of California (Berkeley) and the classification by herpetologist Darrel Frost and The American Museum of Natural History, available as the online reference database Amphibian Species of the World.^[13] The numbers of species cited above follow Frost.

Anatomy and physiology[link]

Integumentary system[link]

The fire salamander has brightly colored yellow spots, indicating that it secretes toxins.

Amphibian skin is permeable to water and contains many mucous glands which keep the skin from drying out. Gas exchange can take place through the skin and this allows adult amphibians to hibernate at the bottom of ponds.^[7] To compensate for their thin and delicate skin, all amphibians have evolved poison glands as a defense mechanism, although toxicity varies by species. Some amphibian toxins can be lethal to humans while others have little effect.^[14] The main poison-producing glands, the paratoids, contain the neurotoxin bufotoxin and are located behind the ears of certain frogs and toads and behind the eyes of salamanders.^[15] The integumentary structure contains some typical characteristics common to terrestrial vertebrates, such as the presence of highly cornified outer layers, renewed periodically through a molting process controlled by the pituitary and thyroid glands. Local thickenings (often called warts) are common, such as those found on toads. The outside of the skin is shed periodically more or less in one piece in contrast to mammals and birds where it is shed in flakes. Amphibians often eat the sloughed skin.^[7]

The skin color of amphibians is produced by three layers of pigment cells called chromatophores. These three cell layers consist of the melanophores (occupying the deepest layer), the guanophores (forming an intermediate layer and containing many granules, producing a blue-green color) and the lipophores (yellow, the most superficial layer). The color change experienced by many species is caused by secretions from the pituitary gland. Unlike bony fish, there is no direct control by the nervous system of the pigment cells. Therefore, the color change is slower. Bright colors usually indicate that the species produces an exceptionally toxic poison.^[16]

Skeletal system[link]

The skeletal system of amphibians is structurally homologous to other tetrapods, though with a number of variations. They possess a cranium, spine, rib cage, long bones such as the humerus and femur, and short bones such as the phalanges, metacarpals, and metatarsals. Most have four limbs except for caecilians. Bones in most amphibians are hollow and lightweight.^[17][18]

The shoulder girdle of early amphibians is almost identical to that of their predecessors the osteolepiformes, except for the presence of a new dermal bone, the interclavicular (which has been lost in modern amphibians). The pelvic girdle is much more developed. In all tetrapods it consists of three main bones: the ilium in the dorsal and ventral, the pubis in the anterior and the ischium in a posterior position. The meeting point of these three bones forms the acetabulum which articulates with the femur.

Circulatory and nervous systems[link]

Amphibians have a juvenile stage and an adult stage and the circulatory systems of the two are distinct. In the juvenile (or tadpole) stage, gills are used to oxygenate blood and movement is similar to that of fish. In the adult stage, amphibians (especially frogs) lose their gills and develop lungs. They have a heart that consists of a ventricle and two atria (it may be considered a single atrium, if not totally or partially divided) that pumps oxygenated blood through arteries and deoxygenated blood through veins to the lungs. Since amphibians are cold-blooded, they must find ways to keep their blood at a constant temperature to maintain homeostasis.^[18]

The nervous system is basically the same as in other vertebrates, with a central brain, a spinal cord, and nerves throughout the body.^[18] The amphibian brain is less developed compared to that of reptiles, birds, and mammals. It consists of a cerebrum, midbrain, and cerebellum of similar sizes. The olfactory lobe is the center of the sense of smell. The cerebrum integrates behavior and learning. The optic lobe processes information from the eyes. The cerebellum is the center of muscular coordination. The medulla oblongata controls some organ functions, such as heart rate and respiration. The brain sends signals through the spinal cord and nerves to regulate activity in the rest of the body. The pineal body, known to regulate sleep patterns in humans, is thought to produce the hormones involved in hibernation and estivation in amphibians.^[19]

Digestive and excretory systems[link]

Amphibians swallow their prey whole, with some chewing done in the oral cavities of some species, so they possess voluminous stomachs. Sphincters separate the esophagus from both the oral cavity and the stomach. The relatively short esophagus is lined with cilia that help transport food and secretions to the stomach. Mucus and pepsin, a digestive enzyme, are secreted by glands lining the esophagus. The stomach is separated from the intestine by a pyloric sphincter. The duodenum controls the transport of food into the intestine from the stomach.^[18]

Amphibians possess a pancreas, liver and gall bladder. Like mammals, the liver functions as the central metabolic organ that regulates blood sugar, and also produces the final metabolic products and transports them through the vascular system to the kidneys, and finally to excretion. The liver in most amphibians is large with two lobes. The size of the liver is determined by its vital function as a glycogen and fat storage unit, and may change proportionally with the seasons with increasing or decreasing activity. In aquatic amphibians, the liver plays only a small role in processing nitrogen for excretion, and ammonia is diffused mainly through the skin and excretion. The liver of terrestrial amphibians converts ammonia to urea, a less toxic, water soluble nitrogenous compound, as a means of water conservation. In some species, urea is further converted into uric acid. The liver secretions from the liver collect in the gall bladder, and flow into the small intestine. Salamanders lack a valve separating the small intestine from the large intestine. In the small intestine, enzymes digest carbohydrates, fats, and proteins. Salt and water absorption occur in the large intestine, as well as mucous secretion to aid in the transport of fecal matter, which is passed out through the cloaca. Amphibians have two kidneys located dorsally, near the roof of the body cavity. Their job is to filter the blood of waste and transport it to the urinary bladder where it passes out of the cloacal vent.^[18]

Respiratory system[link]

The lungs in amphibians are primitive compared to those of amniotes, possessing few internal septa and large alveoli and consequently having a comparatively slow diffusion rate for oxygen entering the blood. Ventilation is accomplished by buccal pumping. Most amphibians, however, are able to exchange gases with the water or air via their skin. To enable sufficient cutaneous respiration, the surface of their highly vascularized skin must remain moist in order to allow the oxygen to diffuse at a sufficiently high rate.^[18] Because oxygen concentration in the water increases at both low temperatures and high flow rates, aquatic amphibians in these situations can rely primarily on cutaneous respiration, as in the Titicaca water frog and the hellbender salamander. In air, where oxygen is more concentrated, some small species can rely solely on cutaneous gas exchange, most famously the plethodontid salamanders, which have neither lungs nor gills. Many aquatic salamanders and all tadpoles have gills in their larval stage, with some (such as the axolotl) retaining gills as aquatic adults.^[18]

Sensory systems[link]

The eyes of amphibians have lids and associated glands and ducts. They are an improvement on invertebrate eyes and were a first step in the development of more advanced vertebrate eyes. They allow colour vision and depth of focus. In the retinas are green rods which are receptive to a wide range of wave lengths.^[20]

The tympani, or eardrums, of many frogs are external and lie just behind the eyes. There is also a patch of papilla amphibiorum in the ear which is unique to amphibians and which can detect low frequency sounds. Another unique feature is the columella-opercular complex, adjoining the auditory capsule, which is involved in the transmission of both airborne and seismic signals. The ears of salamanders and caecilians are less highly developed as they do not normally communicate with each other by sound.^[20]

Reproduction[link]

Frogspawn

For the purpose of reproduction most amphibians require fresh water. A few (e.g. Fejervarya raja) can inhabit brackish water and even survive (though not thrive) in seawater, but there are no true marine amphibians. However, there are reports of particular amphibian populations invading marine waters where their species is normally unable to survive. Such is the case with the Black Sea invasion of the natural hybrid Pelophylax esculentus reported in 2010.^[21]

Several hundred frog species in adaptive radiations (e.g., Eleutherodactylus, the Pacific Platymantines, 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.^[22]

Many amphibians exhibit different kinds of parenting behaviour. After their hatching, the tadpoles of different species of poison dart frogs (family Dendrobatidae) are carried by the adults to a suitable place where they can pass metamorphosis. Such places are the rosettes of many bromeliads in which water is gathered and used by the plant. The Surinam toad raises its young in pores at its back and after enough time they appear out of these pores fully developed. The ringed caecilian (Siphonops annulatus) has developed a unique adaptation for the purposes of reproduction. The progeny feeds on a skin layer that is specially developed by the adult. This phenomenon is known as maternal dermatophagy.

Several species have also adapted to arid and semi-arid environments, but most of them still need water to lay their eggs. Symbiosis with single celled algae that lives in the jelly-like layer of the eggs has evolved several times. The larvae of frogs (tadpoles or polliwogs) breathe with exterior gills at the start, but soon a pouch is formed that covers the gills and the front legs. Lungs are also formed quite early to assist in breathing. Newt larvae have large external gills that gradually disappear and the larvae of newts are quite similar to the adult form from an early age.

Growth and development[link]

Tadpoles

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 thyroxin concentration in the blood, which stimulates metamorphosis, and prolactin, which counteracts its effect. Specific events are dependent on threshold values for different tissues.^[23] Because most embryonic development is outside the parental body, development is subject to many adaptations due to specific ecological circumstances. For this reason tadpoles can have horny ridges for teeth, whiskers and fins. They also make use of the lateral line organ. After metamorphosis, these organs become redundant and will be reabsorbed by controlled cell death, called apoptosis. The extent of adaptations to specific ecological circumstances among amphibians is remarkable, with many discoveries still being made.^[24]

Frogs and toads[link]

Life cycle of a typical frog

With frogs and toads, the external gills of the newly hatched tadpole are covered with a gill sac after a few days, and lungs are quickly formed. Front legs are formed under the gill sac, and hind legs are visible a few days later. Following that there is usually a longer stage during which the tadpole has a vegetarian diet, scraping algae off plant surfaces with their horny tooth ridges. Tadpoles have a relatively long, spiral-shaped gut to enable them to digest this diet.^[25]

Rapid changes in the body can then be observed 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 big jaw, and its gills disappear along with its gill sac. Eyes and legs grow quickly, a tongue is formed, and all this is accompanied by associated changes in the neural networks (development of stereoscopic vision, loss of the lateral line system, etc.) All this can happen in about a day, so it is truly a metamorphosis. It isn't until a few days later that the tail is reabsorbed, due to the higher thyroxin concentrations required for tail resorption.^[25]

Newts and salamanders[link]

In newts and salamanders, there is no true metamorphosis because the larvae already feed as predators and continue doing so as adults. Their gills are never covered by a gill sac and will be reabsorbed only just before the animal leaves the water. Just as in tadpoles, their lungs are functional early, but the larvae don't make as much use of them as tadpoles do. They 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, thyroxin. External gills do not return in subsequent aquatic phases because these are completely absorbed upon leaving the water for the first time.^[26]

Caecilians[link]

Most caecilians lay eggs in burrows formed in moist soil close to water. The eggs are watched over by the female and, on hatching, the young make their way into the adjoining pond or stream. They have gill slits but no external gills. In some species, the young are produced by vivipary. Up to four eggs begin their development in the oviduct. When their yolk has been consumed, the developing larvae feed on maternal secretions and on the cells lining the oviduct, scraping them off with their deciduous teeth.^[27]

Diet[link]

Adult amphibians are predators, feeding mainly on live invertebrates. The diet mostly consists of small items of prey that do not move too fast such as beetles, caterpillars, earthworms and spiders. Many amphibians have extensible tongues with sticky tips which can be flicked forward to catch the prey. They usually swallow the food whole but may chew it lightly first in order to subdue it.^[7] The larvae of frogs and toads have a mostly vegetarian diet, using their specialised mouthparts, consisting of a horny beak edged by several rows of labial teeth, to feed on aquatic plants.^[7]

Vocalization[link]

The noises made by newts, salamanders and caecilians are limited to occasional soft squeaks, grunts or hisses. Frogs and toads however use their voice in the breeding season to attract mates. Each call is characteristic of the species, the presence of which in an area may be easier to detect by its voice than by a fleeting glimpse of the animal itself. In ponds where more than one species breed, it is important for the female to be attracted to the correct partner as male frogs, in their ardour, have been known to attempt to mate with other males, females of the wrong species or inanimate objects.^[7] Besides the attracting call, another vocalization is a release call, used by a frog suffering in this way from advances by the wrong frog. Calling bears the risk of attracting predators. Some males do not call at all but station themselves near others that are calling, with the intention of intercepting a female attracted to the sound. Although calling mainly occurs in the breeding season, sporadic calls may be heard at other times of year in some species.^[7]

The calls are made by vibrations of the larynx but are often amplified by vocal sacs which act as resonators. Sometimes these are fairly small air-filled pouches inside the mouth but in other species they are much larger and cause a ballooning of the elastic skin of the throat and are known as external vocal sacs. Many species have single sacs but members of the family Ranidae have two, one on each side of the mouth.^[7]

Defense mechanisms[link]

Amphibians have soft bodies and are relatively helpless. They are eaten by reptiles, birds and mammals when on land and by fish when in the water. Many are nocturnal and hide during the day, thereby avoiding predators that hunt by sight, apart from owls which are also nocturnal. Other amphibians use camouflage to avoid being detected. They have various colorings such as mottled browns, greys and olives to make themselves inconspicuous so that they blend into the background. Other species contain poison glands and use bright colours to warn potential predators of their toxicity. These are mostly black and yellow or black and red, like the fire salamander (Salamandra salamandra). Once a predator has tried eating one of these, it is likely to remember the coloration next time it encounters one. In some species, such as the fire-bellied toad (Bombina spp.), the warning coloration is on the belly and these animals adopt a defensive pose when attacked to exhibit the bright colors to the attacker. A few salamanders will autotomise their tails when attacked, sacrificing this part of their body to enable the main part to escape. The tail is regenerated later. 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.^[7]

Conservation[link]

The golden toad of Monteverde, Costa Rica, was among the first casualties of amphibian declines. Formerly abundant, it was last seen in 1989.

Dramatic declines in amphibian populations, including population crashes and mass localized extinction, have been noted in the past two decades from locations all over the world, and amphibian declines are thus perceived as one of the most critical threats to global biodiversity.^[28] A number of causes are believed to be involved, including habitat destruction and modification, over-exploitation, pollution, introduced species, climate change, endocrine-disrupting pollutants, destruction of the ozone layer (ultraviolet radiation has shown to be especially damaging to the skin, eyes, and eggs of amphibians), and diseases like chytridiomycosis. However, many of the causes of amphibian declines are still poorly understood, and are a topic of ongoing discussion.^[29]

A global strategy to stem the crisis has been released in the form of the Amphibian Conservation Action Plan. Developed by over 80 leading experts in the field, this call to action details what would be required to curtail amphibian declines and extinctions over the next 5 years—and how much this would cost. The Amphibian Specialist Group of the World Conservation Union (IUCN) is spearheading efforts to implement a comprehensive global strategy for amphibian conservation.^[30] Amphibian Ark is an organization that was formed to implement the ex-situ conservation recommendations of this plan, and they have been working with zoos and aquaria around the world encouraging them to create assurance colonies of threatened amphibians.^[30] One such project is the Panama Amphibian Rescue and Conservation Project that built on existing conservation efforts in Panama to create a country-wide response to the threat of chytridiomycosis rapidly spreading into eastern Panama.^[31]

On January 21, 2008, Evolutionarily Distinct and Globally Endangered (EDGE),^[32] in a statement made by chief Helen Meredith, identified nature's most endangered species: "The EDGE amphibians are amongst the most remarkable and unusual species on the planet and yet an alarming 85% of the top 100 are receiving little or no conservation attention." The top 10 EDGE listed species include the Chinese giant salamander, a distant relative of the newt, the tiny Gardiner's Seychelles, the limbless Sagalla caecilian, South African ghost frogs, lungless Mexican salamanders, the Malagasy rainbow frog, Chile's Darwin frog (Rhinoderma rufum) and the Betic Midwife Toad.^{[33][34][35][36]}

See also[link]

• Vegetal rotation
• List of amphibians

References[link]

Further reading[link]

• Carroll, Robert L. (1988). Vertebrate Paleontology and Evolution. New York: W.H. Freeman & Co.. 
• Carroll, Robert L. (2009). The Rise of Amphibians: 365 Million Years of Evolution. Baltimore: The Johns Hopkins University Press. ISBN 978-0-8018-9140-3. 

• Duellman, William E.; Linda Trueb (1994). Biology of Amphibians. Johns Hopkins University Press. ISBN 978-0-8018-4780-6. 
• Frost, Darrel R.; Taran Grant, Julián Faivovich, Raoul H. Bain, Alexander Haas, Célio F.B. Haddad, Rafael O. De Sá, Alan Channing, Mark Wilkinson, Stephen C. Donnellan, Christopher J. Raxworthy, Jonathan A. Campbell, Boris L. Blotto, Paul Moler, Robert C. Drewes, Ronald A. Nussbaum, John D. Lynch, David M. Green, Ward C. Wheeler (March 2006). "The Amphibian Tree of Life". Bulletin of the American Museum of Natural History 297: 1–291. DOI:10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2. hdl:2246/5781. 
• Pounds, J. Alan; Martín R. Bustamante, Luis A. Coloma, Jamie A. Consuegra, Michael P. L. Fogden, Pru N. Foster, Enrique La Marca, Karen L. Masters, Andrés Merino-Viteri, Robert Puschendorf, Santiago R. Ron, G. Arturo Sánchez-Azofeifa, Christopher J. Still and Bruce E. Young (January 2006). "Widespread amphibian extinctions from epidemic disease driven by global warming". Nature 439 (7073): 161–167. DOI:10.1038/nature04246. PMID 16407945. http://www.nature.com/nature/journal/v439/n7073/full/nature04246.html. 
• San Mauro, Diego; Miguel Vences, Marina Alcobendas, Rafael Zardoya and Axel Meyer (May 2005). "Initial diversification of living amphibians predated the breakup of Pangaea". American Naturalist 165 (5): 590–599. DOI:10.1086/429523. PMID 15795855. 
• San Mauro, Diego (2010). "A multilocus timescale for the origin of extant amphibians". Molecular Phylogenetics and Evolution 56 (2): 554–561. DOI:10.1016/j.ympev.2010.04.019. PMID 20399871. 
• Solomon Berg Martin, Biology
• Stuart, Simon N.; Janice S. Chanson, Neil A. Cox, Bruce E. Young, Ana S. L. Rodrigues, Debra L. Fischman, Robert W. Waller (December 2004). "Status and trends of amphibian declines and extinctions worldwide". Science 306 (5702): 1783–1786. DOI:10.1126/science.1103538. PMID 15486254. http://www.sciencemag.org/cgi/content/full/306/5702/1783. 
• S.N.Stuart, M.Hoffmann, J.S.Chanson, N.A.Cox, R.J.Berridge, P.Ramani, B.E. Young (editors), Collective work (September 2008). Threatened Amphibians of the World. Published by Lynx Edicions, in association with IUCN-The World Conservation Union, Conservation International and NatureServe.. ISBN 978-84-96553-41-5. http://www.hbw.com/lynx/en/lynx-edicions/portada-lynx/MON0017-threatened-amphibians-world.html.  776 pages

External links[link]

Wikispecies has information related to: Amphibia

The Wikibook Dichotomous Key has a page on the topic of Amphibia

• Save the Frogs!
• Amphibian Specialist Group
• AmphibiaWebEcuador
• Amphibians photos Brazil
• Amphibian Ark
• AmphibiaWeb
• EDGE
• Global Amphibian Assessment
• Amphibians of central Europe
• USGS—Online Guide for the Identification of Amphibians in North America north of Mexico
• General amphibian biology information - Living UnderWorld
• Panama Amphibian Rescue and Conservation Project
• Atlanta Botanical Garden Amphibian Conservation Program
• Amphibian vocalisations on Archival Sound Recordings
• Froglife (British conservation charity)
• Salamandridae
• A guide to Amphibians

Sir David Attenborough
Attenborough in May 2003
Born	(1926-05-08) 8 May 1926 (age 86) Isleworth, London
Nationality	British
Alma mater	Clare College, Cambridge (natural sciences) London School of Economics (social anthropology)
Occupation	Broadcaster Naturalist
Title	Member of the Order of Merit Companion of Honour Commander of the Royal Victorian Order Commander of the Order of the British Empire Fellow of the Royal Society Fellow of Zoological Society of London
Spouse	Jane Elizabeth Ebsworth Oriel (m. 1950–1997, her death)
Children	Robert Attenborough Susan Attenborough

Sir David Frederick Attenborough ( /ˈætənbərə/) OM, CH, CVO, CBE, FRS, FZS, FSA (born 8 May 1926) is a British broadcaster and naturalist.

His career as the face and voice of natural history programmes has endured for more than 50 years. He is best known for writing and presenting the nine Life series, in conjunction with the BBC Natural History Unit, which collectively form a comprehensive survey of all life on the planet. He is also a former senior manager at the BBC, having served as controller of BBC Two and director of programming for BBC Television in the 1960s and 1970s.

Attenborough is widely considered a national treasure in Britain, although he himself does not care for the term.^[1][2][3] He is a younger brother of director, producer and actor Richard Attenborough.^[4]

Early life and family[link]

Attenborough was born in Isleworth, west London, but grew up in College House on the campus of the University College, Leicester, where his father, Frederick, was principal.^[5] He is the middle of three sons (his elder brother, Richard, became an actor and his younger brother, John, an executive at Italian car manufacturer Alfa Romeo).^[6] During World War II his parents also adopted two Jewish refugee girls from Europe.^[7][8]

Attenborough spent his childhood collecting fossils, stones and other natural specimens. He received encouragement in this pursuit at age seven, when a young Jacquetta Hawkes admired his "museum". A few years later, one of his adoptive sisters gave him a piece of amber filled with prehistoric creatures; some 50 years later, it would be the focus of his programme The Amber Time Machine.

Attenborough was educated at Wyggeston Grammar School for Boys in Leicester and then won a scholarship to Clare College, Cambridge in 1945, where he studied geology and zoology and obtained a degree in natural sciences.^[9] In 1947, he was called up for national service in the Royal Navy and spent two years stationed in North Wales and the Firth of Forth.

In 1950, Attenborough married Jane Elizabeth Ebsworth Oriel; the marriage lasted until her death in 1997. The couple had two children, Robert and Susan.^[8] Robert is a senior lecturer in bioanthropology for the School of Archaeology and Anthropology at the Australian National University in Canberra.^[10]

First years at the BBC[link]

After leaving the Navy, Attenborough took a position editing children's science textbooks for a publishing company. He soon became disillusioned with the work, however, and in 1950, he applied for a job as a radio talks producer with the BBC. Although he was rejected for this job, his CV later attracted the interest of Mary Adams, head of the Talks (factual broadcasting) department of the BBC's fledgling television service. Attenborough, like most Britons at that time, did not own a television, and he had seen only one programme in his life.^[11] However, he accepted Adams' offer of a three-month training course, and in 1952 he joined the BBC full-time. Initially discouraged from appearing on camera because Adams thought his teeth were too big,^[12] he became a producer for the Talks department, which handled all non-fiction broadcasts. His early projects included the quiz show Animal, Vegetable, Mineral? and Song Hunter, a series about folk music presented by Alan Lomax.

Attenborough's association with natural history programmes began when he produced and presented the three-part series The Pattern of Animals. The studio-bound programme featured animals from London Zoo, with the naturalist Julian Huxley discussing their use of camouflage, aposematism and courtship displays. Through this programme, Attenborough met Jack Lester, the curator of the zoo's reptile house, and they decided to make a series about an animal-collecting expedition. The result was Zoo Quest, first broadcast in 1954, which Attenborough presented at short notice, due to Lester being taken ill.

In 1957, the BBC Natural History Unit was formally established in Bristol. Attenborough was asked to join it, but declined, not wishing to move from London where he and his young family were settled. Instead he formed his own department, the Travel and Exploration Unit,^[13] which allowed him to continue to front Zoo Quest as well as produce other documentaries, notably the Travellers' Tales and Adventure series.

In the early 1960s, Attenborough resigned from the permanent staff of the BBC to study for a postgraduate degree in social anthropology at the London School of Economics, interweaving his study with further filming.^[14] However, he accepted an invitation to return to the BBC as controller of BBC Two before he could finish the degree.

BBC administration[link]

Attenborough became the controller of BBC Two in March 1965, but had a clause inserted in his contract that would allow him to continue making programmes on an occasional basis. Later the same year, he filmed elephants in Tanzania, and in 1969, he made a three-part series on the cultural history of the Indonesian island of Bali. For the 1971 film A Blank on the Map, he joined the first Western expedition to a remote highland valley in New Guinea to seek out a lost tribe.

BBC Two was launched in 1964, but had struggled to capture the public's imagination. When Attenborough arrived as controller, he quickly abolished the channel's quirky kangaroo mascot and shook up the schedule. With a mission to make BBC Two's output diverse and different from that offered by other networks, he began to establish a portfolio of programmes that defined the channel's identity for decades to come. Under his tenure, music, the arts, entertainment, archaeology, experimental comedy, travel, drama, sport, business, science and natural history all found a place in the weekly schedules. Often, an eclectic mix was offered within a single evening's viewing. Programmes he commissioned included Man Alive, Call My Bluff, Chronicle, Life, One Pair of Eyes, The Old Grey Whistle Test, Monty Python's Flying Circus and The Money Programme. When BBC Two became the first British channel to broadcast in colour in 1967, Attenborough took advantage by introducing televised snooker, as well as bringing rugby league to British television on a regular basis via the BBC2 Floodlit Trophy.

One of his most significant decisions was to order a 13-part series on the history of Western art, to show off the quality of the new UHF colour television service that BBC Two offered. Broadcast to universal acclaim in 1969, Civilisation set the blueprint for landmark authored documentaries, which were informally known as "tombstone" or "sledgehammer" projects. Others followed, including Jacob Bronowski's The Ascent of Man (also commissioned by Attenborough), and Alistair Cooke's America. Attenborough thought that the story of evolution would be a natural subject for such a series. He shared his idea with Chris Parsons, a producer at the Natural History Unit, who came up with the title Life on Earth and returned to Bristol to start planning the series. Attenborough harboured a strong desire to present the series himself, but this would not be possible so long as he remained in a management post.

In 1969, Attenborough was promoted to director of programmes, making him responsible for the output of both BBC channels. His tasks, which included agreeing budgets, attending board meetings and firing staff, were now far removed from the business of filming programmes. When Attenborough's name was being suggested as a candidate for the position of Director General of the BBC in 1972, he phoned his brother Richard to confess that he had no appetite for the job. Early the following year, he left his post to return to full-time programme-making, leaving him free to write and present the planned natural history epic.

Return to broadcasting[link]

After his resignation, Attenborough became a freelance broadcaster and immediately started work on his next project, a pre-arranged trip to Indonesia with a crew from the Natural History Unit. It resulted in the 1973 series Eastwards with Attenborough, which was similar in tone to the earlier Zoo Quests but without the animal-collecting element.

On his return, he began to work on the scripts for Life on Earth. Due to the scale of his ambition, the BBC decided to partner with an American network to secure the necessary funding. While the negotiations were proceeding he worked on a number of other television projects. He presented a series on tribal art (The Tribal Eye, 1975) and another on the voyages of discovery (The Explorers, 1975). He also presented a BBC children's series about cryptozoology entitled Fabulous Animals (1975), which featured mythical creatures such as the griffin and kraken.^[15] Eventually, the BBC signed a co-production deal with Turner Broadcasting and Life on Earth moved into production in 1976.

Life series[link]

Beginning with Life on Earth in 1979, Attenborough set about creating a body of work which became a benchmark of quality in wildlife film-making and influenced a generation of documentary film-makers. The series also established many of the hallmarks of the BBC's natural history output. By treating his subject seriously and researching the latest discoveries, Attenborough and his production team gained the trust of the scientific community, who responded by allowing him to feature their subjects in his programmes. In Rwanda, for example, Attenborough and his crew were granted privileged access to film Dian Fossey's research group of mountain gorillas. Innovation was another factor in Life on Earth's success: new film-making techniques were devised to get the shots Attenborough wanted, with a focus on events and animals that were hitherto unfilmed. Computerised airline schedules, which had only recently been introduced, enabled the series to be elaborately devised so that Attenborough visited several locations around the globe in each episode, sometimes even changing continents mid-sentence. Although appearing as the on-screen presenter, he consciously restricted his pieces to camera to give his subjects top billing.

The success of Life on Earth prompted the BBC to consider a follow-up, and five years later, The Living Planet was screened. This time, Attenborough built his series around the theme of ecology, the adaptations of living things to their environment. It was another critical and commercial success, generating huge international sales for the BBC. In 1990, The Trials of Life completed the original Life trilogy, looking at animal behaviour through the different stages of life. The series drew strong reactions from the viewing public for its sequences of killer whales hunting sea lions on a Patagonian beach and chimpanzees hunting and violently killing a colobus monkey.

In the 1990s, Attenborough continued to use the "Life" moniker for a succession of authored documentaries. In 1993, he presented Life in the Freezer, the first television series to survey the natural history of Antarctica. Although past normal retirement age, he then embarked on a number of more specialised surveys of the natural world, beginning with plants. They proved a difficult subject for his producers, who had to deliver five hours of television featuring what are essentially immobile objects. The result, The Private Life of Plants (1995), showed plants as dynamic organisms by using time-lapse photography to speed up their growth.

Prompted by an enthusiastic ornithologist at the BBC Natural History Unit, Attenborough then turned his attention to the animal kingdom and in particular, birds. As he was neither an obsessive twitcher, nor a bird expert, he decided he was better qualified to make The Life of Birds (1998) on the theme of behaviour. The order of the remaining "Life" series was dictated by developments in camera technology. For The Life of Mammals (2002), low-light and infrared cameras were deployed to reveal the behaviour of nocturnal mammals. The series contains a number of memorable two shots of Attenborough and his subjects, which included chimpanzees, a blue whale and a grizzly bear. Advances in macro photography made it possible to capture natural behaviour of very small creatures for the first time, and in 2005, Life in the Undergrowth introduced audiences to the world of invertebrates.

At this point, Attenborough realised that he had spent 20 years unconsciously assembling a collection of programmes on all the major groups of terrestrial animals and plants — only reptiles and amphibians were missing. When Life in Cold Blood was broadcast in 2008, he had the satisfaction of completing the set, brought together in a DVD encyclopaedia called Life on Land. In an interview that year, Attenborough was asked to sum up his achievement, and responded:

The evolutionary history is finished. The endeavour is complete. If you'd asked me 20 years ago whether we'd be attempting such a mammoth task, I'd have said "Don't be ridiculous!" These programmes tell a particular story and I'm sure others will come along and tell it much better than I did, but I do hope that if people watch it in 50 years' time, it will still have something to say about the world we live in.^[16]

However, in 2010 Attenborough asserted that his First Life — dealing with evolutionary history before Life on Earth — should also be included within the "Life" series. In the documentary Attenborough's Journey he stated, "This series, to a degree which I really didn't fully appreciate until I started working on it, really completes the set."^[17]

Other documentaries[link]

Alongside the "Life" series, Attenborough has continued to work on other television documentaries, mainly in the natural history genre. He wrote and presented a series on man's influence on the natural history of the Mediterranean basin, The First Eden, in 1987. Two years later, he demonstrated his passion for fossils in Lost Worlds Vanished Lives.

Attenborough narrated every episode of Wildlife on One, a BBC One wildlife series which ran for 253 episodes between 1977 and 2005. At its peak, it drew a weekly audience of eight to ten million, and the 1987 episode "Meerkats United" was voted the best wildlife documentary of all time by BBC viewers.^[18] He has also narrated over 50 episodes of Natural World, BBC Two's flagship wildlife series. (Its forerunner, The World About Us, was created by Attenborough in 1969, as a vehicle for colour television.^[19]) In 1997, he narrated the BBC Wildlife Specials, each focussing on a charismatic species, and screened to mark the Natural History Unit's 40th anniversary.

As a writer and narrator, he continued to collaborate with the BBC Natural History Unit in the new millennium. Alastair Fothergill, a senior producer with whom Attenborough had worked on The Trials of Life and Life in the Freezer, was making The Blue Planet (2001), the Unit's first comprehensive series on marine life. He decided not to use an on-screen presenter due to difficulties in speaking to camera through diving apparatus, but asked Attenborough to narrate the films. The same team reunited for Planet Earth (2006), the biggest nature documentary ever made for television and the first BBC wildlife series to be shot in high definition. In 2011, Fothergill gave Attenborough a more prominent role in Frozen Planet, a major series on the natural history of the polar regions. Attenborough appeared on screen and authored the final episode, in addition to performing voiceover duties.

In 2009, he co-wrote and narrated Life, a ten-part series focussing on extraordinary animal behaviour,^[20] and narrated Nature's Great Events, which showed how seasonal changes trigger major natural spectacles.^[21]

By the turn of the millennium, Attenborough's authored documentaries were adopting a more overtly environmentalist stance. In State of the Planet (2000), he used the latest scientific evidence and interviews with leading scientists and conservationists to assess the impact of man's activities on the natural world. He later turned to the issues of global warming (The Truth about Climate Change, 2006) and human population growth (How Many People Can Live on Planet Earth?, 2009). He also contributed a programme which highlighted the plight of endangered species to the BBC's Saving Planet Earth project in 2007, the 50th anniversary of the Natural History Unit.

Attenborough is also forging a new partnership with Sky, working on documentaries for the broadcaster's new 3D network, Sky 3D. Their first collaboration was Flying Monsters 3D, a film about pterosaurs which debuted on Christmas Day of 2010.^[22] A second film, The Bachelor King 3D, followed a year later, and further collaborations are planned.

Current projects[link]

Attenborough celebrates his 60th year in broadcasting in 2012, and continues to work on a number of television, film and radio projects. The anniversary will be marked by the BBC with the screening of Attenborough: 60 Years in the Wild, a three part series which looks at how wildlife filmmaking techniques, discoveries in the field of biological sciences and our relationship with the natural world have all developed over the past 60 years.^[23]

Further collaborations with Sky 3D are also in the pipeline. First will come Kingdom of Plants 3D, a three-part series filmed at the Royal Botanical Gardens, Kew which debuts on 26 May 2012 and will be simulcast on Sky Atlantic HD.^[24] A series on the wildlife of the Galapagos Islands has also been announced^[25] and the Sky 3D film The Bachelor King 3D, which Attenborough wrote and narrated, will get a theatrical release in 2012. All Attenborough's 3D projects are being co-produced by Atlantic Productions, as is a second series of First Life, which will explore the origins of the mammals.^[26]

2012 will also herald the third series of David Attenborough's Life Stories for BBC Radio 4 and the new commission David Attenborough's Natural Curiosities on Eden.^[27]

Other work[link]

From 1983, Attenborough worked on two environmentally themed musicals with the WWF and writers Peter Rose and Anne Conlon. Yanomamo was the first, about the Amazon rainforest, and the second, Ocean World, premiered at the Royal Festival Hall in 1991. They were both narrated by Attenborough on their national tour, and recorded on to audio cassette. Ocean World was also filmed for Channel 4 and later released.

In 1990, he highlighted the case of Mahjoub Sharif as part of the BBC's Prisoners of Conscience series.^[28]

In January 2009, the BBC commissioned Attenborough to provide a series of 20 ten-minute monologues covering the history of nature. Entitled David Attenborough's Life Stories, they are broadcast on Radio 4 in the Friday night slot vacated by Alistair Cooke's Letter from America.^[29] Part of Radio 4's A Point of View strand, the talks are also available as podcasts.^[30]

He appeared in the 2009 Children's Prom at the BBC Promenade Concerts and in the Last Night of the Proms on 12 September 2009, playing a floor polisher in Sir Malcolm Arnold's "A Grand, Grand Overture" (after which he was "shot" by Rory Bremner, who was playing the gun).

Attenborough also serves on the advisory board of BBC Wildlife magazine.

Achievements, awards and recognition[link]

Attenborough's contribution to broadcasting and wildlife film-making has brought him international recognition. He has been called "the great communicator, the peerless educator"^[31] and "the greatest broadcaster of our time".^[32] His programmes are often cited as an example of what public service broadcasting should be, even by critics of the BBC, and have influenced a generation of wildlife film-makers.^[33]

Honorary titles[link]

In 2010 it was reported that Attenborough has been given 29 honorary degrees by British universities, more than any other person.^[34] In 1980, he was honoured by the Open University, with whom he has had a close association throughout his career. He also has honorary Doctor of Science awards from the University of Cambridge (1984), the University of Oxford (1988)^[35], the University of Bath (1981)^[36] and the University of St Andrews (2011).^[37] In 2006, the two eldest Attenborough brothers returned to their home city to receive the title of Distinguished Honorary Fellows of the University of Leicester, "in recognition of a record of continuing distinguished service to the University."^[38] David Attenborough was previously awarded an honorary Doctor of Letters degree by the university in 1970, and was made an honorary Freeman of the City of Leicester in 1990.

Attenborough has received the title Honorary Fellow from Clare College, Cambridge (1980), the Zoological Society of London (1998), the Linnean Society (1999), the Institute of Biology (2000) and the Society of Antiquaries (2007).

Recognition[link]

Attenborough has featured as the subject of a number of BBC television programmes. Life on Air (2002) examined the legacy of his work and Attenborough the Controller (2002) focused on his time in charge of BBC Two. He also featured prominently in The Way We Went Wild (2004), a series about natural history television presenters, and 100 Years of Wildlife Films (2007), a special programme marking the centenary of the nature documentary. In 2006, British television viewers were asked to vote for their Favourite Attenborough Moments for a UKTV poll to coincide with the broadcaster's 80th birthday. The winning clip showed Attenborough observing the mimicry skills of the superb lyrebird.

Attenborough was named as the most trusted celebrity in Britain in a 2006 Reader's Digest poll,.^[39] and the following year he won The Culture Show's Living Icon Award.^[40] He has also been named among the 100 Greatest Britons in a 2002 BBC poll and is one of the top ten "Heroes of Our Time" according to New Statesman magazine.^[41]

He has the distinction of having a number of newly discovered species and fossils being named in his honour. In 1993, after discovering that the Mesozoic reptile Plesiosaurus conybeari had not, in fact, been a true plesiosaur, the paleontologist Robert Bakker renamed the species Attenborosaurus conybeari.^[42] A fossilised armoured fish discovered at the Gogo Formation, Western Australia in 2008 was given the name Materpiscis attenboroughi, after Attenborough had filmed at the site and highlighted its scientific importance in Life on Earth.^[43] The Materpiscis fossil is believed to be the earliest organism capable of internal fertilisation.

He has also lent his name to a species of Ecuadorian flowering tree, Blakea attenboroughi, one of the world's largest-pitchered carnivorous plants, Nepenthes attenboroughii,^[44] and one of only four species of long-beaked echidna, the critically endangered Zaglossus attenboroughi, discovered by explorer and zoologist Tim Flannery in the Cyclops Mountains of New Guinea in 1998.

In September 2009, London's Natural History Museum opened the Attenborough Studio, part of its Darwin Centre development.^[45]

Awards[link]

• 1970: BAFTA Desmond Davis Award
• 1974: Commander of the Order of the British Empire (CBE)
• 1980: BAFTA Fellowship
• 1983: Fellow of the Royal Society (FRS)
• 1985: Knighthood^[46]
• 1991: Commander of the Royal Victorian Order (CVO) for producing Queen Elizabeth II's Christmas broadcast for a number of years from 1986
• 1991: Foreign Honorary Member of the American Academy of Arts and Sciences^[47]
• 1996: Companion of Honour (CH) "for services to nature broadcasting"
• 1998: International Cosmos Prize
• 2003: Michael Faraday Prize awarded by the Royal Society
• 2004: Descartes Prize for Outstanding Science Communication Actions
• 2004: Caird Medal of the National Maritime Museum
• 2005: Order of Merit (OM)^[48]
• 2005: Nierenberg Prize for Science in the Public Interest
• 2006: National Outdoor Book Award (Nature and the Environment category), Life in the Underground
• 2006: National Television Awards Special Recognition Award
• 2006: Institute of Ecology and Environmental Management^[49] - Institute Medal in recognition of his outstanding contribution to the public perception and understanding of ecology
• 2006: The Culture Show British Icon Award
• 2007: British Naturalists' Association Peter Scott Memorial Award
• 2009: BAFTA Specialist factual award for Life In Cold Blood
• 2009: Prince of Asturias Award^[50]
• 2010: Fonseca Prize
• 2010: Queensland Museum Medal^[51][52]
•  ???? RSPB Medal^[53][54]
• 2011: Society for the History of Natural History Founders' Medal

Views and advocacy[link]

Environmental causes[link]

Attenborough's programmes have often included references to the impact of human society on the natural world. The last episode of The Living Planet, for example, focuses almost entirely on humans' destruction of the environment and ways that it could be stopped or reversed. Despite this, he has been criticised for not giving enough prominence to environmental messages. Some environmentalists feel that programmes like Attenborough's give a false picture of idyllic wilderness and do not do enough to acknowledge that such areas are increasingly encroached upon by humans.^[55]

However, his closing message from State of the Planet was forthright:

The future of life on earth depends on our ability to take action. Many individuals are doing what they can, but real success can only come if there's a change in our societies and our economics and in our politics. I've been lucky in my lifetime to see some of the greatest spectacles that the natural world has to offer. Surely we have a responsibility to leave for future generations a planet that is healthy, inhabitable by all species.

Attenborough has subsequently become more vocal in his support of environmental causes. In 2005 and 2006, he backed a BirdLife International project to stop the killing of albatross by longline fishing boats.^[56] He gave public support to WWF's campaign to have 220,000 square kilometres of Borneo's rainforest designated a protected area.^[57] He also serves as a vice-president of BTCV, vice-president of Fauna and Flora International, president of Butterfly Conservation and president of Leicestershire and Rutland Wildlife Trust. In 2003 he launched an appeal to create a rainforest reserve in Ecuador in memory of Christopher Parsons, the producer of Life on Earth and a personal friend, who had died the previous year. The same year, he helped to launch ARKive,^[58] a global project instigated by Parsons to gather together natural history media into a digital library. ARKive is an initiative of Wildscreen, of which Attenborough is a patron.^[59] He later became patron of the World Land Trust, and an active supporter. He supported Glyndebourne in their successful application to obtain planning permission for a wind turbine in an Area of Outstanding Natural Beauty, and gave evidence at the planning inquiry arguing in favour of the proposal.

Attenborough has repeatedly said that he considers human overpopulation to be the root cause of many environmental problems. In The Life of Mammals, he made a plea for humans to curb population growth so that other species will not be crowded out. In 2009, he became a patron of Population Matters, (formerly known as the Optimum Population Trust), a UK charity advocating sustainable human populations.

He has written and spoken publicly about the fact that, despite past scepticism, he believes the Earth's climate is warming in a way that is cause for concern, and that this can likely be attributed to human activity.^[60] He summed up his thoughts at the end of his 2006 documentary "Can We Save Planet Earth?" as follows:

In the past, we didn't understand the effect of our actions. Unknowingly, we sowed the wind and now, literally, we are reaping the whirlwind. But we no longer have that excuse: now we do recognise the consequences of our behaviour. Now surely, we must act to reform it — individually and collectively, nationally and internationally — or we doom future generations to catastrophe.

In a 2005 interview with BBC Wildlife magazine, Attenborough said he considered George W. Bush to be the era's top "environmental villain". In 2007, he further elaborated on the USA's consumption of energy in relation to its population. When asked if he thought America to be "the villain of the piece", he responded:

I don't think whole populations are villainous, but Americans are just extraordinarily unaware of all kinds of things. If you live in the middle of that vast continent, with apparently everything your heart could wish for just because you were born there, then why worry? [...] If people lose knowledge, sympathy and understanding of the natural world, they're going to mistreat it and will not ask their politicians to care for it.^[61]

Religion and creationism[link]

In a December 2005 interview with Simon Mayo on BBC Radio Five Live, Attenborough stated that he considers himself an agnostic.^[62] When asked whether his observation of the natural world has given him faith in a creator, he generally responds with some version of this story, making reference to the Loa loa parasitic worm:

My response is that when Creationists talk about God creating every individual species as a separate act, they always instance hummingbirds, or orchids, sunflowers and beautiful things. But I tend to think instead of a parasitic worm that is boring through the eye of a boy sitting on the bank of a river in West Africa, [a worm] that's going to make him blind. And [I ask them], 'Are you telling me that the God you believe in, who you also say is an all-merciful God, who cares for each one of us individually, are you saying that God created this worm that can live in no other way than in an innocent child's eyeball? Because that doesn't seem to me to coincide with a God who's full of mercy'.^[63]

He has explained that he feels the evidence all over the planet clearly shows evolution to be the best way to explain the diversity of life, and that "as far as [he's] concerned, if there is a supreme being then he chose organic evolution as a way of bringing into existence the natural world."

In a BBC Four interview with Mark Lawson, Attenborough was asked if he at any time had any religious faith. He replied simply, "No."^[64] He has also said "It never really occurred to me to believe in God".^[65]

In 2002, Attenborough joined an effort by leading clerics and scientists to oppose the inclusion of creationism in the curriculum of UK state-funded independent schools which receive private sponsorship, such as the Emmanuel Schools Foundation.^[66] In 2009, Attenborough stated that the Book of Genesis, by saying that the world was there for people to dominate, had taught generations that they can "dominate" the environment, and that this has resulted in the devastation of vast areas of the environment.^[67] Attenborough further explained to the science journal Nature, "That's why Darwinism, and the fact of evolution, is of great importance, because it is that attitude which has led to the devastation of so much, and we are in the situation that we are in."^[68][69]

Also in early 2009, the BBC broadcast an Attenborough one-hour special, Charles Darwin and the Tree of Life. In reference to the programme, Attenborough stated that "People write to me that evolution is only a theory. Well, it is not a theory. Evolution is as solid a historical fact as you could conceive. Evidence from every quarter. What is a theory is whether natural selection is the mechanism and the only mechanism. That is a theory. But the historical reality that dinosaurs led to birds and mammals produced whales, that's not theory."^[70] He strongly opposes creationism and its offshoot "intelligent design", saying that a survey that found a quarter of science teachers in state schools believe that creationism should be taught alongside evolution in science lessons was "really terrible".^[70]

In March 2009, Attenborough appeared on Friday Night with Jonathan Ross. Attenborough stated that he felt evolution did not rule out the existence of a God and accepted the title of agnostic saying, "My view is: I don't know one way or the other but I don't think that evolution is against a belief in God."^[71]

Attenborough has joined evolutionary biologist Richard Dawkins and other top scientists in signing a campaign statement coordinated by the British Humanist Association (BHA). The statement calls for "creationism to be banned from the school science curriculum and for evolution to be taught more widely in schools."^[72]

BBC and public service broadcasting[link]

Attenborough is a lifelong supporter of the BBC, public broadcasting and the television licence. He has said:

PSB, to me, is not about selecting individual programme strands here or there, financing them from some outside source and then foisting them upon commercial networks. Public Service Broadcasting, watched by a healthy number of viewers, with programmes financed in proportion to their intrinsic needs and not the size of the audience, can only effectively operate as a network — a network whose aim is to cater for the broadest possible range of interests, popular as well as less popular, a network that measures its success not only by its audience size but by the range of its schedule.^[73]

Public service broadcasting is one of the things that distinguishes this country and makes me want to live here. I have spent all my life in it. I would be very distressed if public service broadcasting was weakened. I have been at the BBC since 1952, and know the BBC is constantly being battered. It is today.^[74]

If you could demonstrate that the BBC was grossly extravagant there might be a case for saying OK take it away. But in fact the BBC per minute in almost every category is as cheap as you can find anywhere in the world and produces the best quality. If you take the money away, which part of the BBC will you remove? The BBC has gone through swingeing staff cuts. It has been cut to the bone, if you divert licence fee money elsewhere, you cut quality and services. There is always that threat from politicians who will say your licence fee is up for grabs. We will take it. There is a lot of people who want to see the BBC weakened. They talk of this terrible tax of the licence fee. Yet it is the best bargain that is going. Four radio channels and god knows how many TV channels. It is piffling.^[74]

There have always been politicians or business people who have wanted to cut the BBC back or stop it saying the sort of things it says. There's always been trouble about the licence and if you dropped your guard you could bet our bottom dollar there'd be plenty of people who'd want to take it away. The licence fee is the basis on which the BBC is based and if you destroy it, broadcasting... becomes a wasteland.^[75]

Attenborough expressed regret at some of the changes made to the BBC in the 1990s by Director-General John Birt, who introduced an internal market at the corporation, slimmed and even closed some departments and outsourced much of the corporation’s output to private production companies, in line with the Broadcasting Act 1990. He has said:

There is no question but that Birtism . . . has had some terrible results. On the other hand, the BBC had to change. Now it has to produce programmes no one else can do. Otherwise, forget the licence fee.^[76]

The Bristol Unit has suffered along with the rest of the BBC from recent staff cuts. Yet it remains confident in the belief that the BBC will maintain it, in spite of the vagaries of fashion, because the Corporation believes that such programmes deserve a place in the schedules of any broadcaster with pretensions of providing a Public Service. In due course, similar specialist Units were also established in London, in order to produce programmes on archaeology and history, on the arts, on music and on science. They too, at one time, had their successes. But they have not survived as well as the Unit in Bristol. The statutory requirement that a certain percentage of programmes must come from independent producers has reduced in-house production and the Units necessarily shrank proportionately in size. As they dwindled, so the critical mass of their production expertise has diminished. The continuity of their archives has been broken, they have lost the close touch they once had worldwide with their subjects and they are no longer regarded internationally as the centres of innovation and expertise that they once were.^[77]

When Birt gets up and says the whole of the BBC was a creative mess and it was wasteful, I never saw any evidence of that. I absolutely know it wasn’t so in my time. Producers now spend all their time worrying about money, and the thing has suffered for it.^[78]

In 2008, he criticised the BBC’s television schedules:

I have to say that there are moments when I wonder — moments when its two senior networks, first set up as a partnership, schedule simultaneously programmes of identical character, thereby contradicting the very reason that the BBC was given a second network. Then there are times when both BBC One and BBC Two, intoxicated by the sudden popularity of a programme genre, allow that genre to proliferate and run rampant through the schedules. The result is that other kinds of programmes are not placed, simply because of a lack of space. Do we really require so many gardening programmes, make-over programmes or celebrity chefs? Is it not a scandal in this day and age, that there seems to be no place for continuing series of programmes about science or serious music or thoughtful in-depth interviews with people other than politicians?^[73]

In 2009, Attenborough commented on the general state of British television, describing the newly introduced product placement on commercial television as something he considered an "appalling" idea 20 years earlier:

I think it's in great trouble. The whole system on which it was built — a limited number of networks, with adequate funding — is under threat. That funding is no longer there. As stations proliferate, so audiences are reduced. The struggle for audiences becomes ever greater, while money diminishes. I think that's a fair recipe for trouble. Inevitably, this has an impact on the BBC … Fortunately, the BBC doesn't think natural history programmes must compete with Strictly Come Dancing in terms of audience. The BBC says, 'Make proper, responsible natural history programmes.'^[79]

Other causes[link]

In May 2005, Attenborough was appointed as patron of the UK's Blood Pressure Association, which provides information and support to people with hypertension.^[80]

Attenborough is also an honorary member of BSES Expeditions, a youth development charity that operates challenging scientific research expeditions to remote wilderness environments.

Parodies and artistic portrayals[link]

Attenborough's accent and hushed, excited delivery have been the subject of frequent parodies by comedians, most notably Spike Milligan, Marty Feldman and The Goodies. Attenborough is portrayed by Michael Palin in the final episode of Monty Python's Flying Circus, where he searches the African jungle for the legendary Walking Tree of Dahomey (Quercus Nicholas Parsonus), sweating excessively and accompanied by native guides wearing saxophones. David Attenborough has also been parodied in a series of GEICO insurance commercials, showing a nature show host, clearly patterned on Attenborough.

"Time Flies", a sketch by David Ives, features a pair of anthropomorphic mayflies engaging in a courtship ritual, while watching themselves on television in a documentary narrated by David Attenborough.^[81]

The character of Nigel Thornberry, a nature documentarian on Nickelodeon's The Wild Thornberrys is strongly influenced by Attenborough.

He has also been parodied by the Australia 1980s sketch show The Comedy Company where Ian McFadyen portrays a character called David Rabbitborough, who goes about examining the "wildlife" of various Australian urban stereotypes.

The video game Discworld, based on the series of books by Terry Pratchett, parodies his unique delivery to explain different aspects of the Discworld Universe, such as L-Space.

Mythbusters host Adam Savage often imitates Attenborough when speaking about his co-host Jamie Hyneman, which Jamie verified in the YouTube Special Episode.

Filmography[link]

Attenborough is known foremost for writing and presenting the ten Life series, which are presented in chronological order below:

• Life on Earth (1979)
• The Living Planet (1984)
• The Trials of Life (1990)
• Life in the Freezer (1993)
• The Private Life of Plants (1995)
• The Life of Birds (1998)
• The Life of Mammals (2002)
• Life in the Undergrowth (2005)
• Life in Cold Blood (2008)
• First Life (2010)

His voice is synonymous with wildlife documentaries for British audiences, and the principal series with which his narration is associated include:

• Wildlife on One (1977–2005)
• BBC Wildlife Specials (1995–2008)
• The Blue Planet (2001)
• Planet Earth (British version) (2006)
• Nature's Great Events (2009)
• Life (2009)
• Frozen Planet (2011)

Books[link]

Bibliography[link]

• Zoo Quest to Guyana (Lutterworth Press, 1956)
• Zoo Quest for a Dragon (Lutterworth Press, 1957)
  • (book club edition with 85 extra pages, Quest for the Paradise Birds, 1959)
• Zoo Quest in Paraguay (Lutterworth Press, 1959)
  • paperback (Penguin Books, 1982)
• Quest in Paradise (1960)
• People of Paradise (Harper & Brothers, 1960)
• Zoo Quest to Madagascar (1961)
• Quest Under Capricorn (1963)
• Fabulous Animals (BBC, 1975) ISBN 0-563-17006-9
• The Tribal Eye (1976)
• Life on Earth (1979)
• Discovering Life on Earth (1981)
• Journeys to the past: Travels in New Guinea, Madagascar, and the northern territory of Australia (1983) Penguin Books ISBN 0-14-00.64133
• The Living Planet (1984)
• The First Eden: The Mediterranean World and Man (1987)(Little Brown & Co (T); 1st American ed edition (March 1990))
• The Atlas of the Living World (1989)
• The Trials of Life (Collins, 1990) ISBN 0-00-219912-2
• The Private Life of Plants (BBC Books, 1994) ISBN 0-563-37023-8
• The Life of Birds (BBC Books, 1998) ISBN 0-563-38792-0
• The Life of Mammals (BBC Books, 2002) ISBN 0-563-53423-0
• Life on Air: Memoirs of a Broadcaster (autobiography; 2002) ISBN 0-563-53461-3
  • paperback: ISBN 0-563-48780-1
  • updated edition: ISBN 978-1-84607-652-7
• Life in the Undergrowth (BBC Books, 2005) ISBN 0-563-52208-9
• Amazing Rare Things - The Art of Natural History in the Age of Discovery with Susan Owens, Martin Clayton and Rea Alexandratos (The Royal Collection, 2007) Hardback - ISBN 978-1-902163-46-8; Softback - ISBN 978-1-902163-99-4
• Life in Cold Blood (BBC Books, 2007) ISBN 978-0-563-53922-3
• Life Stories, (HarperCollins Publishers, 2009) ISBN 978-0-00-733883-2
• New Life Stories, (Collins, 2011) ISBN 978-0-00-742512-9

Introductions[link]

Attenborough has written the introduction or foreword for a number of books, including:

• Buxton, Cindy; Annie Price (1983). Survival South Atlantic. London :: Granada,. pp. xiii,237p.,[64]p. of plates : ill(some col.), maps,col.ports. ; 26cm.. ISBN 0-246-12087-8. 
• African Jigsaw: A Musical Entertainment, Peter Rose and Anne Conlon (published: 1986, Weinberger)
• Tomorrow Is Too Late, Various (The Macmillan Press, 1990)
• Life in the Freezer: Natural History of the Antarctic, Alastair Fothergill (BBC Books, 1993), ISBN 0-563-36431-9
• Birds of Paradise: Paradisaeidae (Bird Families of the World series) Clifford B. Frith, Bruce M. Beehler, William T. Cooper (Illustrator) (Oxford University Press, 1998) ISBN 0-19-854853-2
• The Blue Planet, Andrew Byatt, Alastair Fothergill, Martha Holmes (BBC Books, 2001) ISBN 0-563-38498-0.
• Light on the Earth (BBC Books, 2005), two decades of winning images from the Wildlife Photographer of the Year competition, ISBN 0-563-52260-7
• Planet Earth, Alastair Fothergill (BBC Books, 2006), ISBN 0-563-52212-7

Audio recordings[link]

• Tarka the Otter by Henry Williamson (available on audiocassette, 1978)
• Yanomamo (musical entertainment, 1983) by Peter Rose and Anne Conlon; on-stage narration and published audio recording
• Ocean World (musical entertainment, 1990) by Peter Rose and Anne Conlon; on-stage narration (including at The Royal Festival Hall), for audio recording and video broadcast (both published)
• Peter and the Wolf for BBC Music Magazine (free CD with the June 2000 issue).

In addition, Attenborough has recorded some of his own works in audiobook form, including Life on Earth, Zoo Quest for a Dragon and his autobiography Life on Air: Memoirs of a Broadcaster.

References[link]