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

Conan O'Brien
O'Brien in 2006
Birth name	Conan Christopher O'Brien
Born	(1963-04-18) April 18, 1963 (age 49) Brookline, Massachusetts, United States[1]
Medium	Television
Nationality	American
Years active	1985–present
Genres	Improvisational comedy, sketch comedy, physical comedy, surreal humor, self-deprecation
Influences	Johnny Carson, David Letterman, Robert Smigel, Pee-Wee Herman
Spouse	Elizabeth Ann Powel (since 2002; 2 children)
Notable works and roles	The Simpsons (writer, producer, 1991–1993) Late Night with Conan O'Brien (host, 1993–2009) The Tonight Show with Conan O'Brien (host, 2009–2010) Conan (host, 2010–present)
Signature
Emmy Awards
Outstanding Writing in a Variety or Music Program 1989 Saturday Night Live 2007 Late Night with Conan O'Brien[2]

Conan Christopher O'Brien (born April 18, 1963) is an American television host, comedian, writer, producer and performer. Since November 2010 he has hosted Conan, a late-night talk show that airs on the American cable television station TBS.^[3][4]

O'Brien was born in Brookline, Massachusetts, and raised in an Irish Catholic family. He served as president of the Harvard Lampoon while attending Harvard University, and was a writer for the sketch comedy series Not Necessarily the News. After writing for several comedy shows in Los Angeles, he joined the writing staff of Saturday Night Live, and later of The Simpsons. He hosted Late Night with Conan O'Brien from 1993 to 2009, followed by seven months hosting The Tonight Show with Conan O'Brien, the only person to serve as the permanent host for both NBC programs.

Early life[link]

O'Brien was born in Brookline, Massachusetts (a suburb of Boston) to Thomas O'Brien, a physician, epidemiologist, and professor of medicine at Harvard,^[5][6][7] and Ruth O'Brien (née Reardon), an attorney and partner at the Boston firm Ropes & Gray.^[8] He is the third of six children. O'Brien's family is Irish Catholic and descends from pre-Civil War era immigrants.^[7] In a Late Night episode, O'Brien paid a visit to County Kerry, Ireland, where his ancestors originated.

O'Brien attended Brookline High School, where he served as the managing editor of the school newspaper.^[7] In his senior year, O'Brien won the National Council of Teachers of English writing contest with his short story, "To Bury the Living".^[9] After graduating as valedictorian in 1981, he entered Harvard University.^[10] At Harvard, O'Brien lived in Holworthy Hall during his freshman year^[11] and Mather House during his three upper-class years. He concentrated in history and literature and graduated magna cum laude^[12] in 1985.^[13] His senior thesis concerned the use of children as symbols in the works of William Faulkner and Flannery O'Connor.^[14] Throughout college, O'Brien was a writer for the Harvard Lampoon humor magazine. He also briefly served as the drummer in a band called "The Bad Clams".^[15] During his sophomore and junior years, he served as the Lampoon's president.^[16] At this time, O'Brien's future boss at NBC, Jeff Zucker, was serving as President of the rival The Harvard Crimson.^[17]

Career[link]

[edit] Saturday Night Live (1987–1991)

O'Brien in the offices of The Simpsons writers, 1992

O'Brien moved to Los Angeles after graduation to join the writing staff of HBO's Not Necessarily the News.^[18] He was also a writer on the short-lived The Wilton North Report.^[18] He spent two years with that show and performed regularly with improvisational groups, including The Groundlings. In January 1988, Saturday Night Live's executive producer, Lorne Michaels, hired O'Brien as a writer. During his three years on Saturday Night Live (SNL), he wrote such recurring sketches as "Mr. Short-Term Memory" and "The Girl Watchers"; the latter was first performed by Tom Hanks and Jon Lovitz.^[19] O'Brien also co-wrote the sketch, "Nude Beach", with Robert Smigel, in which the word "penis" was said or sung at least 42 times. While on a writers' strike from Saturday Night Live following the 1987–88 season, O'Brien put on an improvisational comedy revue in Chicago with fellow SNL writers Bob Odenkirk and Robert Smigel called Happy Happy Good Show.^[20][21] While living in Chicago, O'Brien briefly roomed with Jeff Garlin.^[22] In 1989, O'Brien and his fellow SNL writers received an Emmy Award for Outstanding Writing in a Comedy or Variety Series.

O'Brien, like many SNL writers, occasionally appeared as an extra in sketches; his most notable appearance was as a doorman in a sketch in which Tom Hanks was inducted into the SNL "Five-Timers Club" for hosting his fifth episode. O'Brien returned to host the show in 2001 during its 26th season. O'Brien and Robert Smigel wrote the television pilot for Lookwell starring Adam West, which aired on NBC in 1991. The pilot never went to series, but it became a cult hit. It was later screened at The Other Network, a festival of unaired TV pilots produced by Un-Cabaret; it featured an extended interview with O'Brien and was rerun in 2002 on the Trio network.

[edit] The Simpsons (1991–1993)

From 1991 to 1993, O'Brien was a writer and producer for The Simpsons^[19] and was credited as writer or co-writer of four episodes.^[23] Of all the episodes he wrote, he considers "Marge vs. the Monorail" to be his favorite.^[19] Along with this episode, he has sole writing credits on "New Kid on the Block," "Homer Goes to College," and "Treehouse of Horror IV," on which he wrote the episode wraparounds. He was an active producer during seasons 4 and 5 as well, meaning he would frequently contribute to scripts from those seasons as well as come up with story ideas, plot points, and jokes.^[24] The style of the show's comedy during this period was also influenced by his sensibilities, with "Marge vs. the Monorail" being cited by several former writers as the turning point in the show's history where more absurd and visual comedy became acceptable.^[25][26] He also developed a reputation as a "room guy," or a writer who performs comedy bits throughout the day to entertain other writers.^[25]

In his speech given at Class Day at Harvard in 2000,^[27] O'Brien credited The Simpsons with saving him, a reference to the career slump he was experiencing prior to his being hired for the show.^[28]

During his time at The Simpsons, O'Brien also had a side project working with former writing partner Robert Smigel on the script for a musical film based on the "Hans and Franz" sketch from Saturday Night Live. The film was never produced.^[29][30]

[edit] Late Night (1993–2009)

As executive producer, Lorne Michaels invited O'Brien to audition to host the successor show to Late Night with David Letterman.^[7] Premiering on September 13, 1993, Late Night with Conan O'Brien received generally unfavorable critical reviews during its first few years. The show remained on multiweek renewal cycles while NBC decided its fate.^[7] O'Brien later poked fun at the first three years of the show, when on his 10th Anniversary Special Mr. T appeared to give O'Brien a gold necklace with a giant "7" on it. When O'Brien tried to point out that he had actually been on the air for ten years, Mr. T responded, "I know that, fool; but you've only been funny for seven!"^[31]

Beginning in 1996, O'Brien and the Late Night writing team were nominated annually for the Emmy Award for Best Writing in a Comedy or Variety Series, winning the award for the first and only time in 2007. In 1997, 2000, 2002, 2003, and 2004, he and the Late Night writing staff won the Writers Guild Award for Best Writing in a Comedy/Variety Series. In 2001, he formed his own television production company, Conaco, which subsequently shared in the production credits for Late Night.^[7]

A long-running joke, which stems from the recurring segment "Conan O'Brien Hates My Homeland", is that O'Brien resembles the first female president of Finland, Tarja Halonen. After joking about this for several months (which led to his endorsement of her campaign), O'Brien traveled to Finland, appearing on several television shows and meeting President Halonen. The trip was filmed and aired as a special.^[32]

O'Brien ad-libbed the fictional website name "hornymanatee.com" on December 4, 2006, after a sketch about the fictional manatee mascot and its inappropriate webcam site.^[7] NBC opted to purchase the website domain name for $159, since the website did not previously exist. The network was concerned that the Federal Communications Commission would hold NBC liable for promoting inappropriate content if a third party were to register the domain and post such material.^[33] For a period of time, the website hosted material concerning Conan's initial manatee joke and other Tonight Show references, but today the site just redirects to NBC's main web page.

A popular recurring bit on the show was "Pale Force", a series of animated episodes in which comedian Jim Gaffigan and O'Brien are superheroes who fight crime with their "paleness". As Gaffigan introduced each new episode, O'Brien protested the portrayal of his character as cowardly, weak, and impotent. As of October 2005^[update], Late Night with Conan O'Brien had for eleven years consistently attracted an audience averaging about 2.5 million viewers.^[34] O'Brien is an avid guitarist and music listener. When Bruce Springsteen and the Sessions Band appeared on the show as musical guests, O'Brien joined the 17-piece band, along with the Max Weinberg 7 and guests Jimmy Fallon and Thomas Haden Church, playing acoustic guitar and contributing backup vocals for the song "Pay Me My Money Down". On the June 13, 2008, episode of Late Night, O'Brien simply walked onto the stage at the start of the show. Instead of his usual upbeat antics and monologue, O'Brien announced that he had just received news about the sudden death of his good friend, fellow NBC employee and frequent Late Night guest, Tim Russert. O'Brien proceeded to show two clips of his favorite Russert Late Night moments.^[35] On February 20, 2009, NBC aired the last episode of Late Night with Conan O'Brien. The show consisted of a compilation of previous Late Night clips and included a surprise appearance by former sidekick, Andy Richter. Will Ferrell, John Mayer, and the White Stripes also appeared. O'Brien ended the episode by destroying the set with an axe, handing out the pieces of the set to the audience,^[36][37] and thanking a list of people who helped him get to that point in his career. Among those thanked were Lorne Michaels, David Letterman, Jay Leno, and O'Brien's wife and children.

The Colbert / O'Brien / Stewart mock feud[link]

During the writers' strike in 2008, O'Brien staged a mock feud with Comedy Central's Jon Stewart (of The Daily Show) and Stephen Colbert (of The Colbert Report) over a dispute about which of the three were responsible for giving a "bump" to Mike Huckabee's campaign to become the Republican presidential nominee. This feud crossed over all three shows during the 2007–08 Writers Guild of America strike.^[38]

[edit] The Tonight Show (2009–2010)

In 2004, O'Brien negotiated a new contract with NBC. As part of the deal, O'Brien would take over The Tonight Show from Jay Leno in 2009.^[7] O'Brien was a guest on Jay Leno's final episode of The Tonight Show. On June 1, 2009, Will Ferrell became Conan's first Tonight Show guest on the couch and Pearl Jam appeared as his first musical guest.^[39]

During the taping of the Friday, September 25, 2009, episode of The Tonight Show, O'Brien suffered from a mild concussion after he slipped and hit his head while running a race as part of a comedy sketch with guest Teri Hatcher. He was examined at a hospital and released the same day. A rerun was aired that night, but O'Brien returned to work the following Monday and poked fun at the incident.^[40][41]

[edit] Departure from The Tonight Show

O'Brien in Atlanta, June 2010

On Thursday, January 7, 2010, NBC executive Jeff Zucker met with Jay Leno and Conan O'Brien to discuss how to get Leno out of prime time, where his ratings were lackluster, and back into late night. A proposal was made that would see O'Brien remain as host of The Tonight Show, which would be moved to 12:05 am with Leno hosting a 30-minute show at 11:35 pm^[42] On January 10, NBC Universal Television Entertainment Chairman Jeff Gaspin confirmed that The Jay Leno Show would indeed end at the start of the Winter Olympics on February 12, 2010, and be moved to 11:35 pm following the Olympics coverage.^[43] Sources familiar with the situation told the New York Post that O'Brien was unhappy with NBC's plan.^[44]

On January 12, O'Brien released this statement: "I sincerely believe that delaying The Tonight Show into the next day to accommodate another comedy program will seriously damage what I consider to be the greatest franchise in the history of broadcasting. The Tonight Show at 12:05 simply isn’t The Tonight Show."^[46][47] On January 21, 2010, it was announced that Conan had reached a deal with NBC that would see him exit The Tonight Show the next day. The deal also granted him $45 million, of which $12 million was designated for distribution to his staff, who had moved with Conan to Los Angeles from New York when he left Late Night.

The final Tonight Show with Conan aired January 22, 2010, and featured guests Tom Hanks, Steve Carell (who did an exit interview and shredded Conan's ID badge), Neil Young (singing "Long May You Run"), and Will Ferrell. For Ferrell's appearance, Conan played guitar with the band and Ferrell sang "Free Bird" while reprising his SNL cowbell. Ferrell's wife, Viveca Paulin, together with Ben Harper, Beck, and ZZ Top guitarist Billy Gibbons, also joined the band for this final performance.^[48]

Jay Leno returned to The Tonight Show following NBC's coverage of the 2010 Winter Olympic Games. Under the $45 million deal with NBC, Conan was allowed to start working for another network as soon as September 2010.^[49][50][51] Conan's rumored next networks ranged anywhere from Fox^[52] to Comedy Central.^[53][54]

[edit] After Tonight

On February 8, 2010, it was reported that O'Brien was attempting to sell his Central Park West penthouse in New York with an asking price of $35 million.^[55] He purchased the apartment in 2007 for $10 million.^[55] Two years earlier, O'Brien had purchased a home in the Brentwood section of Los Angeles for over $10.5 million.^[56] Some industry insiders have speculated that O'Brien had chosen to stay on the west coast in order to facilitate a return to late night television^[55] and because he did not want to put his children through another move.^[57]

On February 24, 2010, O'Brien attracted media attention for starting a Twitter account.^[58][59] His tweets, although primarily jokes, amounted to his first public statements since leaving The Tonight Show one month earlier.^[60] After about one hour, O'Brien's subscriber list had rocketed to over 30,000 members and approximately 30 minutes later, he was on the brink of passing 50,000 followers, already 20,000 more than the verified @jayleno account.^[61][62][63] After 24 hours, O'Brien had well over 300,000 followers.^[60] In late May 2010, he surpassed the one million mark for number of Twitter followers,^[64] and as of April 2012^[update] he has over 5.3 million followers.^[65]

O'Brien has been named to the 2010 Time 100, a list compiled by TIME of the 100 most influential people in the world as voted on by readers.^[66] After being prohibited from making television appearances of any kind until May, O'Brien spoke about the Tonight Show conflict on the CBS newsmagazine 60 Minutes on May 2, 2010.^[67] During the interview with Steve Kroft, O'Brien said the situation felt "like a marriage breaking up suddenly, violently, quickly. And I was just trying to figure out what happened." He also said he "absolutely" expected NBC to give him more of a chance and that, if in Jay Leno's position, he would not have come back to The Tonight Show. However, Conan said he did not feel he got shafted. "It's crucial to me that anyone seeing this, if they take anything away from this, it's I'm fine. I'm doing great," said O'Brien. "I hope people still find me comedically absurd and ridiculous. And I don't regret anything."^[68]

The Legally Prohibited from Being Funny on Television Tour[link]

O'Brien performing in a replica of the costume Eddie Murphy wore in Eddie Murphy Raw.

On March 11, 2010, O'Brien announced via his Twitter account that he would embark on a 30-city live tour beginning April 12, 2010, entitled "The Legally Prohibited from Being Funny on Television Tour".^[69][70] Co-host Andy Richter, along with members of the former Tonight Show Band, joined O'Brien on the tour.^[71] Max Weinberg, however, was not able to join,^[72] except for a guest appearance at one of Conan's New York City shows. VIP tickets for $695 offered the opportunity to meet O'Brien in person.^[73] On the same day, teamcoco.com—an apparently official website—was launched.^[74]

On April 12, 2010, O'Brien opened his two-month comedy tour in Eugene, Oregon, with a crowd of 2,500 and no TV cameras. The tour traveled through America's Northwest and Canada before moving on to larger cities, including Los Angeles and New York City, where he performed on the campuses that house both of the NBC-owned studios he formerly occupied. The tour ended in Atlanta on June 14. With ticket prices starting at $40, "The Legally Prohibited from Being Funny on Television Tour" was effectively sold out.^[3]

[edit] Conan (2010–present)

On April 12, 2010, just hours before the start of his tour in Eugene, Oregon, O'Brien announced that he would host a new show on cable station TBS.^[3] The show debuted on November 8, 2010,^[75] and airs Monday through Thursday beginning at 11:00 pm ET/10:00 pm CT. O'Brien's addition moved Lopez Tonight with George Lopez back one hour to midnight ET/11:00 pm CT.^[4] Refusing at first to do to Lopez what had happened to him at NBC, O'Brien agreed to join the network after Lopez called to persuade him to come to TBS.^[76] In Canada, CTV will air the show^[77] and in Turkey, CNBC-e will air the show.

Other networks that were reportedly interested in O'Brien include TBS' sister networks TNT and HBO, Fox, FX, Comedy Central, Showtime, Revision3,^[78] and even the NBC Universal-owned USA Network.^[79]

On September 1, 2010, O'Brien announced via his Twitter account and Team Coco YouTube page that the title of his new show on TBS would simply be Conan.

Television writer/producer (2002–present)[link]

In 2004, O'Brien apologized to Canadians for engaging in Quebec bashing, something which some felt reflected prejudice against Francophones.^[80] On March 7, 2006, NBC announced a new adventure/comedy series entitled Andy Barker, P.I., starring O'Brien's former sidekick, Andy Richter. O'Brien was executive producer and co-wrote the pilot. After six episodes and low ratings, the show was canceled despite being named by Entertainment Weekly as one of the Top Ten Shows of 2007.^[81] USA Network has handed out a 90-minute, cast-contingent pilot order to the medical-themed Operating Instructions from O'Brien's production banner. O'Brien will serve as an executive producer through his Conaco label. The script comes from Just Shoot Me veterans Judd Pillot and John Peaslee who will also executive produce."^[82] NBC ordered two pilots from Conaco in January 2010, the one-hour courtroom drama, Outlaw, and a half-hour comedy.^[83] Outlaw was produced in eight episodes and premiered on September 15, 2010.^[84]

Voice work and guest appearances[link]

O'Brien's first guest appearance after beginning his late-night career was on the show he used to write for, The Simpsons. He played himself in the season five episode "Bart Gets Famous", interviewing Bart Simpson during his rise to fame as a catchphrase comedian. The episode was produced after O'Brien's audition to replace David Letterman, but before he was hired for the show. O'Brien recorded his part after his own show went on the air, though he believed his show would be canceled by the time "Bart Gets Famous" aired.^[85] In 2006, he voiced himself in a short South Park scene as part of the opening of the 2006 Emmy Awards. In 2005, he provided the voice of Robert Todd Lincoln in the audio book version of Assassination Vacation by Sarah Vowell.^[86]

O'Brien has made multiple voice appearances on the Adult Swim series Robot Chicken, including the specials Robot Chicken: Star Wars, and Robot Chicken: Star Wars Episode II as the voice of the bounty hunter Zuckuss.^[87] On the TV show 30 Rock, O'Brien is depicted as an ex-boyfriend of lead character Liz Lemon, who works in the same building. In the episode "Tracy Does Conan," Conan appears as himself, awkwardly reunited with Lemon and coerced by network executive Jack Donaghy into having the character Tracy Jordan on Late Night, despite having been assaulted in Jordan's previous appearance.^[88]

O'Brien made an appearance on Futurama in the second-season episode "Xmas Story". O'Brien plays himself as a head in a jar and still alive in the year 3000. O'Brien performs a stand-up routine at a futuristic ski lodge while being heckled by Bender the robot.^[89]

O'Brien also made a cameo appearance on the U.S. version of The Office. In the episode "Valentine's Day", Michael believes that he spots former SNL cast member, Tina Fey, but has actually mistaken another woman for her. In the meantime, Conan has a quick walk-on and the camera crew informs Michael, when he returns from talking to the Tina Fey lookalike.^[90]

In January 2010, O'Brien appeared in The Simpsons 20th Anniversary Special – In 3-D! On Ice! to honor the show he had written for in the early 1990s.^[91]

O'Brien created a superhero character with veteran DC Comics animator Bruce Timm during one episode of Conan. Named "The Flaming C", the superhero bears a likeness to O'Brien, with a typically muscular superhero body and costume with chest insignia, but also with idiosyncrasies arbitrarily suggested by O'Brien like an oven mitt, a jai alai glove, marijuana leaf buckle, golf shoes, sock garters and fishnet stockings.^[92] O'Brien later aired a clip in which the character appears in Young Justice.^[93]

In 2011, O'Brien hosted the Christmas in Washington special for TBS' sister network, TNT, featuring performances from Justin Bieber, Victoria Justice, Cee-Lo Green, The Band Perry and Jennifer Hudson; with a special appearance by the Obama family.^[94]

Emmy host[link]

O'Brien hosted the 58th Primetime Emmy Awards on August 27, 2006, to critical acclaim.^[95] He had previously hosted the Primetime Emmys in 2002, and co-hosted in 2003.

Commercials and product placements[link]

According to Marketing Evolutions, the company that produces Q Scores measuring the familiarity and appeal of celebrities and brands, almost 74% of U.S. consumers are familiar with O'Brien.^[96] In spite of that high score, O'Brien hasn't done many commercials for corporate America; two recent examples are a 2009 appearance in a Super Bowl XLIII commercial for Bud Light, and one for American Express, which debuted on the same day as Conan.^[96] He donated his proceeds from the Super Bowl commercial to the Fresh Air Fund, a charity which sends inner-city New York children to the country for vacations.^[97] According to The Wall Street Journal, O'Brien was paid more than one million dollars to do the American Express commercial, which required him to spend three days in Jaipur, India filming it.^[96]

While O'Brien has done few commercials, he "does do plenty of promoting, weaving product pitches into his show"; he has said "it's increasingly incumbent to help with tie-ins [but] if it can't be funny, I'd rather go hungry."^[96]

Comedy and mannerisms[link]

On Late Night, O'Brien became known for his active and spontaneous hosting style,^[7] which has been characterized as "awkward, self-deprecating humor".^[98]

Personal life[link]

O'Brien with his wife Liza in 2007

O'Brien met Elizabeth Ann 'Liza' Powel in 2000 when she appeared on Late Night with Conan O'Brien in an advertising skit involving Foote, Cone & Belding, where she worked as senior copywriter.^[99] The couple dated for nearly 18 months before their 2002 marriage in Powel's hometown of Seattle. O'Brien and Powel have a daughter, Neve (born in 2003)^[100] and a son, Beckett (born in 2005).^[101]

O'Brien repeatedly affirms his Irish Catholic heritage on his show. On a 2009 episode of Inside the Actors Studio, he stated that both sides of his family moved to America from Ireland in the 1850s, subsequently marrying only other Irish Catholics, and that his lineage is thus 100% Irish Catholic.^[7]

He has been a staunch Democrat since casting his first vote for President in 1984 for Walter Mondale, although he considers himself a moderate on the political spectrum.^[7] O'Brien's longtime friend and former roommate at Harvard is Father Paul B. O' Brien, with whom he founded Labels Are For Jars,^[102] an antihunger organization based in Lawrence, Massachusetts, and helped open the Cor Unum meal center in 2006.^[103] The two are not related.

In January 2008, after his show was put on hold for two months owing to the strike by the Writers Guild of America, he reemerged on late-night TV sporting a beard, which guest Tom Brokaw described as making him look like "a draft dodger from the Civil War." After leaving The Tonight Show with Conan O'Brien on Jan. 22, 2010, O'Brien again grew a beard, which he kept until May 2, 2011, when it was partially shaved on the set of his TBS talk show, Conan, by Will Ferrell with battery-operated clippers (and completely shaved off-screen by a professional barber).^[104] The event was dubbed on the show as "Beardocalypse," and included a contest for fan-submitted artwork.^[105]

O'Brien purchased a $10.5-million mansion in Brentwood, Los Angeles, California, to prepare for his move there in 2009 from New York City to host The Tonight Show at Universal Studios Hollywood.^[106] As part of a long running gag, he brought his 1992 Ford Taurus SHO with him to California, showcasing it on both the inaugural episodes of The Tonight Show and Conan.

On October 21, 2011, O'Brien was ordained as a minister by the Universal Life Church,^[107] allowing him to perform a same-sex marriage while back in New York (where gay marriage is legal) to tape a week's worth of shows. The wedding, between a member of O'Brien's staff and his partner, was held on the stage of the Beacon Theatre on November 3, 2011, and broadcast on Conan.^[108]

Conan vs. Bear[link]

Conan vs. Bear is a comedic website, lampooning the prospect of Conan O'Brien fighting bears. Artists from around the world have sent in images and videos with their own take on this theme. Currently the site boasts over 400 unique renderings, as well as several original videos inspired by the site. The website was noted by O'Brien on Late Night with Conan O'Brien.^[109][110]

Victim of stalking[link]

It was reported that since September 2006, O'Brien had allegedly been stalked by Father David Ajemian of the Archdiocese of Boston, who, despite multiple warnings to stop, had been sending O'Brien letters signed as "your priest stalker" and coming in contact with O'Brien's parents. Frustrated that he had been denied a spot in the Late Night audience, Ajemian sent a letter to O'Brien stating that he flew to New York "in the dimming hope that you might finally acknowledge me." He stated in another letter, "Is this the way you treat your most dangerous fans??? You owe me big time, pal." In another letter, Ajemian seemed to make a death threat, saying, "Remember Frank Costello once dodged a bullet in your building and so can you." Ajemian then tried to forcefully enter a taping of Late Night but was caught and arrested. He was previously warned by the NBC security team to stay away from the studio. After a psychological evaluation, he was deemed fit to stand trial. He has since been bailed out of jail.^[111] He was then reported missing by his father on November 10, 2007. He was found and underwent evaluation at a hospital. It is known that the two had attended Harvard University at the same time. He was found fit to stand trial on April 4, 2008. On April 8, 2008, Ajemian pleaded guilty to stalking, stating that he "never meant to cause anxiety or to upset anyone." He was ordered to pay a $95 US$ court charge and to sign a two-year restraining order barring him from coming near O'Brien.^[112] On September 11, 2008, Ajemian checked himself out of his treatment at a hospital against the wishes of his bishop, Seán Patrick O'Malley. Cardinal O'Malley then released a statement saying that because he violated his bishop's wishes, Ajemian could no longer serve as a priest in the Catholic Church.^[113] As of late 2010, Ajemian has moved on to stalking Anthony Everett.^[114]

Nicknames[link]

A poster created by Mike Mitchell during the 2010 Tonight Show conflict displaying his "Coco" nickname.

Conan acquired the nickname "Coco" after its use in the first "Twitter Tracker" sketch during the second episode of his Tonight Show run.^[115] Guest Tom Hanks used the nickname during his subsequent interview, even getting the audience to chant it. In reaction to the moniker, Conan remarked to Hanks in jest, "If that catches on, I'll sue you."^[116][117]

In a March 23, 2011, interview with WWE Champion The Miz on Conan, The Miz dubbed Conan "The Ginja Ninja", a reference to Conan's red hair and the fact that he came back fighting to get his new late-night talk show. A week later, "Team Ginja Ninja" T-shirts were available on TeamCoco.com.^[118]

Filmography[link]

Appearances on talk shows

• Guest appearance on The Tonight Show with Jay Leno (1993,^[119] 2007, 2008, 2009)
• Guest appearance on Late Night with David Letterman (1993)
• Guest appearance on The Jon Stewart Show (1993)
• Late Night with Conan O'Brien (1993–2009)
• Guest appearance on Last Call with Carson Daly (2002)
• Guest appearance on The Daily Show with Jon Stewart (2007, 2008)
• Guest appearance on The Colbert Report (2007, 2008)
• Late Night with Jimmy Fallon (2009)
• The Tonight Show with Conan O'Brien (2009–2010)
• Guest appearance on Lopez Tonight (2010, 2011)
• Conan (November 8, 2010–)
• Guest appearance on Late Night with Jimmy Fallon (2011)
• Guest appearance on Piers Morgan Tonight (2012)
• Guest appearance on Late Show with David Letterman (2012)

Other shows

• Saturday Night Live (1991, 2001)
• The Simpsons (as himself) (1994)
• Space Ghost: Coast to Coast (as himself) (1999)
• Mr. Show with Bob and David (1995)
• The Single Guy (1996)
• Arli$$ (1996)
• Spin City (1999)
• Futurama (1999)
• Storytelling (2001)
• Vanilla Sky (2001)
• Sugar & Spice (2001) (cameo in end credits)
• Dr. Katz (as himself) (2002)
• Andy Richter Controls the Universe (2003)
• End of the Century (2003)
• Bewitched (as himself) (2005)
• Robot Chicken (2005)
• The Denial Twist (The White Stripes music video) (2005)
• O'Grady (2006)
• The Office (cameo) (2006)
• Queer Duck (as himself) (2006)
• 30 Rock (as himself) (2007, 2009)
• Robot Chicken Star Wars (as Zuckuss) (2007)
• Sesame Street (2008)
• Robot Chicken: Star Wars Episode II (as Zuckuss, Star Wars nerd) (2008)
• The Backyardigans (2009)
• 60 Minutes (interview) (2010)
• Eagleheart (himself) (2010)
• How I Met Your Mother (cameo) (2012)

Awards and nominations[link]