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Common Name:

Baltimore Oriole

Scientific Name:

Icterus galbula

Size relative to a teacup:

The Baltimore oriole is Maryland's official state bird. This popular animal has also been the namesake of the state's professional baseball team, the Baltimore Orioles, since the late 19th century. Male orioles have brilliant orange-golden underparts and shoulder patches, with black wings and a black head. Females are not as brightly colored. Though they are partially orange, they also have and brownish-olive plumage.

Population Range

Baltimore orioles inhabit Maryland and the rest of the eastern United States only in the summer months. (They are also found, less commonly, in the central U.S. and Canada). In winter, some of these migrating birds live in the southeastern U.S., but most fly further afield in search of neotropical climates.

Diet

These attractive birds frequent woodlands and eat common creatures including caterpillars and insects supplemented by fruits and berries. The Baltimore oriole's appetite for caterpillars may help protect forests from some destructive pests. In the backyard, they can be enticed to visit feeders with oranges, nectars, or peanut butter.

Nesting

Each spring a female oriole constructs a hanging nest at the end of a tree branch. From this perch, she will guard her eggs (typically four) for about two weeks. When the young birds hatch, both parents will feed and watch over them for an additional two weeks.

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Common Name: Baltimore Oriole Scientific Name: Icterus galbula Size relative to a teacup: The Baltimore oriole is Maryland's official state bird. This popular animal has also been the namesake of the state's professional baseball team, the Baltimore Orioles, since the late 19th century.

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What is a hornbill?

Hornbills are tropical birds named for their unusually large, curved bills, some of which are adorned with a large casque, a hollow structure made from keratin.

Casques, which vary in size and color, serve a number of functions, such as attracting mates and resonating sound, making the birds' territorial calls more audible. Casques may also provide support when the birds use their beaks to peel off tree bark or dig in soil for insects.

Some hornbill species don't have casques, such as the red-billed hornbill, which was the model for the character Zazu in The Lion King. (The Walt Disney Company is majority owner of National Geographic Partners.)

Most of the 62 hornbill species spend their time in the trees, except for the family Bucorvidae, the ground-dwelling hornbills.

The largest hornbill, southern Africa's southern ground hornbill, stands at three feet tall with a four-foot wingspan. The red-billed dwarf hornbill, found in East and Central Africa, is the smallest species, at just under one foot.

Hornbills inhabit a variety of landscapes, including forests, savannas, shrublands, and rocky and high-altitude areas. Almost all species nest in natural cavities, mostly in trees but sometimes on rocky cliffs. These omnivores may dine on anything from fruit and seeds to insects, reptiles, amphibians, and small mammals.

Courtship and reproduction

There are many courtship behaviors, such as clattering bills together or alternating calls in a sort of duet.

Many hornbill species are considered monogamous, but only Montiero's hornbill of southern Africa is genetically monogamous, meaning neither the male nor the female mate with any other birds. A male will often bring food to its mate, sometimes for an entire year.

Once mated, most hornbill pairs will find an existing tree cavity, and the male will seal the female inside using its own dung mixed with soil or regurgitated food. The male will leave only a small slit, through which he'll will deliver her food. Protected from predators in this private suite, she'll generally lay two eggs, which will incubate for about 42 days.

Usually only one chick will hatch or, if both emerge, only one chick will usually survive. Great hornbills of Southeast Asia and Malayan wreathed hornbills have a 40-day egg incubation period, and will brood chicks for as long as four months in the mother's lair.

About a week before the chick is ready to strike out on its own, the female will tear the door down and leave the shelter. The chick will rebuild the protective wall and stay inside by itself, while both parents provide their offspring with food. Chicks will then fledge after roughly 80 days.

Southern ground hornbills have a different strategy, living in small flocks of up to nine individuals. Only the largest male and female-the alpha pair-will breed, while the rest of the birds help rear the offspring.

Conservation status

Of the 62 hornbill species, 51 species are decreasing in population, eight are stable, and three are data deficient.

The forest-dwelling helmeted hornbill of Southeast Asia is critically endangered due to an illegal trade in the birds' casques, which has led to an increase in poaching of wild birds. The helmeted hornbills' casques are softer than ivory and easy to carve into trinkets, jewelry, and other decorative items.

Also critically endangered are the rufous-headed hornbill and Sulu hornbill, both of the Philippines. There are only about 50 Sulu hornbills left in the wild, victims of hunting, human land use, and climate change. In 2019, the International Union for Conservation of Nature launched a decade-long effort to save the Sulu.

- San Diego Zoo Wildlife Alliance

- Museum of Osteology

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Hornbills are tropical birds named for their unusually large, curved bills, some of which are adorned with a large casque, a hollow structure made from keratin. Casques, which vary in size and color, serve a number of functions, such as attracting mates and resonating sound, making the birds' territorial calls more audible.

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The common emu may not be able to soar, but for such a big bird it sure can run. This flightless bird has small wings relative to the size of its body. Its long, powerful legs, though, allow it to run up to about 30 miles (50 kilometers) per hour. Each emu foot has three forward-facing toes that allow it to grip the ground, thrusting the bird forward. A powerful kick is also handy for keeping predators at bay.

Territory

Emus are members of the ratite family, which also includes ostriches, cassowaries, and rheas. They are among the largest birds in the world, and they're found primarily in Australia, but also in New Guinea, Indonesia, Solomon Islands, and the Philippines. Emus beat the heat in hot, arid habitats: Their grayish-to-brownish body feathers provide almost complete protection from solar radiation. Though equipped with this natural sunscreen, they also live in grasslands, savannas, and forests, preferring areas with access to standing water.

Emus range over large areas, foraging on fruits, seeds, plant shoots, small animals, animal droppings, and insects. They mate and nest over the Australian winter, and it's not always a loving affair-females have been known to fight viciously over unpaired males.

Reproduction

Mating pairs stay together for up to five months, after which females lay large, emerald-green eggs in expansive ground nests. The males incubate the eggs for about seven weeks without drinking, feeding, defecating, or leaving the nest. The females, meanwhile, have often moved on, sometimes mating with a different male in the same season. Chicks stay with dad for about four months, until they are able to eat on their own.

Threats

Predators of the emu include dingoes and wedge-tailed eagles. Snakes and other nest-raiders devour emu eggs, but they're not the only ones. Humans farm emus for their meat and eggs, too. One emu egg can make an omelet big enough to feed four to six adults.

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The common emu may not be able to soar, but for such a big bird it sure can run. This flightless bird has small wings relative to the size of its body. Its long, powerful legs, though, allow it to run up to about 30 miles (50 kilometers) per hour.

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Scientific Name:

Cyanocitta cristata

Size:

10 to 12 inches; wingspan: 13 to 17 inches

Weight:

2.5 to 3.5 ounces

Size relative to a teacup:

Blue jays are natural forest dwellers, but they are also highly adaptable and intelligent birds. They are a familiar and noisy presence around many North American bird feeders. The blue jay's "Jay! Jay!" call is only one of a wide variety of sounds the bird employs-including excellent imitations of several hawk calls.

Diet

Blue jays are sometimes known to eat eggs or nestlings, and it is this practice that has tarnished their reputation. In fact, they are largely vegetarian birds. Most of their diet is composed of acorns, nuts, and seeds-though they also eat small creatures such as caterpillars, grasshoppers, and beetles. Blue jays sometimes store acorns in the ground and may fail to retrieve them, thus aiding the spread of forests.

Behavior and Migration

Common in much of eastern and central North America, blue jays are gradually extending their range to the Northwest. They are fairly social and are typically found in pairs or in family groups or small flocks. Most northern birds head south for the winter and join in large flocks of up to 250 birds to make the long journey. However, this migration is a bit of a mystery to scientists. Some birds winter in all parts of the blue jay's range, and some individual birds may migrate one year and not the next. It is unclear what factors determine whether each blue jay or family decides to migrate.

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Scientific Name: Cyanocitta cristata Size: 10 to 12 inches; wingspan: 13 to 17 inches Weight: 2.5 to 3.5 ounces Size relative to a teacup: Blue jays are natural forest dwellers, but they are also highly adaptable and intelligent birds. They are a familiar and noisy presence around many North American bird feeders.

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Watch to discover interesting facts about animals from all over the world.

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Watch to discover interesting facts about animals from all over the world.

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Geckos can't blink ... so how do they clean their eyeballs? You probably sleep eight hours a night ... but which animal sleeps the most? Get the answers to these questions and more in this series of weird (but true!) videos!

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Watch what happens when animals are caught on camera doing wacky activities! This music video series captures the funny moments animals have in their day-to-day lives.

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A Day of Seals and Sea Lions | The Aquarium: A Deeper Dive

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Illegal Pet Trade: Help Stop Wildlife Trafficking

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Illegal Pet Trade: Help Stop Wildlife Trafficking

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The Science of Animal Movement: How Snakes Slither

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The Science of Animal Movement: How Snakes Slither

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National Geographic Kids Reader | Dinosaurs | Books Read Aloud #dinosaurbooksforkids

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Why the Dinosaurs’ Extinction is an Ongoing Puzzle | Nat Geo Explores

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The Rare and Exotic Animals " White Lions"-[HD]National Geographic[Full Documentary]

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The Rare and Exotic Animals " White Lions"-[HD]National Geographic[Full Documentary]

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The Rare and Exotic Animals " White Lions"-[HD]National Geographic[Full Documentary]

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Mammals for Kids | Learn all about the unique characteristics of mammals and what mammals are!

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Find Out About Insects! | Nat Geo Kids Insects Playlist

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Rare Pigs Caught on Video for the First Time | National Geographic

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Although electric eels have the power to be the bullies of the Amazon, they are actually not very aggressive animals. The eel uses its shock to stun prey and keep predators at bay.

Electric eels are nocturnal, live in muddy, dark waters, and have poor eyesight. So instead of using eyes, electric eels emit a weak electric signal, which they use like radar to navigate, to find a mate, and to find prey. Once prey is found, electric eels will use a much stronger shock to stun prey, usually smaller fish.

Electric eels can grow up to 8 feet (2.5 meters) in length. Despite their appearance, electric eels aren't actually eels at all! They are more closely related to carp and catfish.

The shock of an electric eel has been known to knock a horse off its feet. Now that's shocking!

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The electric eel gets its name from its shocking abilities! Special organs in the eel's body release powerful electric charges of up to 650 volts-that's more than five times the power of a standard United States wall socket! Although eels have the power to be the bullies of the Amazon, they are actually not very aggressive animals.

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Beautiful “Underwater Kaleidoscope” | National Geographic

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Why you Should NEVER Release Pets into the Wild

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Abundant fossil bones, teeth, trackways, and other hard evidence have revealed that Earth was the domain of the dinosaurs for at least 230 million years. But so far, not a single trace of dinosaur remains has been found in rocks younger than about 66 million years. At that point, as the Cretaceous period yielded to the Paleogene, it seems that all nonavian dinosaurs suddenly ceased to exist.

Along with them went fearsome marine reptiles such as the mosasaurs, ichthyosaurs, and plesiosaurs, as well as all the flying reptiles known as pterosaurs. Ancient forests seem to have flamed out across much of the planet. And while some mammals, birds, small reptiles, fish, and amphibians survived, diversity among the remaining life-forms dropped precipitously. In total, this mass extinction event claimed three quarters of life on Earth.

Piecing together what happened has been a massive effort for paleontologists, and theories for what killed the dinosaurs and the rest of the planet's Cretaceous inhabitants have ranged from the plausible to the downright zany. For now, two leading ideas are battling it out within the scientific community: Were dinosaurs victims of interplanetary violence, or more Earthly woes?

Death from above

One of the most well-known theories for the death of the dinosaurs is the Alvarez hypothesis, named after the father-and-son duo Luis and Walter Alvarez. In 1980, these two scientists proposed the notion that a meteor the size of a mountain slammed into Earth 66 million years ago, filling the atmosphere with gas, dust, and debris that drastically altered the climate.

Their key piece of evidence is an oddly high amount of the metal iridium in what's known as the Cretaceous-Paleogene, or K-Pg, layer-the geologic boundary zone that seems to cap any known rock layers containing dinosaur fossils. Iridium is relatively rare in Earth's crust but is more abundant in stony meteorites, which led the Alvarezs to conclude that the mass extinction was caused by an extraterrestrial object. The theory gained even more steam when scientists were able to link the extinction event to a huge impact crater along the coast of Mexico's Yucatán Peninsula. At about 93 miles wide, the Chicxulub crater seems to be the right size and age to account for the dino die-off.

In 2016, scientists drilled a rock core inside the underwater part of Chicxulub, pulling up a sample stretching deep beneath the seabed. This rare peek inside the guts of the crater showed that the impact would have been powerful enough to send deadly amounts of vaporized rock and gases into the atmosphere, and that the effects would have persisted for years. And in 2019, paleontologists digging in North Dakota found a treasure trove of fossils extremely close to the K-Pg boundary, essentially capturing the remains of an entire ecosystem that existed shortly before the mass extinction. Tellingly, the fossil-bearing layers contain loads of tiny glass bits called tektites-likely blobs of melted rock kicked up by the impact that solidified in the atmosphere and then rained down over Earth.

Volcanic fury

However, other scientists maintain that the evidence for a massive meteor impact event is inconclusive, and that the more likely culprit may be Earth itself.

Ancient lava flows in India known as the Deccan Traps also seem to match nicely in time with the end of the Cretaceous, with massive outpourings of lava spewing forth between 60 and 65 million years ago. Today, the resulting volcanic rock covers nearly 200,000 square miles in layers that are in places more than 6,000 feet thick. Such a vast eruptive event would have choked the skies with carbon dioxide and other gases that would have dramatically changed Earth's climate.

Proponents of this theory point to multiple clues that suggest volcanism is a better fit. For one, some studies show that Earth's temperature was changing even before the proposed impact event. Other research has found evidence for mass die-offs much earlier than 66 million years ago, with some signs that dinosaurs in particular were already in a slow decline in the late Cretaceous. What's more, volcanic activity is frequent on this planet and is a plausible culprit for other ancient extinctions, while giant meteor strikes are much more rare. This all makes sense, supporters say, if ongoing volcanic eruptions were the root cause of the world-wide K-Pg extinctions.

Why not both?

Increasingly, scientists trying to unravel this prehistoric mystery are seeing room for a combination of these ideas. It's possible the dinosaurs were the unlucky recipients of a geologic one-two punch, with volcanism weakening ecosystems enough to make them vulnerable to an incoming meteor.

But that notion depends a lot on more precise dating of the Deccan Traps and the Chicxulub crater. In 2019, two independent studies looked at geochemical clues from Deccan Traps lava and came to slightly different conclusions, with one paper suggesting the volcanoes played a supporting role in the dinosaurs' demise by causing pre-impact declines, and the other saying the eruptions came after the impact event and may have played only a small role in ushering along their end.

This debate may rage for years, as scientists dig up new clues and develop new techniques for understanding the past. But whether space invaders or loads of lava are to blame, it's clear that scientists studying the dinosaurs' last gasp are revealing vital lessons about the effects of dramatic climate change on Earth's inhabitants.

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Learn about the mass extinction event 66 million years ago and the evidence for what ended the age of the dinosaurs.

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This Baby Elephant Is Learning How To Use Her Trunk | The Zoo: San Diego

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Built on a collection of 40,000+ popular, high-quality books from 250+ of the world's best publishers, Epic safely fuels curiosity and reading confidence for kids 12 and under.

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LIVE 4K Wild Animal & Bird Feeder Cam - Recke, Germany - Hedgehog & Bird Watching, 24/7, ASMR

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Feed My Pet Friday: Softshell, Painted, Map, and Musk Turtles!

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bird, (class Aves), any of the more than 10,400 living species unique in having feathers, the major characteristic that distinguishes them from all other animals. A more-elaborate definition would note that they are warm-blooded vertebrates more related to reptiles than to mammals and that they have a four-chambered heart (as do mammals), forelimbs modified into wings (a trait shared with bats), a hard-shelled egg, and keen vision, the major sense they rely on for information about the environment. Their sense of smell is not highly developed, and auditory range is limited. Most birds are diurnal in habit. More than 1,000 extinct species have been identified from fossil remains.

Since earliest times birds have been not only a material but also a cultural resource. Bird figures were created by prehistoric humans in the Lascaux Grotto of France and have featured prominently in the mythology and literature of societies throughout the world. Long before ornithology was practiced as a science, interest in birds and the knowledge of them found expression in conversation and stories, which then crystallized into the records of general culture. Ancient Egyptian hieroglyphs and paintings, for example, include bird figures. The Bible refers to Noah's use of the raven and dove to bring him information about the proverbial Flood.

Various bird attributes, real or imagined, have led to their symbolic use in language as in art. Aesop's fables abound in bird characters. The Physiologus and its descendants, the bestiaries of the Middle Ages, contain moralistic writings that use birds as symbols for conveying ideas but indicate little knowledge of the birds themselves. Supernatural beliefs about birds probably took hold as early as recognition of the fact that some birds were good to eat. Australian Aborigines, for example, drove the black-and-white flycatcher from camp, lest it overhear conversation and carry the tales to enemies. Peoples of the Pacific Islands saw frigate birds as symbols of the Sun and as carriers of omens and frequently portrayed them in their art. The raven-a common symbol of dark prophecy-was the most important creature to the Indians of the Pacific Northwest and was immortalized in Edgar Allan Poe's poem "The Raven." Eagles have long been symbols of power and prestige in many parts of the world, including Europe, where their representations are often seen in heraldry. Native Americans sprinkled eagle down before guests as a sign of peace and friendship, and eagle feathers were commonly used in rituals and headdresses. The resplendent quetzal-the national bird of Guatemala, which shares its name with the currency and is a popular motif in art, fabric, and jewelry-was worshipped and deified by the ancient Mayans and Aztecs. Highly symbolic birds include the phoenix, representing resurrection, and the owl, a common symbol of wisdom but also a reminder of death in Native American mythology. The bird in general has long been a common Christian symbol of the transcendent soul, and in medieval iconography a bird entangled in foliage symbolized the soul embroiled in the materialism of the secular world.

In modern times the recreational pleasures of bird-watching have grown in tandem with the rise of environmentalism. Evolving from the American and European "shoot-and-stuff" mania of the 19th century, bird-watching became a sportlike activity based on rapid identification-the rarest being the most rewarding-with the aid of binoculars and spotting scopes. The change from shooting to sighting coincided with campaigns, beginning about 1900, to halt the slaughter of wild birds for food and millinery. Bird-watching was advanced by the publication of excellent field guides and improvements in photography and sound recording. By mid-century the watcher's enjoyable but rather unsophisticated tallying of "year lists" and "life lists" of species personally observed was being augmented, if not replaced, by interest in careful studies of bird behaviour, migration, ecology, and the like. This trend was abetted by bird banding (called ringing in the United Kingdom) and by such organizations as the British Trust for Ornithology and the National Audubon Society, which coordinate professional and amateur observations and efforts with scientific studies.

General features

Birds arose as warm-blooded, arboreal, flying creatures with forelimbs adapted for flight and hind limbs for perching. This basic plan has become so modified during the course of evolution that in some forms it is difficult to recognize.

Among flying birds, the wandering albatross has the greatest wingspan, up to 3.5 metres (11.5 feet), and the trumpeter swan perhaps the greatest weight, 17 kg (37 pounds). In the largest flying birds, part of the bone is replaced by air cavities (pneumatic skeletons) because the maximum size attainable by flying birds is limited by the fact that wing area varies as the square of linear proportions, and weight or volume as the cube. During the Pleistocene Epoch (2.6 million to 11,700 years ago) lived a bird called Teratornis incredibilis. Though similar to the condors of today, it had a larger estimated wingspan of about 5 metres (16.5 feet) and was by far the largest known flying bird.

The smallest living bird is generally acknowledged to be the bee hummingbird of Cuba, which is 6.3 cm (2.5 inches) long and weighs less than 3 grams (about 0.1 ounce). The minimum size is probably governed by another aspect of the surface-volume ratio: the relative increase, with decreasing size, in surface through which heat can be lost. The small size of some hummingbirds may be facilitated by a decrease in heat loss resulting from their becoming torpid at night.

When birds lose the power of flight, the limit on their maximum size is increased, as can be seen in the ostrich and other ratites such as the emu, cassowary, and rhea. The ostrich is the largest living bird and may stand 2.75 metres (9 feet) tall and weigh 150 kg (330 pounds). Some recently extinct birds were even larger: the largest moas of New Zealand and the elephant birds of Madagascar may have reached over 3 metres (10 feet) in height.

The ability to fly has permitted an almost unlimited diversification of birds, so that they are now found virtually everywhere on Earth, from occasional stragglers over the polar ice caps to complex communities in tropical forests. In general the number of species found breeding in a given area is directly proportional to the size of the area and the diversity of habitats available. The total number of species is also related to such factors as the position of the area with respect to migration routes and to wintering grounds of species that nest outside the area. In the United States, Texas and California have the most-approximately 620 for each (the figure varies based on criteria used for inclusion on state lists, such as unconfirmed, accidental, hypothetical, extirpated, and extinct species). More than 920 species have been recorded from North America north of Mexico. The figure for Europe west of the Ural Mountains and including most of Turkey is 514. More than 700 species live in Russia. At least 4,400 species live in North and South America. Although several South American countries boast well over 1,000 species, Costa Rica, with an area of only about 51,000 square km (about 20,000 square miles) and a known avifauna of more than 800 species, probably has the most diversity for its size. Asia accounts for more than 25 percent of the world's species, with 2,700 species, and Africa slightly less, with about 2,300.

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bird | Description, Species, Classification, Types, & Facts

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bird, (class Aves), any of the more than 10,400 living species unique in having feathers, the major characteristic that distinguishes them from all other animals.

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Instantly access Galapagos Tortoises plus over 40,000 of the best books & videos for kids.

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Many amphibians-like this Ceratophrys cranwelli-exhibit biofluorescence.[3]

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

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

The earliest amphibians evolved in the Devonian period from sarcopterygian fish with lungs and bony-limbed fins, features that were helpful in adapting to dry land. They diversified and became ecologically dominant during the Carboniferous and Permian periods, but were later displaced in terrestrial environments by early reptiles and basal synapsids (mammal predecessors). The origin of modern amphibians belonging to Lissamphibia, which first appeared during the Early Triassic, around 250 million years ago, has long been contentious. However the emerging consensus is that they likely originated from temnospondyls, the most diverse group of prehistoric amphibians, during the Permian period.

The three modern orders of amphibians are Anura (the frogs), Urodela (the salamanders), and Apoda (the caecilians). A fourth group, the Albanerpetontidae, became extinct around 2 million years ago. The number of known amphibian species is approximately 8,000, of which nearly 90% are frogs. The smallest amphibian (and vertebrate) in the world is a frog from New Guinea (Paedophryne amauensis) with a length of just 7.7 mm (0.30 in). The largest living amphibian is the 1.8 m (5 ft 11 in) South China giant salamander ( Andrias sligoi), but this is dwarfed by prehistoric temnospondyls such as Mastodonsaurus which could reach up to 6 m (20 ft) in length. The study of amphibians is called batrachology, while the study of both reptiles and amphibians is called herpetology.

Classification

The world's smallest known vertebrate, Paedophryne amauensis, sitting on a U.S. dime. The dime is 17.9 mm in diameter, for scale

The word amphibian is derived from the Ancient Greek term ( amphíbios), which means 'both kinds of life', meaning 'of both kinds' and meaning 'life'. The term was initially used as a general adjective for animals that could live on land or in water, including seals and otters. Traditionally, the class Amphibia includes all tetrapod vertebrates that are not amniotes. Amphibia in its widest sense (sensu lato) was divided into three subclasses, two of which are extinct:

These three subclasses do not include all extinct amphibians. Other extinct amphibian groups include Embolomeri (Late Paleozoic large aquatic predators), Seymouriamorpha (semiaquatic to terrestrial Permian forms related to amniotes), among others. Names such as Tetrapoda and Stegocephalia encompass the entirety of amphibian-grade tetrapods, while Reptiliomorpha or Anthracosauria have been used to describe extinct amphibians more closely related to amniotes than to lissamphibians.

Subclass Lepospondyli† (A potentially polyphyletic Late Paleozoic group of small forms, likely more closely related to amniotes than Lissamphibia)

Subclass Temnospondyli† (diverse Late Paleozoic and early Mesozoic grade, some of which were large predators)

Subclass Lissamphibia (all modern amphibians, including frogs, toads, salamanders, newts and caecilians)

Salientia (frogs, toads and relatives): Early Triassic to present-7,360 current species in 53 families. Modern (crown group) salientians are described via the name Anura.

Caudata (salamanders, newts and relatives): Late Triassic to present-764 current species in 9 families. Modern (crown group) caudatans are described via the name Urodela.

Gymnophiona (caecilians and relatives): Late Triassic to present-215 current species in 10 families. The name Apoda is also sometimes used for caecilians.

Allocaudata† (Albanerpetontidae) Middle Jurassic - Early Pleistocene

These three subclasses do not include all extinct amphibians. Other extinct amphibian groups include Embolomeri (Late Paleozoic large aquatic predators), Seymouriamorpha (semiaquatic to terrestrial Permian forms related to amniotes), among others. Names such as Tetrapoda and Stegocephalia encompass the entirety of amphibian-grade tetrapods, while Reptiliomorpha or Anthracosauria are variably used to describe extinct amphibians more closely related to amniotes than to lissamphibians.

The actual number of species in each group depends on the taxonomic classification followed. The two most common systems are the classification adopted by 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". The numbers of species cited above follows Frost and the total number of known (living) amphibian species as of March 31, 2019, is exactly 8,000, of which nearly 90% are frogs.

With the phylogenetic classification, the taxon Labyrinthodontia has been discarded as it is a polyparaphyletic 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 uncertain, and Lissamphibia may possibly fall within extinct groups, like the Temnospondyli (traditionally placed in the subclass Labyrinthodontia) or the Lepospondyli, and in some analyses even in 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. If the common ancestor of amphibians and amniotes is included in Amphibia, it becomes a paraphyletic group.

All modern amphibians are included in the subclass Lissamphibia, which is usually considered a clade, a group of species that have evolved from a common ancestor. The three modern orders are Anura (the frogs), Caudata (or Urodela, the salamanders), and Gymnophiona (or Apoda, the caecilians). It has 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. Although the fossils of several older proto-frogs with primitive characteristics are known, the oldest "true frog", with hopping adaptations is Prosalirus bitis, from the Early Jurassic Kayenta Formation of Arizona. It is anatomically very similar to modern frogs. The oldest known caecilians are Funcusvermis gilmorei (from the Late Triassic) and Eocaecilia micropodia (from the Early Jurassic), both from Arizona. The earliest salamander is Beiyanerpeton jianpingensis from the Late Jurassic of northeastern China.

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

Evolutionary history

The first major groups of amphibians developed in the Devonian period, around 370 million years ago, from lobe-finned fish which were similar to the modern coelacanth and lungfish. These ancient lobe-finned fish had evolved multi-jointed leg-like fins with digits that enabled them to crawl along the sea bottom. Some fish had developed primitive lungs that help them breathe air when the stagnant pools of the Devonian swamps were low in oxygen. They could also use their strong fins to hoist themselves out of the water and onto dry land if circumstances so required. Eventually, their bony fins would evolve into limbs and they would become the ancestors to all tetrapods, including modern 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. They had started to develop lungs, but still breathed predominantly with gills.

Many examples of species showing transitional features have been discovered. Ichthyostega was one of the first primitive amphibians, with nostrils and more efficient lungs. It had four sturdy limbs, a neck, a tail with fins and a skull very similar to that of the lobe-finned fish, Eusthenopteron. Amphibians evolved adaptations that allowed them to stay out of the water for longer periods. Their lungs improved and their skeletons became heavier and stronger, better able to support the weight of their bodies on land. They developed "hands" and "feet" with five or more digits; the skin became more capable of retaining body fluids and resisting desiccation. The fish's hyomandibula bone in the hyoid region behind the gills diminished in size and became the stapes of the amphibian ear, an adaptation necessary for hearing on dry land. An affinity between the amphibians and the teleost fish is the multi-folded structure of the teeth and the paired supra-occipital bones at the back of the head, neither of these features being found elsewhere in the animal kingdom.

At the end of the Devonian period (360 million years ago), the seas, rivers and lakes were teeming with life while the land was the realm of early plants and devoid of vertebrates, though some, such as Ichthyostega, may have sometimes hauled themselves out of the water. It is thought they may have propelled themselves with their forelimbs, dragging their hindquarters in a similar manner to that used by the elephant seal. In the early Carboniferous (360 to 323 million years ago), the climate was relatively wet and warm. Extensive swamps developed with mosses, ferns, horsetails and calamites. Air-breathing arthropods evolved and invaded the land where they provided food for the carnivorous amphibians that began to adapt to the terrestrial environment. There were no other tetrapods on the land and the amphibians were at the top of the food chain, with some occupying ecological positions currently held by crocodiles. Though equipped with limbs and the ability to breathe air, most still had a long tapering body and strong tail. Others were the top land predators, sometimes reaching several metres in length, preying on the large insects of the period and the many types of fish in the water. They still needed to return to water to lay their shell-less eggs, and even most modern amphibians have a fully aquatic larval stage with gills like their fish ancestors. It was the development of the amniotic egg, which prevents the developing embryo from drying out, that enabled the reptiles to reproduce on land and which led to their dominance in the period that followed.

After the Carboniferous rainforest collapse amphibian dominance gave way to reptiles, and amphibians were further devastated by the Permian-Triassic extinction event. During the Triassic Period (252 to 201 million years ago), the reptiles continued to out-compete the amphibians, leading to a reduction in both the amphibians' size and their importance in the biosphere. According to the fossil record, Lissamphibia, which includes all modern amphibians and is the only surviving lineage, may have branched off from the extinct groups Temnospondyli and Lepospondyli at some period between the Late Carboniferous and the Early Triassic. The relative scarcity of fossil evidence precludes precise dating, but the most recent molecular study, based on multilocus sequence typing, suggests a Late Carboniferous/ Early Permian origin for extant amphibians.

The origins and evolutionary relationships between the three main groups of amphibians is a matter of debate. A 2005 molecular phylogeny, based on rDNA analysis, suggests that salamanders and caecilians are more closely related to each other than they are to frogs. It also appears that the divergence of the three groups took place in the Paleozoic or early Mesozoic (around 250 million years ago), before the breakup of the supercontinent Pangaea and soon after their divergence from the lobe-finned fish. The briefness of this period, and the swiftness with which radiation took place, would help account for the relative scarcity of primitive amphibian fossils. There are large gaps in the fossil record, the discovery of the dissorophoid temnospondyl Gerobatrachus from the Early Permian in Texas in 2008 provided a missing link with many of the characteristics of modern frogs. Molecular analysis suggests that the frog-salamander divergence took place considerably earlier than the palaeontological evidence indicates. One study suggested suggested that the last common ancestor of all modern amphibians lived about 315 million years ago, and that stereospondyl temnospondyls are the closest relatives to the caecilians. However, most studies support a single monophyletic origin of all modern amphibians within the dissorophoid temnospondyls.

As they evolved from lunged fish, amphibians had to make certain adaptations for living on land, including the need to develop new means of locomotion. In the water, the sideways thrusts of their tails had propelled them forward, but on land, quite different mechanisms were required. Their vertebral columns, limbs, limb girdles and musculature needed to be strong enough to raise them off the ground for locomotion and feeding. Terrestrial adults discarded their lateral line systems and adapted their sensory systems to receive stimuli via the medium of the air. They needed to develop new methods to regulate their body heat to cope with fluctuations in ambient temperature. They developed behaviours suitable for reproduction in a terrestrial environment. Their skins were exposed to harmful ultraviolet rays that had previously been absorbed by the water. The skin changed to become more protective and prevent excessive water loss.

Characteristics

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

Modern amphibians have a simplified anatomy compared to their ancestors due to paedomorphosis, caused by two evolutionary trends: miniaturization and an unusually large genome, which result in a slower growth and development rate compared to other vertebrates. Another reason for their size is associated with their rapid metamorphosis, which seems to have evolved only in the ancestors of lissamphibia; in all other known lines the development was much more gradual. Because a remodeling of the feeding apparatus means they do not eat during the metamorphosis, the metamorphosis has to go faster the smaller the individual is, so it happens at an early stage when the larvae are still small. (The largest species of salamanders do not go through a metamorphosis.) Amphibians that lay eggs on land often go through the whole metamorphosis inside the egg. An anamniotic terrestrial egg is less than 1 cm in diameter due to diffusion problems, a size which puts a limit on the amount of posthatching growth.

The smallest amphibian (and vertebrate) in the world is a microhylid frog from New Guinea (Paedophryne amauensis) first discovered in 2012. It has an average length of 7.7 mm (0.30 in) and is part of a genus that contains four of the world's ten smallest frog species. The largest living amphibian is the 1.8 m (5 ft 11 in) Chinese giant salamander ( Andrias davidianus) but this is a great deal smaller than the largest amphibian that ever existed-the extinct 9 m (30 ft) Prionosuchus, a crocodile-like temnospondyl dating to 270 million years ago from the middle Permian of Brazil. The largest frog is the African Goliath frog ( Conraua goliath), which can reach 32 cm (13 in) and weigh 3 kg (6.6 lb).

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

The order Anura (from the Ancient Greek meaning "without" and oura meaning "tail") comprises the frogs and toads. They usually have long hind limbs that fold underneath them, shorter forelimbs, webbed toes with no claws, no tails, large eyes and glandular moist skin. Members of this order with smooth skins are commonly referred to as frogs, while those with warty skins are known as toads. The difference is not a formal one taxonomically and there are numerous exceptions to this rule. Members of the family Bufonidae are known as the "true toads". Frogs range in size from the 30-centimetre (12 in) Goliath frog ( Conraua goliath) of West Africa to the 7.7-millimetre (0.30 in) Paedophryne amauensis, first described in Papua New Guinea in 2012, which is also the smallest known vertebrate. Although most species are associated with water and damp habitats, some are specialised to live in trees or in deserts. They are found worldwide except for polar areas.

Anura is divided into three suborders that are broadly accepted by the scientific community, but the relationships between some families remain unclear. Future molecular studies should provide further insights into their evolutionary relationships. The suborder Archaeobatrachia contains four families of primitive frogs. These are Ascaphidae, Bombinatoridae, Discoglossidae and Leiopelmatidae which have few derived features and are probably paraphyletic with regard to other frog lineages. The six families in the more evolutionarily advanced suborder Mesobatrachia are the fossorial Megophryidae, Pelobatidae, Pelodytidae, Scaphiopodidae and Rhinophrynidae and the obligatorily aquatic Pipidae. These have certain characteristics that are intermediate between the two other suborders. Neobatrachia is by far the largest suborder and includes the remaining families of modern frogs, including most common species. Ninety-six percent of the over 5,000 extant species of frog are neobatrachians.

The order Caudata (from the Latin cauda meaning "tail") consists of the salamanders-elongated, low-slung animals that mostly resemble lizards in form. This is a symplesiomorphic trait and they are no more closely related to lizards than they are to mammals. Salamanders lack claws, have scale-free skins, either smooth or covered with tubercles, and tails that are usually flattened from side to side and often finned. They range in size from the Chinese giant salamander ( Andrias davidianus), which has been reported to grow to a length of 1.8 metres (5 ft 11 in), to the diminutive Thorius pennatulus from Mexico which seldom exceeds 20 mm (0.8 in) in length. Salamanders have a mostly Laurasian distribution, being present in much of the Holarctic region of the northern hemisphere. The family Plethodontidae is also found in Central America and South America north of the Amazon basin; South America was apparently invaded from Central America by about the start of the Miocene, 23 million years ago. Urodela is a name sometimes used for all the extant species of salamanders. Members of several salamander families have become paedomorphic and either fail to complete their metamorphosis or retain some larval characteristics as adults. Most salamanders are under 15 cm (6 in) long. They may be terrestrial or aquatic and 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.

The suborder Cryptobranchoidea contains the primitive salamanders. A number of fossil cryptobranchids have been found, but there are only three living species, the Chinese giant salamander ( Andrias davidianus), the Japanese giant salamander ( Andrias japonicus) and the hellbender ( Cryptobranchus alleganiensis) from North America. These large amphibians retain several larval characteristics in their adult state; gills slits are present and the eyes are unlidded. A unique feature is their ability to feed by suction, depressing either the left side of their lower jaw or the right. The males excavate nests, persuade females to lay their egg strings inside them, and guard them. As well as breathing with lungs, they respire through the many folds in their thin skin, which has capillaries close to the surface.

The suborder Salamandroidea contains the advanced salamanders. They differ from the cryptobranchids by having fused prearticular bones in the lower jaw, and by using internal fertilisation. In salamandrids, the male deposits a bundle of sperm, the spermatophore, and the female picks it up and inserts it into her cloaca where the sperm is stored until the eggs are laid. The largest family in this group is Plethodontidae, the lungless salamanders, which includes 60% of all salamander species. The family Salamandridae includes the true salamanders and the name " newt" is given to members of its subfamily Pleurodelinae.

The third suborder, Sirenoidea, contains the four species of sirens, which are in a single family, Sirenidae. Members of this order are eel-like aquatic salamanders with much reduced forelimbs and no hind limbs. Some of their features are primitive while others are derived. Fertilisation is likely to be external as sirenids lack the cloacal glands used by male salamandrids to produce spermatophores and the females lack spermathecae for sperm storage. Despite this, the eggs are laid singly, a behaviour not conducive for external fertilisation.

Gymnophiona

The order Gymnophiona (from the Greek gymnos meaning "naked" and ophis meaning "serpent") or Apoda comprises the caecilians. These are long, cylindrical, limbless animals with a snake- or worm-like form. The adults vary in length from 8 to 75 centimetres (3 to 30 inches) with the exception of Thomson's caecilian ( Caecilia thompsoni), which can reach 150 centimetres (4.9 feet). A caecilian's skin has a large number of transverse folds and in some species contains tiny embedded dermal scales. It has rudimentary eyes covered in skin, which are probably limited to discerning differences in light intensity. It also has a pair of short tentacles near the eye that can be extended and which have tactile and olfactory functions. Most caecilians live underground in burrows in damp soil, in rotten wood and under plant debris, but some are aquatic. Most species lay their eggs underground and when the larvae hatch, they make their way to adjacent bodies of water. Others brood their eggs and the larvae undergo metamorphosis before the eggs hatch. A few species give birth to live young, nourishing them with glandular secretions while they are in the oviduct. Caecilians have a mostly Gondwanan distribution, being found in tropical regions of Africa, Asia and Central and South America.

Anatomy and physiology

The bright colours of the common reed frog (Hyperolius viridiflavus) are typical of a toxic species

The integumentary structure contains some typical characteristics common to terrestrial vertebrates, such as the presence of highly cornified outer layers, renewed periodically through a moulting 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 mostly in one piece, in contrast to mammals and birds where it is shed in flakes. Amphibians often eat the sloughed skin. Caecilians are unique among amphibians in having mineralized dermal scales embedded in the dermis between the furrows in the skin. The similarity of these to the scales of bony fish is largely superficial. Lizards and some frogs have somewhat similar osteoderms forming bony deposits in the dermis, but this is an example of convergent evolution with similar structures having arisen independently in diverse vertebrate lineages.

Cross section of frog skin:

Amphibian skin is permeable to water. Gas exchange can take place through the skin ( cutaneous respiration) and this allows adult amphibians to respire without rising to the surface of water and to hibernate at the bottom of ponds. To compensate for their thin and delicate skin, amphibians have evolved mucous glands, principally on their heads, backs and tails. The secretions produced by these help keep the skin moist. In addition, most species of amphibian have granular glands that secrete distasteful or poisonous substances. Some amphibian toxins can be lethal to humans while others have little effect. The main poison-producing glands, the parotoids, produce the neurotoxin bufotoxin and are located behind the ears of toads, along the backs of frogs, behind the eyes of salamanders and on the upper surface of caecilians.

The skin colour 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 colour) and the lipophores (yellow, the most superficial layer). The colour change displayed by many species is initiated by hormones secreted by the pituitary gland. Unlike bony fish, there is no direct control of the pigment cells by the nervous system, and this results in the colour change taking place more slowly than happens in fish. A vividly coloured skin usually indicates that the species is toxic and is a warning sign to predators.

Skeletal system and locomotion

Amphibians have a skeletal system that is structurally homologous to other tetrapods, though with a number of variations. They all have four limbs except for the legless caecilians and a few species of salamander with reduced or no limbs. The bones are hollow and lightweight. The musculoskeletal system is strong to enable it to support the head and body. The bones are fully ossified and the vertebrae interlock with each other by means of overlapping processes. The pectoral girdle is supported by muscle, and the well-developed pelvic girdle is attached to the backbone by a pair of sacral ribs. The ilium slopes forward and the body is held closer to the ground than is the case in mammals.

In most amphibians, there are four digits on the fore foot and five on the hind foot, but no claws on either. Some salamanders have fewer digits and the amphiumas are eel-like in appearance with tiny, stubby legs. The sirens are aquatic salamanders with stumpy forelimbs and no hind limbs. The caecilians are limbless. They burrow in the manner of earthworms with zones of muscle contractions moving along the body. On the surface of the ground or in water they move by undulating their body from side to side.

In frogs, the hind legs are larger than the fore legs, especially so in those species that principally move by jumping or swimming. In the walkers and runners the hind limbs are not so large, and the burrowers mostly have short limbs and broad bodies. The feet have adaptations for the way of life, with webbing between the toes for swimming, broad adhesive toe pads for climbing, and keratinised tubercles on the hind feet for digging (frogs usually dig backwards into the soil). In most salamanders, the limbs are short and more or less the same length and project at right angles from the body. Locomotion on land is by walking and the tail often swings from side to side or is used as a prop, particularly when climbing. In their normal gait, only one leg is advanced at a time in the manner adopted by their ancestors, the lobe-finned fish. Some salamanders in the genus and certain plethodontids climb trees and have long limbs, large toepads and prehensile tails. In aquatic salamanders and in frog tadpoles, the tail has dorsal and ventral fins and is moved from side to side as a means of propulsion. Adult frogs do not have tails and caecilians have only very short ones.

Didactic model of an amphibian heart.

Salamanders use their tails in defence and some are prepared to jettison them to save their lives in a process known as autotomy. Certain species in the Plethodontidae have a weak zone at the base of the tail and use this strategy readily. The tail often continues to twitch after separation which may distract the attacker and allow the salamander to escape. Both tails and limbs can be regenerated. Adult frogs are unable to regrow limbs but tadpoles can do so.

Circulatory system

Juvenile amphibian circulatory systems are single loop systems which resemble fish.

1. Internal gills where the blood is reoxygenated

2. Point where the blood is depleted of oxygen and returns to the heart via veins

3. Two chambered heart

Red indicates oxygenated blood, and blue represents oxygen depleted blood.

Amphibians have a juvenile stage and an adult stage, and the circulatory systems of the two are distinct. In the juvenile (or tadpole) stage, the circulation is similar to that of a fish; the two-chambered heart pumps the blood through the gills where it is oxygenated, and is spread around the body and back to the heart in a single loop. In the adult stage, amphibians (especially frogs) lose their gills and develop lungs. They have a heart that consists of a single ventricle and two atria. When the ventricle starts contracting, deoxygenated blood is pumped through the pulmonary artery to the lungs. Continued contraction then pumps oxygenated blood around the rest of the body. Mixing of the two bloodstreams is minimized by the anatomy of the chambers.

Nervous and sensory systems

The nervous system is basically the same as in other vertebrates, with a central brain, a spinal cord, and nerves throughout the body. The amphibian brain is relatively simple but broadly the same structurally as in reptiles, birds and mammals. Their brains are elongated, except in caecilians, and contain the usual motor and sensory areas of tetrapods. The pineal body, known to regulate sleep patterns in humans, is thought to produce the hormones involved in hibernation and aestivation in amphibians.

Tadpoles retain the lateral line system of their ancestral fishes, but this is lost in terrestrial adult amphibians. Some caecilians possess electroreceptors that allow them to locate objects around them when submerged in water. The ears are well developed in frogs. There is no external ear, but the large circular eardrum lies on the surface of the head just behind the eye. This vibrates and sound is transmitted through a single bone, the stapes, to the inner ear. Only high-frequency sounds like mating calls are heard in this way, but low-frequency noises can be detected through another mechanism. There is a patch of specialized haircells, called papilla amphibiorum, in the inner ear capable of detecting deeper sounds. Another feature, unique to frogs and salamanders, is the columella-operculum 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 than those of frogs as they do not normally communicate with each other through the medium of sound.

The eyes of tadpoles lack lids, but at metamorphosis, the cornea becomes more dome-shaped, the lens becomes flatter, and eyelids and associated glands and ducts develop. The adult eyes 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 wavelengths.

Digestive and excretory systems

Dissected frog:

1. Right atrium

2. Liver

3. Aorta

4. Egg mass

5. Colon

6. Left atrium

7. Ventricle

8. Stomach

9. Left lung

10. Spleen

11. Small intestine

12. Cloaca

Many amphibians catch their prey by flicking out an elongated tongue with a sticky tip and drawing it back into the mouth before seizing the item with their jaws. Some use inertial feeding to help them swallow the prey, repeatedly thrusting their head forward sharply causing the food to move backwards in their mouth by inertia. Most amphibians swallow their prey whole without much chewing so they possess voluminous stomachs. The short oesophagus is lined with cilia that help to move the food to the stomach and mucus produced by glands in the mouth and pharynx eases its passage. The enzyme chitinase produced in the stomach helps digest the chitinous cuticle of arthropod prey.

Amphibians possess a pancreas, liver and gall bladder. The liver is usually large with two lobes. Its size is determined by its function as a glycogen and fat storage unit, and may change with the seasons as these reserves are built or used up. Adipose tissue is another important means of storing energy and this occurs in the abdomen (in internal structures called fat bodies), under the skin and, in some salamanders, in the tail.

There are two kidneys located dorsally, near the roof of the body cavity. Their job is to filter the blood of metabolic waste and transport the urine via ureters to the urinary bladder where it is stored before being passed out periodically through the cloacal vent. Larvae and most aquatic adult amphibians excrete the nitrogen as ammonia in large quantities of dilute urine, while terrestrial species, with a greater need to conserve water, excrete the less toxic product urea. Some tree frogs with limited access to water excrete most of their metabolic waste as uric acid.

Urinary bladder

Respiratory system

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 vascularised skin must remain moist to allow the oxygen to diffuse at a sufficiently high rate. 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.

Reproduction

Male orange-thighed frog (Litoria xanthomera) grasping the female during amplexus

For the purpose of reproduction most amphibians require fresh water although some lay their eggs on land and have developed various means of keeping them moist. A few (e.g. Fejervarya raja) can inhabit brackish water, but there are no true marine amphibians. There are reports, however, of particular amphibian populations unexpectedly invading marine waters. Such was the case with the Black Sea invasion of the natural hybrid Pelophylax esculentus reported in 2010.

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

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

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

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

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

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

Sexual selection has been studied in the red back salamander

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

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

Frogspawn, a mass of eggs surrounded by jelly

Amphibian egg:

1. Jelly capsule

2. Vitelline membrane

3. Perivitelline fluid

4. Yolk plug

5. Embryo

In the egg, the embryo is suspended in perivitelline fluid and surrounded by semi-permeable gelatinous capsules, with the yolk mass providing nutrients. As the larvae hatch, the capsules are dissolved by enzymes secreted from gland at the tip of the snout. The eggs of some salamanders and frogs contain unicellular green algae. These penetrate the jelly envelope after the eggs are laid and may increase the supply of oxygen to the embryo through photosynthesis. They seem to both speed up the development of the larvae and reduce mortality. In the wood frog ( Rana sylvatica), the interior of the globular egg cluster has been found to be up to 6 °C (11 °F) warmer than its surroundings, which is an advantage in its cool northern habitat.

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

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

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

Iodine and T4 (over stimulate the spectacular apoptosis [programmed cell death] of the cells of the larval gills, tail and fins) also stimulate the evolution of nervous systems transforming the aquatic, vegetarian tadpole into the terrestrial, carnivorous frog with better neurological, visuospatial, olfactory and cognitive abilities for hunting.

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

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

Salamanders

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

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

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

Caecilians

The caecilian Ichthyophis glutinosus with eggs and developing embryo

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

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

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

Parental care

The care of offspring among amphibians has been little studied but, in general, the larger the number of eggs in a batch, the less likely it is that any degree of parental care takes place. Nevertheless, it is estimated that in up to 20% of amphibian species, one or both adults play some role in the care of the young. Those species that breed in smaller water bodies or other specialised habitats tend to have complex patterns of behaviour in the care of their young.

Many woodland salamanders lay clutches of eggs under dead logs or stones on land. The black mountain salamander ( Desmognathus welteri) does this, the mother brooding the eggs and guarding them from predation as the embryos feed on the yolks of their eggs. When fully developed, they break their way out of the egg capsules and disperse as juvenile salamanders. The male hellbender, a primitive salamander, excavates an underwater nest and encourages females to lay there. The male then guards the site for the two or three months before the eggs hatch, using body undulations to fan the eggs and increase their supply of oxygen.

The male Colostethus subpunctatus, a tiny frog, protects the egg cluster which is hidden under a stone or log. When the eggs hatch, the male transports the tadpoles on his back, stuck there by a mucous secretion, to a temporary pool where he dips himself into the water and the tadpoles drop off. The male midwife toad ( Alytes obstetricans) winds egg strings round his thighs and carries the eggs around for up to eight weeks. He keeps them moist and when they are ready to hatch, he visits a pond or ditch and releases the tadpoles. The female gastric-brooding frog ( Rheobatrachus spp.) reared larvae in her stomach after swallowing either the eggs or hatchlings; however, this stage was never observed before the species became extinct. The tadpoles secrete a hormone that inhibits digestion in the mother whilst they develop by consuming their very large yolk supply. The pouched frog ( Assa darlingtoni) lays eggs on the ground. When they hatch, the male carries the tadpoles around in brood pouches on his hind legs. The aquatic Surinam toad ( Pipa pipa) raises its young in pores on its back where they remain until metamorphosis. The granular poison frog ( Oophaga granulifera) is typical of a number of tree frogs in the poison dart frog family Dendrobatidae. Its eggs are laid on the forest floor and when they hatch, the tadpoles are carried one by one on the back of an adult to a suitable water-filled crevice such as the axil of a leaf or the rosette of a bromeliad. The female visits the nursery sites regularly and deposits unfertilised eggs in the water and these are consumed by the tadpoles.

Genetics and genomics

Amphibians are notable among vertebrates for their diversity of chromosomes and genomes. The karyotypes ( chromosomes) have been determined for at least 1,193 (14.5%) of the ≈8,200 known ( diploid) species, including 963 anurans, 209 salamanders, and 21 caecilians. Generally, the karyotypes of diploid amphibians are characterized by 20-26 bi-armed chromosomes. Amphibians have also very large genomes compared to other taxa of vertebrates and corresponding variation in genome size ( C-value: picograms of DNA in haploid nuclei). The genome sizes range from 0.95 to 11.5 pg in frogs, from 13.89 to 120.56 pg in salamanders, and from 2.94 to 11.78 pg in caecilians.

The large genome sizes have prevented whole- genome sequencing of amphibians although a number of genomes have been published recently. The 1.7GB draft genome of Xenopus tropicalis was the first to be reported for amphibians in 2010. Compared to some salamanders this frog genome is tiny. For instance, the genome of the Mexican axolotl turned out to be 32 Gb, which is more than 10 times larger than the human genome (3GB).

Feeding and diet

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

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

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

Edible frog (Pelophylax esculentus) exhibiting cannibalism

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

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

Audio showing Brazilian torrent frog males executing advertisement, peep, and squeal calls.

Vocalization

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

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

Territorial behaviour

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

In salamanders, defence of a territory involves adopting an aggressive posture and if necessary attacking the intruder. This may involve snapping, chasing and sometimes biting, occasionally causing the loss of a tail. The behaviour of red back salamanders ( Plethodon cinereus) has been much studied. 91% of marked individuals that were later recaptured were within a metre (yard) of their original daytime retreat under a log or rock. A similar proportion, when moved experimentally a distance of 30 metres (98 ft), found their way back to their home base. The salamanders left odour marks around their territories which averaged 0.16 to 0.33 square metres (1.7 to 3.6 sq ft) in size and were sometimes inhabited by a male and female pair. These deterred the intrusion of others and delineated the boundaries between neighbouring areas. Much of their behaviour seemed stereotyped and did not involve any actual contact between individuals. An aggressive posture involved raising the body off the ground and glaring at the opponent who often turned away submissively. If the intruder persisted, a biting lunge was usually launched at either the tail region or the naso-labial grooves. Damage to either of these areas can reduce the fitness of the rival, either because of the need to regenerate tissue or because it impairs its ability to detect food.

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

Defence mechanisms

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

Perhaps the most poisonous animal in the world, the golden poison frog (Phyllobates terribilis) is endemic to Colombia.[154]

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

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

In amphibians, there is evidence of habituation, associative learning through both classical and instrumental learning, and discrimination abilities. Amphibians are widely considered to be sentient, able to feel emotions such as anxiety and fear.

In one experiment, when offered live fruit flies (Drosophila virilis), salamanders chose the larger of 1 vs 2 and 2 vs 3. Frogs can distinguish between low numbers (1 vs 2, 2 vs 3, but not 3 vs 4) and large numbers (3 vs 6, 4 vs 8, but not 4 vs 6) of prey. This is irrespective of other characteristics, i.e. surface area, volume, weight and movement, although discrimination among large numbers may be based on surface area.

Conservation

Dramatic declines in amphibian populations, including population crashes and mass localized extinction, have been noted since the late 1980s from locations all over the world, and amphibian declines are thus perceived to be one of the most critical threats to global biodiversity. In 2004, the International Union for Conservation of Nature (IUCN) reported stating that currently birds, mammals, and amphibians extinction rates were at minimum 48 times greater than natural extinction rates-possibly 1,024 times higher. In 2006, there were believed to be 4,035 species of amphibians that depended on water at some stage during their life cycle. Of these, 1,356 (33.6%) were considered to be threatened and this figure is likely to be an underestimate because it excludes 1,427 species for which there was insufficient data to assess their status. A number of causes are believed to be involved, including habitat destruction and modification, over-exploitation, pollution, introduced species, global warming, 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.

Food webs and predation

Any decline in amphibian numbers will affect the patterns of predation. The loss of carnivorous species near the top of the food chain will upset the delicate ecosystem balance and may cause dramatic increases in opportunistic species.

Predators that feed on amphibians are affected by their decline. The western terrestrial garter snake ( Thamnophis elegans) in California is largely aquatic and depends heavily on two species of frog that are decreasing in numbers, the Yosemite toad ( Bufo canorus) and the mountain yellow-legged frog ( Rana muscosa), putting the snake's future at risk. If the snake were to become scarce, this would affect birds of prey and other predators that feed on it. Meanwhile, in the ponds and lakes, fewer frogs means fewer tadpoles. These normally play an important role in controlling the growth of algae and also forage on detritus that accumulates as sediment on the bottom. A reduction in the number of tadpoles may lead to an overgrowth of algae, resulting in depletion of oxygen in the water when the algae later die and decompose. Aquatic invertebrates and fish might then die and there would be unpredictable ecological consequences.

Pollution and pesticides

The decline in amphibian and reptile populations has led to an awareness of the effects of pesticides on reptiles and amphibians. In the past, the argument that amphibians or reptiles were more susceptible to any chemical contamination than any land aquatic vertebrate was not supported by research until recently. Amphibians and reptiles have complex life cycles, live in different climate and ecological zones, and are more vulnerable to chemical exposure. Certain pesticides, such as organophosphates, neonicotinoids, and carbamates, react via cholinesterase inhibition. Cholinesterase is an enzyme that causes the hydrolysis of acetylcholine, an excitatory neurotransmitter that is abundant in the nervous system. AChE inhibitors are either reversible or irreversible, and carbamates are safer than organophosphorus insecticides, which are more likely to cause cholinergic poisoning. Reptile exposure to an AChE inhibitory pesticide may result in disruption of neural function in reptiles. The buildup of these inhibitory effects on motor performance, such as food consumption and other activities.

Conservation and protection strategies

The Amphibian Specialist Group of the IUCN is spearheading efforts to implement a comprehensive global strategy for amphibian conservation. 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. 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.

Another measure would be to stop exploitation of frogs for human consumption. In the Middle East, a growing appetite for eating frog legs and the consequent gathering of them for food was already linked to an increase in mosquitoes and thus has direct consequences for human health.

References

Cited texts

Further reading

Carroll, Robert L. (1988). Vertebrate Paleontology and Evolution. W. H. Freeman. ISBN 978-0-7167-1822-2.

Carroll, Robert L. (2009). The Rise of Amphibians: 365 Million Years of Evolution. 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.; Grant, Taran; Faivovich, Julián; Bain, Raoul H.; Haas, Alexander; Haddad, Célio F.B.; De Sá, Rafael O.; Channing, Alan; Wilkinson, Mark; Donnellan, Stephen C.; Raxworthy, Christopher J.; Campbell, Jonathan A.; Blotto, Boris L.; Moler, Paul; Drewes, Robert C.; Nussbaum, Ronald A.; Lynch, John D.; Green, David M.; Wheeler, Ward C. (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. S2CID 86140137.

Pounds, J. Alan; Bustamante, Martín R.; Coloma, Luis A.; Consuegra, Jamie A.; Fogden, Michael P. L.; Foster, Pru N.; La Marca, Enrique; Masters, Karen L.; Merino-Viteri, Andrés; Puschendorf, Robert; Ron, Santiago R.; Sánchez-Azofeifa, G. Arturo; Still, Christopher J.; Young, Bruce E. (2006). "Widespread amphibian extinctions from epidemic disease driven by global warming". Nature. 439 (7073): 161-167. Bibcode:2006Natur.439..161A. doi:10.1038/nature04246. PMID 16407945. S2CID 4430672.

Stuart, Simon N.; Chanson, Janice S.; Cox, Neil A.; Young, Bruce E.; Rodrigues, Ana S. L.; Fischman, Debra L.; Waller, Robert W. (2004). "Status and trends of amphibian declines and extinctions worldwide". Science. 306 (5702): 1783-1786. Bibcode:2004Sci...306.1783S. CiteSeerX 10.1.1.225.9620. doi:10.1126/science.1103538. PMID 15486254. S2CID 86238651.

Stuart, S. N.; Hoffmann, M.; Chanson, J. S.; Cox, N. A.; Berridge, R. J.; Ramani, P.; Young, B. E., eds. (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. Archived from the original on October 30, 2014.

External links

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Amphibians are ectothermic, anamniotic, four-limbed vertebrate animals that constitute the class Amphibia. They are a paraphyletic group encompassing all tetrapods excluding the amniotes (tetrapods with an amniotic membrane, such as modern reptiles, birds, and mammals). All extant (living) amphibians belong to the subclass Lissamphibia, with three living orders: Anura ( frogs), Urodela ( salamanders), and Gymnophiona ( caecilians).

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Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus . Other rat genera include (pack rats), (bandicoot rats) and (kangaroo rats).

Rats are typically distinguished from mice by their size. Usually the common name of a large muroid rodent will include the word "rat", while a smaller muroid's name will include "mouse". The common terms rat and mouse are not taxonomically specific. There are 56 known species of rats in the world.

Species and description

A rat in a suburb of Vancouver

Skeleton of a black rat (Rattus rattus) on display at the Museum of Osteology.

The best-known rat species are the black rat ( Rattus rattus) and the brown rat ( Rattus norvegicus). This group, generally known as the Old World rats or true rats, originated in Asia. Rats are bigger than most Old World mice, which are their relatives, but seldom weigh over 500 grams ( oz) in the wild.

The term rat is also used in the names of other small mammals that are not true rats. Examples include the North American pack rats (aka wood rats) and a number of species loosely called kangaroo rats. Rats such as the bandicoot rat ( Bandicota bengalensis) are murine rodents related to true rats but are not members of the genus Rattus.

Male rats are called bucks; unmated females, does, pregnant or parent females, dams; and infants, kittens or pups. A group of rats is referred to as a mischief.

The common species are opportunistic survivors and often live with and near humans; therefore, they are known as commensals. They may cause substantial food losses, especially in developing countries. However, the widely distributed and problematic commensal species of rats are a minority in this diverse genus. Many species of rats are island endemics, some of which have become endangered due to habitat loss or competition with the brown, black, or Polynesian rat.

Wild rodents, including rats, can carry many different zoonotic pathogens, such as Leptospira, Toxoplasma gondii, and Campylobacter. The Black Death is traditionally believed to have been caused by the microorganism Yersinia pestis, carried by the tropical rat flea (Xenopsylla cheopis), which preyed on black rats living in European cities during the epidemic outbreaks of the Middle Ages; these rats were used as transport hosts. Another zoonotic disease linked to the rat is foot-and-mouth disease.

Rats become sexually mature at age 6 weeks, but reach social maturity at about 5 to 6 months of age. The average lifespan of rats varies by species, but many only live about a year due to predation.

The black and brown rats diverged from other Old World rats in the forests of Asia during the beginning of the Pleistocene.

A closeup of a rat tail

The characteristic long tail of most rodents is a feature that has been extensively studied in various rat species models, which suggest three primary functions of this structure: thermoregulation, minor proprioception, and a nocifensive-mediated degloving response. Rodent tails-particularly in rat models-have been implicated with a thermoregulation function that follows from its anatomical construction. This particular tail morphology is evident across the family Muridae, in contrast to the bushier tails of Sciuridae, the squirrel family. The tail is hairless and thin skinned but highly vascularized, thus allowing for efficient countercurrent heat exchange with the environment. The high muscular and connective tissue densities of the tail, along with ample muscle attachment sites along its plentiful caudal vertebrae, facilitate specific proprioceptive senses to help orient the rodent in a three-dimensional environment. Murids have evolved a unique defense mechanism termed degloving that allows for escape from predation through the loss of the outermost integumentary layer on the tail. However, this mechanism is associated with multiple pathologies that have been the subject of investigation.[citation needed ]

Microscopic cross section of Rattus rattus tail, delineating tendon bundles, vasculature, and vertebral canal.

Multiple studies have explored the thermoregulatory capacity of rodent tails by subjecting test organisms to varying levels of physical activity and quantifying heat conduction via the animals' tails. One study demonstrated a significant disparity in heat dissipation from a rat's tail relative to its abdomen. This observation was attributed to the higher proportion of vascularity in the tail, as well as its higher surface-area-to-volume ratio, which directly relates to heat's ability to dissipate via the skin. These findings were confirmed in a separate study analyzing the relationships of heat storage and mechanical efficiency in rodents that exercise in warm environments. In this study, the tail was a focal point in measuring heat accumulation and modulation.[citation needed ]

On the other hand, the tail's ability to function as a proprioceptive sensor and modulator has also been investigated. As aforementioned, the tail demonstrates a high degree of muscularization and subsequent innervation that ostensibly collaborate in orienting the organism. Specifically, this is accomplished by coordinated flexion and extension of tail muscles to produce slight shifts in the organism's center of mass, orientation, etc., which ultimately assists it with achieving a state of proprioceptive balance in its environment. Further mechanobiological investigations of the constituent tendons in the tail of the rat have identified multiple factors that influence how the organism navigates its environment with this structure. A particular example is that of a study in which the morphology of these tendons is explicated in detail. Namely, cell viability tests of tendons of the rat's tail demonstrate a higher proportion of living fibroblasts that produce the collagen for these fibers. As in humans, these tendons contain a high density of golgi tendon organs that help the animal assess stretching of muscle in situ and adjust accordingly by relaying the information to higher cortical areas associated with balance, proprioception, and movement.[citation needed ]

The characteristic tail of murids also displays a unique defense mechanism known as in which the outer layer of the integument can be detached in order to facilitate the animal's escape from a predator. This evolutionary selective pressure has persisted despite a multitude of pathologies that can manifest upon shedding part of the tail and exposing more interior elements to the environment. Paramount among these are bacterial and viral infection, as the high density of vascular tissue within the tail becomes exposed upon avulsion or similar injury to the structure. The degloving response is a nocifensive response, meaning that it occurs when the animal is subjected to acute pain, such as when a predator snatches the organism by the tail.[citation needed ]

A domesticated rat

Specially bred rats have been kept as pets at least since the late 19th century. Pet rats are typically variants of the species brown rat, but black rats and giant pouched rats are also sometimes kept. Pet rats behave differently from their wild counterparts depending on how many generations they have been kept as pets. Pet rats do not pose any more of a risk of zoonotic diseases than pets such as cats or dogs. Tamed rats are generally friendly and can be taught to perform selected behaviors.

Selective breeding has brought about different color and marking varieties in rats. Genetic mutations have also created different fur types, such as rex and hairless. Congenital malformation in selective breeding have created the dumbo rat, a popular pet choice due to their low, saucer-shaped ears. A breeding standard exists for rat fanciers wishing to breed and show their rat at a rat show.

As subjects for scientific research

A laboratory rat strain, known as a Zucker rat, bred to be genetically prone to diabetes, a metabolic disorder also found among humans.

In 1895, Clark University in Worcester, Massachusetts, established a population of domestic albino brown rats to study the effects of diet and for other physiological studies. Over the years, rats have been used in many experimental studies, adding to our understanding of genetics, diseases, the effects of drugs, and other topics that have provided a great benefit for the health and wellbeing of humankind.

The aortic arches of the rat are among the most commonly studied in murine models due to marked anatomical homology to the human cardiovascular system. Both rat and human aortic arches exhibit subsequent branching of the brachiocephalic trunk, left common carotid artery, and left subclavian artery, as well as geometrically similar, nonplanar curvature in the aortic branches. Aortic arches studied in rats exhibit abnormalities similar to those of humans, including altered pulmonary arteries and double or absent aortic arches. Despite existing anatomical analogy in the inthrathoracic position of the heart itself, the murine model of the heart and its structures remains a valuable tool for studies of human cardiovascular conditions.

The rat's larynx has been used in experimentations that involve inhalation toxicity, allograft rejection, and irradiation responses. One experiment described four features of the rat's larynx. The first being the location and attachments of the thyroarytenoid muscle, the alar cricoarytenoid muscle, and the superior cricoarytenoid muscle, the other of the newly named muscle that ran from the arytenoid to a midline tubercle on the cricoid. The newly named muscles were not seen in the human larynx. In addition, the location and configuration of the laryngeal alar cartilage was described. The second feature was that the way the newly named muscles appear to be familiar to those in the human larynx. The third feature was that a clear understanding of how MEPs are distributed in each of the laryngeal muscles was helpful in understanding the effects of botulinum toxin injection. The MEPs in the posterior cricoarytenoid muscle, lateral cricoarytenoid muscle, cricothyroid muscle, and superior cricoarytenoid muscle were focused mostly at the midbelly. In addition, the medial thyroarytenoid muscle were focused at the midbelly while the lateral thyroarytenoid muscle MEPs were focused at the anterior third of the belly. The fourth and final feature that was cleared up was how the MEPs were distributed in the thyroarytenoid muscle.

Laboratory rats have also proved valuable in psychological studies of learning and other mental processes (Barnett 2002), as well as to understand group behavior and overcrowding (with the work of John B. Calhoun on behavioral sink). A 2007 study found rats to possess metacognition, a mental ability previously only documented in humans and some primates.

Domestic rats differ from wild rats in many ways. They are calmer and less likely to bite; they can tolerate greater crowding; they breed earlier and produce more offspring; and their brains, livers, kidneys, adrenal glands, and hearts are smaller (Barnett 2002).

Brown rats are often used as model organisms for scientific research. Since the publication of the rat genome sequence, and other advances, such as the creation of a rat SNP chip, and the production of knockout rats, the laboratory rat has become a useful genetic tool, although not as popular as mice. When it comes to conducting tests related to intelligence, learning, and drug abuse, rats are a popular choice due to their high intelligence, ingenuity, aggressiveness, and adaptability. Their psychology seems in many ways similar to that of humans.

Entirely new breeds or "lines" of brown rats, such as the Wistar rat, have been bred for use in laboratories. Much of the genome of Rattus norvegicus has been sequenced.

General intelligence

Early studies found evidence both for and against measurable intelligence using the "g factor" in rats. Part of the difficulty of understanding animal cognition, generally, is determining what to measure. One aspect of intelligence is the ability to learn, which can be measured using a maze like the T-maze. Experiments done in the 1920s showed that some rats performed better than others in maze tests, and if these rats were selectively bred, their offspring also performed better, suggesting that in rats an ability to learn was heritable in some way.

Rat meat is a food that, while taboo in some cultures, is a dietary staple in others.

Working rats

Rats have been used as working animals. Tasks for working rats include the sniffing of gunpowder residue, demining, acting and animal-assisted therapy. Rats have a keen sense of smell and are easy to train. These characteristics have been employed, for example, by the Belgian non-governmental organization APOPO, which trains rats (specifically African giant pouched rats) to detect landmines and diagnose tuberculosis through smell.

Rodent Bait Station, Chennai, India

Rats have long been considered deadly pests. Once considered a modern myth, the rat flood in India occurs every fifty years, as armies of bamboo rats descend upon rural areas and devour everything in their path. Rats have long been held up as the chief villain in the spread of the Bubonic Plague; however, recent studies show that rats alone could not account for the rapid spread of the disease through Europe in the Middle Ages. Still, the Centers for Disease Control does list nearly a dozen diseases directly linked to rats.

Most urban areas battle rat infestations. A 2015 study by the American Housing Survey (AHS) found that eighteen percent of homes in Philadelphia showed evidence of rodents. Boston, New York City, and Washington, D.C., also demonstrated significant rodent infestations. Indeed, rats in New York City are famous for their size and prevalence. The urban legend that the rat population in Manhattan equals that of its human population was definitively refuted by Robert Sullivan in his book Rats but illustrates New Yorkers' awareness of the presence, and on occasion boldness and cleverness, of the rodents. New York has specific regulations for eradicating rats; multifamily residences and commercial businesses must use a specially trained and licensed rat catcher.

Chicago was declared the "rattiest city" in the US by the pest control company Orkin in 2020, for the sixth consecutive time. It's followed by Los Angeles, New York, Washington, DC, and San Francisco. To help combat the problem, a Chicago animal shelter has placed more than 1000 feral cats (sterilized and vaccinated) outside of homes and businesses since 2012, where they hunt and catch rats while also providing a deterrent simply by their presence.

Rats have the ability to swim up sewer pipes into toilets. Rats will infest any area that provides shelter and easy access to sources of food and water, including under sinks, near garbage, and inside walls or cabinets.

In the spread of disease

Rats can serve as zoonotic vectors for certain pathogens and thus spread disease, such as bubonic plague, Lassa fever, leptospirosis, and Hantavirus infection. Researchers studying New York City wastewater have also cited rats as the potential source of "cryptic" SARS-CoV-2 lineages, due to unknown viral RNA fragments in sewage matching mutations previously shown to make SARS-CoV-2 more adept at rodent-based transmission.

Rats are also associated with human dermatitis because they are frequently infested with blood feeding rodent mites such as the tropical rat mite (Ornithonyssus bacoti) and spiny rat mite ( Laelaps echidnina), which will opportunistically bite and feed on humans, where the condition is known as rat mite dermatitis.

As invasive species

Rat-catching, 1823, by Edwin Landseer, engraving, published by Hurst, Robinson & Co.

When introduced into locations where rats previously did not exist, they can wreak an enormous degree of environmental degradation. Rattus rattus, the black rat, is considered to be one of the world's worst invasive species. Also known as the ship rat, it has been carried worldwide as a stowaway on seagoing vessels for millennia and has usually accompanied men to any new area visited or settled by human beings by sea. Rats first got to countries such as America and Australia by stowing away on ships. The similar species Rattus norvegicus, the brown rat or wharf rat, has also been carried worldwide by ships in recent centuries.

The ship or wharf rat has contributed to the extinction of many species of wildlife, including birds, small mammals, reptiles, invertebrates, and plants, especially on islands. True rats are omnivorous, capable of eating a wide range of plant and animal foods, and have a very high birth rate. When introduced to a new area, they quickly reproduce to take advantage of the new food supply. In particular, they prey on the eggs and young of forest birds, which on isolated islands often have no other predators and thus have no fear of predators. Some experts believe that rats are to blame for between forty percent and sixty percent of all seabird and reptile extinctions, with ninety percent of those occurring on islands. Thus man has indirectly caused the extinction of many species by accidentally introducing rats to new areas.

Rat-free areas

Rats are found in nearly all areas of Earth which are inhabited by human beings. The only rat-free continent is Antarctica, which is too cold for rat survival outdoors, and its lack of human habitation does not provide buildings to shelter them from the weather. However, rats have been introduced to many of the islands near Antarctica, and because of their destructive effect on native flora and fauna, efforts to eradicate them are ongoing. In particular, Bird Island (just off rat-infested South Georgia Island), where breeding seabirds could be badly affected if rats were introduced, is subject to special measures and regularly monitored for rat invasions.

As part of island restoration, some islands' rat populations have been eradicated to protect or restore the ecology. Hawadax Island, Alaska was declared rat free after 229 years and Campbell Island, New Zealand after almost 200 years. Breaksea Island in New Zealand was declared rat free in 1988 after an eradication campaign based on a successful trial on the smaller Hawea Island nearby.

In January 2015, an international "Rat Team" set sail from the Falkland Islands for the British Overseas Territory of South Georgia and the South Sandwich Islands on board a ship carrying three helicopters and 100 tons of rat poison with the objective of "reclaiming the island for its seabirds". Rats have wiped out more than 90% of the seabirds on South Georgia, and the sponsors hope that once the rats are gone, it will regain its former status as home to the greatest concentration of seabirds in the world. The South Georgia Heritage Trust, which organized the mission describes it as "five times larger than any other rodent eradication attempted worldwide". That would be true if it were not for the rat control program in Alberta (see below).

The Canadian province of Alberta is notable for being the largest inhabited area on Earth which is free of true rats due to very aggressive government rat control policies. It has large numbers of native pack rats, also called bushy-tailed wood rats, but they are forest-dwelling vegetarians which are much less destructive than true rats.

Alberta was settled relatively late in North American history and only became a province in 1905. Black rats cannot survive in its climate at all, and brown rats must live near people and in their structures to survive the winters. There are numerous predators in Canada's vast natural areas which will eat non-native rats, so it took until 1950 for invading rats to make their way over land from Eastern Canada. Immediately upon their arrival at the eastern border with Saskatchewan, the Alberta government implemented an extremely aggressive rat control program to stop them from advancing further. A systematic detection and eradication system was used throughout a control zone about 600 kilometres (400 mi) long and 30 kilometres (20 mi) wide along the eastern border to eliminate rat infestations before the rats could spread further into the province. Shotguns, bulldozers, high explosives, poison gas, and incendiaries were used to destroy rats. Numerous farm buildings were destroyed in the process. Initially, tons of arsenic trioxide were spread around thousands of farm yards to poison rats, but soon after the program commenced the rodenticide and medical drug warfarin was introduced, which is much safer for people and more effective at killing rats than arsenic.

Forceful government control measures, strong public support and enthusiastic citizen participation continue to keep rat infestations to a minimum. The effectiveness has been aided by a similar but newer program in Saskatchewan which prevents rats from even reaching the Alberta border. Alberta still employs an armed rat patrol to control rats along Alberta's borders. About ten single rats are found and killed per year, and occasionally a large localized infestation has to be dug out with heavy machinery, but the number of permanent rat infestations is zero.

Ancient Romans did not generally differentiate between rats and mice, instead referring to the former as mus maximus (big mouse) and the latter as mus minimus (little mouse).

Chinese zodiac pendant with 5 rats climbing ruyi, bat at top of pendant

Asian cultures

Two mice, Vietnam Museum of Ethnology - Hanoi

Chuột rước đèn (The mouse carries the lamp), Vietnamese Đông Hồ painting

The rat (sometimes referred to as a mouse) is the first of the twelve animals of the Chinese zodiac. People born in this year are expected to possess qualities associated with rats, including creativity, intelligence, honesty, generosity, ambition, a quick temper and wastefulness. People born in a year of the rat are said to get along well with "monkeys" and "dragons", and to get along poorly with "horses".

Indigenous rats are allowed to run freely throughout the Karni Mata Temple.

In Indian tradition, rats are seen as the vehicle of Ganesha, and a rat's statue is always found in a temple of Ganesh. In the northwestern Indian city of Deshnoke, the rats at the Karni Mata Temple are held to be destined for reincarnation as Sadhus ( Hindu holy men). The attending priests feed milk and grain to the rats, of which the pilgrims also partake.

European cultures

European associations with the rat are generally negative. For instance, "Rats!" is used as a substitute for various vulgar interjections in the English language. These associations do not draw, per se, from any biological or behavioral trait of the rat, but possibly from the association of rats (and fleas) with the 14th-century medieval plague called the Black Death. Rats are seen as vicious, unclean, parasitic animals that steal food and spread disease. In 1522, the rats in Autun, France were charged and put on trial for destroying crops. However, some people in European cultures keep rats as pets and conversely find them to be tame, clean, intelligent, and playful.

Rats are often used in scientific experiments; animal rights activists allege the treatment of rats in this context is cruel. The term "lab rat" is used, typically in a self-effacing manner, to describe a person whose job function requires them to spend a majority of their work time engaged in bench-level research (such as postgraduate students in the sciences).

Terminology

Rats are frequently blamed for damaging food supplies and other goods, or spreading disease. Their reputation has carried into common parlance: in the English language, "rat" is often an insult or is generally used to signify an unscrupulous character; it is also used, as a synonym for the term , to mean an individual who works as a police informant or who has turned state's evidence. Writer/director Preston Sturges created the humorous alias "Ratskywatsky" for a soldier who seduced, impregnated, and abandoned the heroine of his 1944 film, The Miracle of Morgan's Creek. It is a term ( noun and verb) in criminal slang for an informant - "to rat on someone" is to betray them by informing the authorities of a crime or misdeed they committed. Describing a person as "rat-like" usually implies he or she is unattractive and suspicious.

Among trade unions, the word "rat" is also a term for nonunion employers or breakers of union contracts, and this is why unions use inflatable rats.

Imperial Japan depicted as a rat in a World War II United States Navy propaganda poster.

Depictions of rats in fiction are historically inaccurate and negative. The most common falsehood is the squeaking almost always heard in otherwise realistic portrayals (i.e. non anthropomorphic). While the recordings may be of actual squeaking rats, the noise is uncommon - they may do so only if distressed, hurt, or annoyed. Normal vocalizations are very high-pitched, well outside the range of human hearing. Rats are also often cast in vicious and aggressive roles when in fact, their shyness helps keep them undiscovered for so long in an infested home.

The actual portrayals of rats vary from negative to positive with a majority in the negative and ambiguous. The rat plays a villain in several mouse societies; from Brian Jacques's and Robin Jarvis's The Deptford Mice, to the roles of Disney's Professor Ratigan and Kate DiCamillo's Roscuro and Botticelli. They have often been used as a mechanism in horror; being the titular evil in stories like or H.P. Lovecraft's The Rats in the Walls and in films like and . Another terrifying use of rats is as a method of torture, for instance in Room 101 in George Orwell's Nineteen Eighty-Four or The Pit and the Pendulum by Edgar Allan Poe.

Selfish helpfulness -those willing to help for a price- has also been attributed to fictional rats. Templeton, from E. B. White's Charlotte's Web, repeatedly reminds the other characters that he is only involved because it means more food for him, and the cellar-rat of John Masefield's The Midnight Folk requires bribery to be of any assistance.

By contrast, the rats appearing in the Doctor Dolittle books tend to be highly positive and likeable characters, many of whom tell their remarkable life stories in the Mouse and Rat Club established by the animal-loving doctor.

Some fictional works use rats as the main characters. Notable examples include the society created by O'Brien's Mrs. Frisby and the Rats of NIMH, and others include , and Rizzo the Rat from The Muppets. Pixar's 2007 animated film Ratatouille is about a rat described by Roger Ebert as "earnest... lovable, determined, [and] gifted" who lives with a Parisian garbage-boy-turned-chef.

Mon oncle d'Amérique (" My American Uncle"), a 1980 French film, illustrates Henri Laborit's theories on evolutionary psychology and human behaviors by using short sequences in the storyline showing lab rat experiments.

In Harry Turtledove's science fiction novel Homeward Bound, humans unintentionally introduce rats to the ecology at the home world of an alien race which previously invaded Earth and introduced some of its own fauna into its environment. A. Bertram Chandler pitted the space-bound protagonist of a long series of novels, Commodore Grimes, against giant, intelligent rats who took over several stellar systems and enslaved their human inhabitants. " The Stainless Steel Rat" is nickname of the (human) protagonist of a series of humorous science fiction novels written by Harry Harrison.

Wererats, therianthropic creatures able to take the shape of a rat, have appeared in the fantasy or horror genre since the 1970s. The term is a neologism coined in analogy to werewolf.[citation needed ] The concept has since become common in role playing games like Dungeons & Dragons and fantasy fiction like the Anita Blake series.

The Pied Piper

One of the oldest and most historic stories about rats is " The Pied Piper of Hamelin", in which a rat-catcher leads away an infestation with enchanted music. The piper is later refused payment, so he in turn leads away the town's children. This tale, traced to Germany around the late 13th century, has inspired adaptations in film, theatre, literature, and even opera. The subject of much research, some theories have intertwined the tale with events related to the Black Plague, in which black rats played an important role. Fictional works based on the tale that focus heavily on the rat aspect include Pratchett's The Amazing Maurice and his Educated Rodents, and Belgian graphic novel Le Bal du Rat Mort ( The Ball of the Dead Rat). Furthermore, a linguistic phenomenon when a wh-expression drags with it an entire encompassing phrase to the front of the clause has been named pied-piping after "Pied Piper of Hamlin" (see also pied-piping with inversion).

References

Further reading

External links

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Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus . Other rat genera include (pack rats), (bandicoot rats) and (kangaroo rats). Rats are typically distinguished from mice by their size.

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Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is a subject of active research. They became the dominant terrestrial vertebrates after the Triassic-Jurassic extinction event 201.3 mya and their dominance continued throughout the Jurassic and Cretaceous periods. The fossil record shows that birds are feathered dinosaurs, having evolved from earlier theropods during the Late Jurassic epoch, and are the only dinosaur lineage known to have survived the Cretaceous-Paleogene extinction event approximately 66 mya. Dinosaurs can therefore be divided into avian dinosaurs-birds-and the extinct non-avian dinosaurs, which are all dinosaurs other than birds.

Dinosaurs are varied from taxonomic, morphological and ecological standpoints. Birds, at over 10,700 living species, are among the most diverse groups of vertebrates. Using fossil evidence, paleontologists have identified over 900 distinct genera and more than 1,000 different species of non-avian dinosaurs. Dinosaurs are represented on every continent by both extant species (birds) and fossil remains. Through the first half of the 20th century, before birds were recognized as dinosaurs, most of the scientific community believed dinosaurs to have been sluggish and cold-blooded. Most research conducted since the 1970s, however, has indicated that dinosaurs were active animals with elevated metabolisms and numerous adaptations for social interaction. Some were herbivorous, others carnivorous. Evidence suggests that all dinosaurs were egg-laying, and that nest-building was a trait shared by many dinosaurs, both avian and non-avian.

While dinosaurs were ancestrally bipedal, many extinct groups included quadrupedal species, and some were able to shift between these stances. Elaborate display structures such as horns or crests are common to all dinosaur groups, and some extinct groups developed skeletal modifications such as bony armor and spines. While the dinosaurs' modern-day surviving avian lineage (birds) are generally small due to the constraints of flight, many prehistoric dinosaurs (non-avian and avian) were large-bodied-the largest sauropod dinosaurs are estimated to have reached lengths of 39.7 meters (130 feet) and heights of 18 m (59 ft) and were the largest land animals of all time. The misconception that non-avian dinosaurs were uniformly gigantic is based in part on preservation bias, as large, sturdy bones are more likely to last until they are fossilized. Many dinosaurs were quite small, some measuring about 50 centimeters (20 inches) in length.

The first dinosaur fossils were recognized in the early 19th century, with the name "dinosaur" (meaning "terrible lizard") being coined by Sir Richard Owen in 1842 to refer to these "great fossil lizards". Since then, mounted fossil dinosaur skeletons have been major attractions at museums worldwide, and dinosaurs have become an enduring part of popular culture. The large sizes of some dinosaurs, as well as their seemingly monstrous and fantastic nature, have ensured their regular appearance in best-selling books and films, such as Jurassic Park. Persistent public enthusiasm for the animals has resulted in significant funding for dinosaur science, and new discoveries are regularly covered by the media.

Definition

Under phylogenetic nomenclature, dinosaurs are usually defined as the group consisting of the most recent common ancestor (MRCA) of Triceratops and modern birds (Neornithes), and all its descendants. It has also been suggested that Dinosauria be defined with respect to the MRCA of Megalosaurus and , because these were two of the three genera cited by Richard Owen when he recognized the Dinosauria. Both definitions result in the same set of animals being defined as dinosaurs: "Dinosauria = Ornithischia + Saurischia". This definition includes major groups such as ankylosaurians (armored herbivorous quadrupeds), stegosaurians (plated herbivorous quadrupeds), ceratopsians (bipedal or quadrupedal herbivores with neck frills), pachycephalosaurians (bipedal herbivores with thick skulls), ornithopods (bipedal or quadrupedal herbivores including " duck-bills"), theropods (mostly bipedal carnivores and birds), and sauropodomorphs (mostly large herbivorous quadrupeds with long necks and tails).

Birds are the sole surviving dinosaurs. In traditional taxonomy, birds were considered a separate class that had evolved from dinosaurs, a distinct superorder. However, a majority of contemporary paleontologists concerned with dinosaurs reject the traditional style of classification in favor of phylogenetic taxonomy; this approach requires that, for a group to be natural, all descendants of members of the group must be included in the group. Birds belong to the dinosaur subgroup Maniraptora, which are coelurosaurs, which are theropods, which are saurischians.

Research by Matthew G. Baron, David B. Norman, and Paul M. Barrett in 2017 suggested a radical revision of dinosaurian systematics. Phylogenetic analysis by Baron et al. recovered the Ornithischia as being closer to the Theropoda than the Sauropodomorpha, as opposed to the traditional union of theropods with sauropodomorphs. This would cause sauropods and kin to fall outside traditional dinosaurs, so they re-defined Dinosauria as the last common ancestor of Triceratops horridus, Passer domesticus and Diplodocus carnegii, and all of its descendants, to ensure that sauropods and kin remain included as dinosaurs. They also resurrected the clade Ornithoscelida to refer to the group containing Ornithischia and Theropoda.

General description

Using one of the above definitions, dinosaurs can be generally described as archosaurs with hind limbs held erect beneath the body. Other prehistoric animals, including pterosaurs, mosasaurs, ichthyosaurs, plesiosaurs, and Dimetrodon, while often popularly conceived of as dinosaurs, are not taxonomically classified as dinosaurs. Pterosaurs are distantly related to dinosaurs, being members of the clade Ornithodira. The other groups mentioned are, like dinosaurs and pterosaurs, members of Sauropsida (the reptile and bird clade), except Dimetrodon (which is a synapsid). None of them had the erect hind limb posture characteristic of true dinosaurs.

Dinosaurs were the dominant terrestrial vertebrates of the Mesozoic Era, especially the Jurassic and Cretaceous periods. Other groups of animals were restricted in size and niches; mammals, for example, rarely exceeded the size of a domestic cat, and were generally rodent-sized carnivores of small prey. They have always been recognized as an extremely varied group of animals; over 900 non-avian dinosaur genera have been identified with certainty as of 2018, and the total number of genera preserved in the fossil record has been estimated at around 1850, nearly 75% of which remain to be discovered, and 1124 species by 2016. A 1995 study predicted that about 3,400 dinosaur genera ever existed, including many that would not have been preserved in the fossil record.

In 2016, the estimated number of dinosaur species that existed in the Mesozoic was 1,543-2,468. In 2021, the number of modern-day birds (avian dinosaurs) was estimated to be at 10,806 species. Some are herbivorous, others carnivorous, including seed-eaters, fish-eaters, insectivores, and omnivores. While dinosaurs were ancestrally bipedal (as are all modern birds), some prehistoric species were quadrupeds, and others, such as Anchisaurus and Iguanodon, could walk just as easily on two or four legs. Cranial modifications like horns and crests are common dinosaurian traits, and some extinct species had bony armor. Although known for large size, many Mesozoic dinosaurs were human-sized or smaller, and modern birds are generally small in size. Dinosaurs today inhabit every continent, and fossils show that they had achieved global distribution by at least the Early Jurassic epoch. Modern birds inhabit most available habitats, from terrestrial to marine, and there is evidence that some non-avian dinosaurs (such as Microraptor) could fly or at least glide, and others, such as spinosaurids, had semiaquatic habits.

Distinguishing anatomical features

In the skull, a supratemporal fossa (excavation) is present in front of the supratemporal fenestra, the main opening in the rear skull roof

Epipophyses, obliquely backward-pointing processes on the rear top corners of the anterior (front) neck vertebrae behind the atlas and axis, the first two neck vertebrae

Apex of a deltopectoral crest (a projection on which the deltopectoral muscles attach) located at or more than 30% down the length of the humerus (upper arm bone)

Radius, a lower arm bone, shorter than 80% of humerus length

Fourth trochanter (projection where the caudofemoralis muscle attaches on the inner rear shaft) on the femur (thigh bone) is a sharp flange

Fourth trochanter asymmetrical, with distal, lower, margin forming a steeper angle to the shaft

On the astragalus and calcaneum, upper ankle bones, the proximal articular facet, the top connecting surface, for the fibula occupies less than 30% of the transverse width of the element

Exoccipitals (bones at the back of the skull) do not meet along the midline on the floor of the endocranial cavity, the inner space of the braincase

In the pelvis, the proximal articular surfaces of the ischium with the ilium and the pubis are separated by a large concave surface (on the upper side of the ischium a part of the open hip joint is located between the contacts with the pubic bone and the ilium)

Cnemial crest on the tibia (protruding part of the top surface of the shinbone) arcs anterolaterally (curves to the front and the outer side)

Distinct proximodistally oriented (vertical) ridge present on the posterior face of the distal end of the tibia (the rear surface of the lower end of the shinbone)

Concave articular surface for the fibula of the calcaneum (the top surface of the calcaneum, where it touches the fibula, has a hollow profile)

A variety of other skeletal features are shared by dinosaurs. However, because they either are common to other groups of archosaurs or were not present in all early dinosaurs, these features are not considered to be synapomorphies. For example, as diapsids, dinosaurs ancestrally had two pairs of Infratemporal fenestrae (openings in the skull behind the eyes), and as members of the diapsid group Archosauria, had additional openings in the snout and lower jaw. Additionally, several characteristics once thought to be synapomorphies are now known to have appeared before dinosaurs, or were absent in the earliest dinosaurs and independently evolved by different dinosaur groups. These include an elongated scapula, or shoulder blade; a sacrum composed of three or more fused vertebrae (three are found in some other archosaurs, but only two are found in Herrerasaurus); and a perforate acetabulum, or hip socket, with a hole at the center of its inside surface (closed in Saturnalia tupiniquim, for example). Another difficulty of determining distinctly dinosaurian features is that early dinosaurs and other archosaurs from the Late Triassic epoch are often poorly known and were similar in many ways; these animals have sometimes been misidentified in the literature.

Dinosaurs stand with their hind limbs erect in a manner similar to most modern mammals, but distinct from most other reptiles, whose limbs sprawl out to either side. This posture is due to the development of a laterally facing recess in the pelvis (usually an open socket) and a corresponding inwardly facing distinct head on the femur. Their erect posture enabled early dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels that surpassed those of "sprawling" reptiles. Erect limbs probably also helped support the evolution of large size by reducing bending stresses on limbs. Some non-dinosaurian archosaurs, including rauisuchians, also had erect limbs but achieved this by a "pillar-erect" configuration of the hip joint, where instead of having a projection from the femur insert on a socket on the hip, the upper pelvic bone was rotated to form an overhanging shelf.

History of study

Pre-scientific history

Dinosaur fossils have been known for millennia, although their true nature was not recognized. The Chinese considered them to be dragon bones and documented them as such. For example, Huayang Guo Zhi (華陽國志), a gazetteer compiled by Chang Qu (常璩) during the Western Jin Dynasty (265-316), reported the discovery of dragon bones at Wucheng in Sichuan Province. Villagers in central China have long unearthed fossilized "dragon bones" for use in traditional medicines. In Europe, dinosaur fossils were generally believed to be the remains of giants and other biblical creatures.

Early dinosaur research

William Buckland

Scholarly descriptions of what would now be recognized as dinosaur bones first appeared in the late 17th century in England. Part of a bone, now known to have been the femur of a Megalosaurus, was recovered from a limestone quarry at Cornwell near Chipping Norton, Oxfordshire, in 1676. The fragment was sent to Robert Plot, Professor of Chemistry at the University of Oxford and first curator of the Ashmolean Museum, who published a description in his The Natural History of Oxford-shire (1677). He correctly identified the bone as the lower extremity of the femur of a large animal, and recognized that it was too large to belong to any known species. He, therefore, concluded it to be the femur of a huge human, perhaps a Titan or another type of giant featured in legends. Edward Lhuyd, a friend of Sir Isaac Newton, published Lithophylacii Britannici ichnographia (1699), the first scientific treatment of what would now be recognized as a dinosaur when he described and named a sauropod tooth, " Rutellum impicatum", that had been found in Caswell, near Witney, Oxfordshire.

Between 1815 and 1824, the Rev William Buckland, the first Reader of Geology at the University of Oxford, collected more fossilized bones of Megalosaurus and became the first person to describe a non-avian dinosaur in a scientific journal. The second non-avian dinosaur genus to be identified, Iguanodon, was according to legend discovered in 1822 by Mary Ann Mantell - the wife of English geologist Gideon Mantell who in fact had required remains years earlier. Gideon Mantell recognized similarities between his fossils and the bones of modern iguanas. He published his findings in 1825.

The study of these "great fossil lizards" soon became of great interest to European and American scientists, and in 1842 the English paleontologist Sir Richard Owen coined the term "dinosaur", using it to refer to the "distinct tribe or sub-order of Saurian Reptiles" that were then being recognized in England and around the world. The term is derived from Ancient Greek δεινός (deinos) 'terrible, potent or fearfully great', and σαῦρος (sauros) 'lizard or reptile'. Though the taxonomic name has often been interpreted as a reference to dinosaurs' teeth, claws, and other fearsome characteristics, Owen intended it also to evoke their size and majesty. Owen recognized that the remains that had been found so far, Iguanodon, Megalosaurus and Hylaeosaurus, shared distinctive features, and so decided to present them as a distinct taxonomic group. As clarified by British geologist and historian Hugh Torrens, Owen had given a presentation about fossil reptiles to the British Association for the Advancement of Science in 1841, but reports of the time show that Owen did not mention the word "dinosaur", nor recognize dinosaurs as a distinct group of reptiles in his address. He introduced the Dinosauria only in the revised text version of his talk published in April of 1842. With the backing of Prince Albert, the husband of Queen Victoria, Owen established the Natural History Museum, London, to display the national collection of dinosaur fossils and other biological and geological exhibits.

Discoveries in North America

In 1858, William Parker Foulke discovered the first known American dinosaur, in marl pits in the small town of Haddonfield, New Jersey. (Although fossils had been found before, their nature had not been correctly discerned.) The creature was named Hadrosaurus foulkii. It was an extremely important find: Hadrosaurus was one of the first nearly complete dinosaur skeletons found ( the first was in 1834, in Maidstone, England), and it was clearly a bipedal creature. This was a revolutionary discovery as, until that point, most scientists had believed dinosaurs walked on four feet, like other lizards. Foulke's discoveries sparked a wave of interests in dinosaurs in the United States, known as dinosaur mania.

Dinosaur mania was exemplified by the fierce rivalry between Edward Drinker Cope and Othniel Charles Marsh, both of whom raced to be the first to find new dinosaurs in what came to be known as the Bone Wars. This fight between the two scientists lasted for over 30 years, ending in 1897 when Cope died after spending his entire fortune on the dinosaur hunt. Many valuable dinosaur specimens were damaged or destroyed due to the pair's rough methods: for example, their diggers often used dynamite to unearth bones. Modern paleontologists would find such methods crude and unacceptable, since blasting easily destroys fossil and stratigraphic evidence. Despite their unrefined methods, the contributions of Cope and Marsh to paleontology were vast: Marsh unearthed 86 new species of dinosaur and Cope discovered 56, a total of 142 new species. Cope's collection is now at the American Museum of Natural History in New York City, while Marsh's is at the Peabody Museum of Natural History at Yale University.

"Dinosaur renaissance" and beyond

World War II caused a pause in palaeontological research; after the war, research attention was also diverted increasingly to fossil mammals rather than dinosaurs, which were seen as sluggish and cold-blooded. At the end of the 1960s, however, the field of dinosaur research experienced a surge in activity that remains ongoing. Several seminal studies led to this activity. First, John Ostrom discovered the bird-like dromaeosaurid theropod Deinonychus and described it in 1969. Its anatomy indicated that it was an active predator that was likely warm-blooded, in marked contrast to the then-prevailing image of dinosaurs. Concurrently, Robert T. Bakker published a series of studies that likewise argued for active lifestyles in dinosaurs based on anatomical and ecological evidence (see § Physiology), which were subsequently summarized in his 1986 book The Dinosaur Heresies.

Paleontologist Robert T. Bakker with a mounted skeleton of a tyrannosaurid (Gorgosaurus libratus)

New revelations were supported by an increase in dinosaur discoveries. Major new dinosaur discoveries have been made by paleontologists working in previously unexplored regions, including India, South America, Madagascar, Antarctica, and most significantly China. Across theropods, sauropodomorphs, and ornithischians, the number of named genera began to increase exponentially in the 1990s. As of 2008, over 30 new species of dinosaurs were named each year. At least sauropodomorphs experienced a further increase in the number of named species in the 2010s, with an average of 9.3 new species having been named each year between 2009 and 2020. As a consequence, more sauropodomorphs were named between 1990 and 2020 than in all previous years combined. These new localities also led to improvements in overall specimen quality, with new species being increasingly named not on scrappy fossils but on more complete skeletons, sometimes from multiple individuals. Better specimens also led to new species being invalidated less frequently. Asian localities have produced the most complete theropod specimens, while North American localities have produced the most complete sauropodomorph specimens.

Prior to the dinosaur renaissance, dinosaurs were mostly classified using the traditional rank-based system of Linnaean taxonomy. The renaissance was also accompanied by the increasingly widespread application of cladistics, a more objective method of classification based on ancestry and shared traits, which has proved tremendously useful in the study of dinosaur systematics and evolution. Cladistic analysis, among other techniques, helps to compensate for an often incomplete and fragmentary fossil record. Reference books summarizing the state of dinosaur research, such as David B. Weishampel and colleagues' The Dinosauria, made knowledge more accessible and spurred further interest in dinosaur research. The release of the first and second editions of The Dinosauria in 1990 and 2004, and of a review paper by Paul Sereno in 1998, were accompanied by increases in the number of published phylogenetic trees for dinosaurs.

Soft tissue and molecular preservation

Dinosaur fossils are not limited to bones, but also include imprints or mineralized remains of skin coverings, organs, and other tissues. Of these, skin coverings based on keratin proteins are most easily preserved because of their cross-linked, hydrophobic molecular structure. Fossils of keratin-based skin coverings or bony skin coverings are known from most major groups of dinosaurs. Dinosaur fossils with scaly skin impressions have been found since the 19th century. Samuel Beckles discovered a sauropod forelimb with preserved skin in 1852 that was incorrectly attributed to a crocodile; it was correctly attributed by Marsh in 1888 and subject to further study by Reginald Hooley in 1917. Among ornithischians, in 1884 Jacob Wortman found skin impressions on the first known specimen of Edmontosaurus annectens, which were largely destroyed during the specimen's excavation. Owen and Hooley subsequently described skin impressions of Hypsilophodon and Iguanodon in 1885 and 1917. Since then, scale impressions have been most frequently found among hadrosaurids, where the impressions are known from nearly the entire body across multiple specimens.

Starting from the 1990s, major discoveries of exceptionally preserved fossils in deposits known as conservation Lagerstätten contributed to research on dinosaur soft tissues. Chiefly among these were the rocks that produced the Jehol (Early Cretaceous) and Yanliao (Mid-to-Late Jurassic) biotas of northeastern China, from which hundreds of dinosaur specimens bearing impressions of feather-like structures (both closely related to birds and otherwise, see § Origin of birds) have been described by Xing Xu and colleagues. In living reptiles and mammals, pigment-storing cellular structures known as melanosomes are partially responsible for producing colouration. Both chemical traces of melanin and characteristically-shaped melanosomes have been reported from feathers and scales of Jehol and Yanliao dinosaurs, including both theropods and ornithischians. This has enabled multiple full-body reconstructions of dinosaur colouration, such as for Sinosauropteryx and Psittacosaurus by Jakob Vinther and colleagues, and similar techniques have also been extended to dinosaur fossils from other localities. (However, some researchers have also suggested that fossilized melanosomes represent bacterial remains.) Stomach contents in some Jehol and Yanliao dinosaurs closely related to birds have also provided indirect indications of diet and digestive system anatomy (e.g., crops). More concrete evidence of internal anatomy has been reported in from the Pietraroja Plattenkalk of Italy. It preserves portions of the intestines, colon, liver, muscles, and windpipe.

Concurrently, a line of work led by Mary Higby Schweitzer, Jack Horner, and colleagues reported various occurrences of preserved soft tissues and proteins within dinosaur bone fossils. Various mineralized structures that likely represented red blood cells and collagen fibres had been found by Schweitzer and others in tyrannosaurid bones as early as 1991. However, in 2005, Schweitzer and colleagues reported that a femur of Tyrannosaurus preserved soft, flexible tissue within, including blood vessels, bone matrix, and connective tissue (bone fibers) that had retained their microscopic structure. This discovery suggested that original soft tissues could be preserved over geological time, with multiple mechanisms having been proposed. Later, in 2009, Schweitzer and colleagues reported that a Brachylophosaurus femur preserved similar microstructures, and immunohistochemical techniques (based on antibody binding) demonstrated the presence of proteins such as collagen, elastin, and laminin. Both specimens yielded collagen protein sequences that were viable for molecular phylogenetic analyses, which grouped them with birds as would be expected. The extraction of fragmentary DNA has also been reported for both of these fossils, along with a specimen of Hypacrosaurus. In 2015, Sergio Bertazzo and colleagues reported the preservation of collagen fibres and red blood cells in eight Cretaceous dinosaur specimens that did not show any signs of exceptional preservation, indicating that soft tissue may be preserved more commonly than previously thought. Suggestions that these structures represent bacterial biofilms have been rejected, but cross-contamination remains a possibility that is difficult to detect.

Evolutionary history

Origins and early evolution

The early dinosaurs Herrerasaurus (large), Eoraptor (small) and a Plateosaurus skull, from the Triassic

Dinosaurs diverged from their archosaur ancestors during the Middle to Late Triassic epochs, roughly 20 million years after the devastating Permian-Triassic extinction event wiped out an estimated 96% of all marine species and 70% of terrestrial vertebrate species approximately 252 million years ago. The oldest dinosaur fossils known from substantial remains date to the Carnian epoch of the Triassic period and have been found primarily in the Ischigualasto and Santa Maria Formations of Argentina, and the Pebbly Arkose Formation of Zimbabwe.

The Ischigualasto Formation ( radiometrically dated at 231-230 million years old) has produced the early saurischian , originally considered a member of the Herrerasauridae but now considered to be an early sauropodomorph, along with the herrerasaurids Herrerasaurus and Sanjuansaurus, and the sauropodomorphs Chromogisaurus, Eodromaeus, and . Eoraptor's likely resemblance to the common ancestor of all dinosaurs suggests that the first dinosaurs would have been small, bipedal predators. The Santa Maria Formation (radiometrically dated to be older, at 233.23 million years old) has produced the herrerasaurids Gnathovorax and Staurikosaurus, along with the sauropodomorphs Bagualosaurus, Buriolestes, Guaibasaurus, Macrocollum, Nhandumirim, Pampadromaeus, Saturnalia, and Unaysaurus. The Pebbly Arkose Formation, which is of uncertain age but was likely comparable to the other two, has produced the sauropodomorph Mbiresaurus, along with an unnamed herrerasaurid.

Less well-preserved remains of the sauropodomorphs Jaklapallisaurus and , along with the early saurischian Alwalkeria, are known from the Upper Maleri and Lower Maleri Formations of India. The Carnian-aged Chañares Formation of Argentina preserves primitive, dinosaur-like ornithodirans such as Lagosuchus and Lagerpeton in Argentina, making it another important site for understanding dinosaur evolution. These ornithodirans support the model of early dinosaurs as small, bipedal predators. Dinosaurs may have appeared as early as the Anisian epoch of the Triassic, approximately 243 million years ago, which is the age of Nyasasaurus from the Manda Formation of Tanzania. However, its known fossils are too fragmentary to identify it as a dinosaur or only a close relative. The referral of the Manda Formation to the Anisian is also uncertain. Regardless, dinosaurs existed alongside non-dinosaurian ornithodirans for a period of time, with estimates ranging from 5-10 million years to 21 million years.

When dinosaurs appeared, they were not the dominant terrestrial animals. The terrestrial habitats were occupied by various types of archosauromorphs and therapsids, like cynodonts and rhynchosaurs. Their main competitors were the pseudosuchians, such as aetosaurs, ornithosuchids and rauisuchians, which were more successful than the dinosaurs. Most of these other animals became extinct in the Triassic, in one of two events. First, at about 215 million years ago, a variety of basal archosauromorphs, including the protorosaurs, became extinct. This was followed by the Triassic-Jurassic extinction event (about 201 million years ago), that saw the end of most of the other groups of early archosaurs, like aetosaurs, ornithosuchids, phytosaurs, and rauisuchians. Rhynchosaurs and dicynodonts survived (at least in some areas) at least as late as early -mid Norian and late Norian or earliest Rhaetian stages, respectively, and the exact date of their extinction is uncertain. These losses left behind a land fauna of crocodylomorphs, dinosaurs, mammals, pterosaurians, and turtles. The first few lines of early dinosaurs diversified through the Carnian and Norian stages of the Triassic, possibly by occupying the niches of the groups that became extinct. Also notably, there was a heightened rate of extinction during the Carnian pluvial event.

Evolution and paleobiogeography

The supercontinent Pangaea in the early Mesozoic (around 200 million years ago)

Dinosaur evolution after the Triassic followed changes in vegetation and the location of continents. In the Late Triassic and Early Jurassic, the continents were connected as the single landmass Pangaea, and there was a worldwide dinosaur fauna mostly composed of coelophysoid carnivores and early sauropodomorph herbivores. Gymnosperm plants (particularly conifers), a potential food source, radiated in the Late Triassic. Early sauropodomorphs did not have sophisticated mechanisms for processing food in the mouth, and so must have employed other means of breaking down food farther along the digestive tract. The general homogeneity of dinosaurian faunas continued into the Middle and Late Jurassic, where most localities had predators consisting of ceratosaurians, megalosauroids, and allosauroids, and herbivores consisting of stegosaurian ornithischians and large sauropods. Examples of this include the Morrison Formation of North America and Tendaguru Beds of Tanzania. Dinosaurs in China show some differences, with specialized metriacanthosaurid theropods and unusual, long-necked sauropods like Mamenchisaurus. Ankylosaurians and ornithopods were also becoming more common, but primitive sauropodomorphs had become extinct. Conifers and pteridophytes were the most common plants. Sauropods, like earlier sauropodomorphs, were not oral processors, but ornithischians were evolving various means of dealing with food in the mouth, including potential cheek-like organs to keep food in the mouth, and jaw motions to grind food. Another notable evolutionary event of the Jurassic was the appearance of true birds, descended from maniraptoran coelurosaurians.

By the Early Cretaceous and the ongoing breakup of Pangaea, dinosaurs were becoming strongly differentiated by landmass. The earliest part of this time saw the spread of ankylosaurians, iguanodontians, and brachiosaurids through Europe, North America, and northern Africa. These were later supplemented or replaced in Africa by large spinosaurid and carcharodontosaurid theropods, and rebbachisaurid and titanosaurian sauropods, also found in South America. In Asia, maniraptoran coelurosaurians like dromaeosaurids, troodontids, and oviraptorosaurians became the common theropods, and ankylosaurids and early ceratopsians like Psittacosaurus became important herbivores. Meanwhile, Australia was home to a fauna of basal ankylosaurians, hypsilophodonts, and iguanodontians. The stegosaurians appear to have gone extinct at some point in the late Early Cretaceous or early Late Cretaceous. A major change in the Early Cretaceous, which would be amplified in the Late Cretaceous, was the evolution of flowering plants. At the same time, several groups of dinosaurian herbivores evolved more sophisticated ways to orally process food. Ceratopsians developed a method of slicing with teeth stacked on each other in batteries, and iguanodontians refined a method of grinding with dental batteries, taken to its extreme in hadrosaurids. Some sauropods also evolved tooth batteries, best exemplified by the rebbachisaurid Nigersaurus.

There were three general dinosaur faunas in the Late Cretaceous. In the northern continents of North America and Asia, the major theropods were tyrannosaurids and various types of smaller maniraptoran theropods, with a predominantly ornithischian herbivore assemblage of hadrosaurids, ceratopsians, ankylosaurids, and pachycephalosaurians. In the southern continents that had made up the now-splitting supercontinent Gondwana, abelisaurids were the common theropods, and titanosaurian sauropods the common herbivores. Finally, in Europe, dromaeosaurids, rhabdodontid iguanodontians, nodosaurid ankylosaurians, and titanosaurian sauropods were prevalent. Flowering plants were greatly radiating, with the first grasses appearing by the end of the Cretaceous. Grinding hadrosaurids and shearing ceratopsians became very diverse across North America and Asia. Theropods were also radiating as herbivores or omnivores, with therizinosaurians and ornithomimosaurians becoming common.

The Cretaceous-Paleogene extinction event, which occurred approximately 66 million years ago at the end of the Cretaceous, caused the extinction of all dinosaur groups except for the neornithine birds. Some other diapsid groups, including crocodilians, dyrosaurs, sebecosuchians, turtles, lizards, snakes, sphenodontians, and choristoderans, also survived the event.

The surviving lineages of neornithine birds, including the ancestors of modern ratites, ducks and chickens, and a variety of waterbirds, diversified rapidly at the beginning of the Paleogene period, entering ecological niches left vacant by the extinction of Mesozoic dinosaur groups such as the arboreal enantiornithines, aquatic hesperornithines, and even the larger terrestrial theropods (in the form of , eogruiids, bathornithids, ratites, geranoidids, mihirungs, and " terror birds"). It is often stated that mammals out-competed the neornithines for dominance of most terrestrial niches but many of these groups co-existed with rich mammalian faunas for most of the Cenozoic Era. Terror birds and bathornithids occupied carnivorous guilds alongside predatory mammals, and ratites are still fairly successful as mid-sized herbivores; eogruiids similarly lasted from the Eocene to Pliocene, becoming extinct only very recently after over 20 million years of co-existence with many mammal groups.

Classification

Dinosaurs belong to a group known as archosaurs, which also includes modern crocodilians. Within the archosaur group, dinosaurs are differentiated most noticeably by their gait. Dinosaur legs extend directly beneath the body, whereas the legs of lizards and crocodilians sprawl out to either side.

Collectively, dinosaurs as a clade are divided into two primary branches, Saurischia and Ornithischia. Saurischia includes those taxa sharing a more recent common ancestor with birds than with Ornithischia, while Ornithischia includes all taxa sharing a more recent common ancestor with Triceratops than with Saurischia. Anatomically, these two groups can be distinguished most noticeably by their pelvic structure. Early saurischians-"lizard-hipped", from the Greek () meaning "lizard" and () meaning "hip joint"-retained the hip structure of their ancestors, with a pubis bone directed cranially, or forward. This basic form was modified by rotating the pubis backward to varying degrees in several groups ( Herrerasaurus, therizinosauroids, dromaeosaurids, and birds). Saurischia includes the theropods (exclusively bipedal and with a wide variety of diets) and sauropodomorphs (long-necked herbivores which include advanced, quadrupedal groups).

By contrast, ornithischians-"bird-hipped", from the Greek ornitheios (ὀρνίθειος) meaning "of a bird" and ischion (ἰσχίον) meaning "hip joint"-had a pelvis that superficially resembled a bird's pelvis: the pubic bone was oriented caudally (rear-pointing). Unlike birds, the ornithischian pubis also usually had an additional forward-pointing process. Ornithischia includes a variety of species that were primarily herbivores.

Despite the terms "bird hip" (Ornithischia) and "lizard hip" (Saurischia), birds are not part of Ornithischia. Birds instead belong to Saurischia, the "lizard-hipped" dinosaurs-birds evolved from earlier dinosaurs with "lizard hips".

The following is a simplified classification of dinosaur groups based on their evolutionary relationships, and those of the main dinosaur groups Theropoda, Sauropodomorpha and Ornithischia, compiled by Justin Tweet. Further details and other hypotheses of classification may be found on individual articles.

Restoration of six ornithopods; far left: Camptosaurus, left: Iguanodon, center background: Shantungosaurus, center foreground: Dryosaurus, right: Corythosaurus, far right (large) Tenontosaurus.

†Ornithischia ("bird-hipped"; diverse bipedal and quadrupedal herbivores)

†Thyreophora (armored dinosaurs; bipeds and quadrupeds)

†Eurypoda (heavy, quadrupedal thyreophorans)

Restoration of four ceratopsids: top left - Triceratops, top right - Styracosaurus, bottom left - Anchiceratops, bottom right - Chasmosaurus.

†Ornithopoda (various sizes; bipeds and quadrupeds; evolved a method of chewing using skull flexibility and numerous teeth)

†Elasmaria (mostly southern ornithopods with mineralized plates along the ribs; may be thescelosaurids)

†Rhabdodontomorpha (with distinctive dentition)

†Hadrosauriformes (ancestrally had a thumb spike; large quadrupedal herbivores, with teeth merged into dental batteries)

Restoration of four macronarian sauropods: from left to right Camarasaurus, Brachiosaurus, Giraffatitan, and Euhelopus

†Sauropoda (very large and heavy; quadrupedal)

†Diplodocoidea (skulls and tails elongated; teeth typically narrow and pencil-like)

†Macronaria (boxy skulls; spoon- or pencil-shaped teeth)

†Euhelopodidae (stocky, mostly Asian)

†Titanosauria (diverse; stocky, with wide hips; most common in the Late Cretaceous of southern continents)

†Ceratosauria (generally elaborately horned carnivores that existed from the Jurassic to Cretaceous periods, originally included Coelophysoidea)

Coelurosauria (feathered theropods, with a range of body sizes and niches)

†"Nexus of basal coelurosaurs" (used by Tweet to denote well-known taxa with unstable positions at the base of Coelurosauria)

Tyrannoraptora ("tyrant thieves")

†Ornithomimosauria (small-headed, mostly toothless, omnivorous or possible herbivores)

Restoration of six dromaeosaurid theropods: from left to right Microraptor, Velociraptor, Austroraptor, Dromaeosaurus, Utahraptor, and Deinonychus

†Caenagnathidae (toothless oviraptorosaurs known from North America and Asia)

†Oviraptoridae (characterized by two bony projections at the back of the mouth; exclusive to Asia)

Paraves (avialans and their closest relatives)

†Microraptoria (characterized by large wings on both the arms and legs; may have been capable of powered flight)

†Eudromaeosauria (hunters with greatly enlarged sickle claws)

Avialae (modern birds and extinct relatives)

Timeline of major groups

Timeline of major dinosaur groups per Holtz (2007).

Paleobiology

Knowledge about dinosaurs is derived from a variety of fossil and non-fossil records, including fossilized bones, feces, trackways, gastroliths, feathers, impressions of skin, internal organs and other soft tissues. Many fields of study contribute to our understanding of dinosaurs, including physics (especially biomechanics), chemistry, biology, and the Earth sciences (of which paleontology is a sub-discipline). Two topics of particular interest and study have been dinosaur size and behavior.

Scale diagram comparing the average human to the longest known dinosaurs in five major clades:

Current evidence suggests that dinosaur average size varied through the Triassic, Early Jurassic, Late Jurassic and Cretaceous. Predatory theropod dinosaurs, which occupied most terrestrial carnivore niches during the Mesozoic, most often fall into the 100 to kg (220 to lb) category when sorted by estimated weight into categories based on order of magnitude, whereas recent predatory carnivoran mammals peak in the 10 to 100 kg (22 to 220 lb) category. The mode of Mesozoic dinosaur body masses is between 1 and 10 metric tons (1.1 and 11.0 short tons). This contrasts sharply with the average size of Cenozoic mammals, estimated by the National Museum of Natural History as about 2 to 5 kg (4.4 to 11.0 lb).

The sauropods were the largest and heaviest dinosaurs. For much of the dinosaur era, the smallest sauropods were larger than anything else in their habitat, and the largest was an order of magnitude more massive than anything else that has since walked the Earth. Giant prehistoric mammals such as Paraceratherium (the largest land mammal ever) were dwarfed by the giant sauropods, and only modern whales approach or surpass them in size. There are several proposed advantages for the large size of sauropods, including protection from predation, reduction of energy use, and longevity, but it may be that the most important advantage was dietary. Large animals are more efficient at digestion than small animals, because food spends more time in their digestive systems. This also permits them to subsist on food with lower nutritive value than smaller animals. Sauropod remains are mostly found in rock formations interpreted as dry or seasonally dry, and the ability to eat large quantities of low-nutrient browse would have been advantageous in such environments.

Largest and smallest

The tallest and heaviest dinosaur known from good skeletons is Giraffatitan brancai (previously classified as a species of Brachiosaurus). Its remains were discovered in Tanzania between 1907 and 1912. Bones from several similar-sized individuals were incorporated into the skeleton now mounted and on display at the Museum für Naturkunde in Berlin; this mount is 12 meters (39 ft) tall and 21.8 to 22.5 meters (72 to 74 ft) long, and would have belonged to an animal that weighed between and kilograms ( and lb). The longest complete dinosaur is the 27 meters (89 ft) long Diplodocus, which was discovered in Wyoming in the United States and displayed in Pittsburgh's Carnegie Museum of Natural History in 1907. The longest dinosaur known from good fossil material is Patagotitan: the skeleton mount in the American Museum of Natural History in New York is 37 meters (121 ft) long. The Museo Municipal Carmen Funes in Plaza Huincul, Argentina, has an Argentinosaurus reconstructed skeleton mount that is 39.7 meters (130 ft) long.

An adult bee hummingbird, the smallest known dinosaur

There were larger dinosaurs, but knowledge of them is based entirely on a small number of fragmentary fossils. Most of the largest herbivorous specimens on record were discovered in the 1970s or later, and include the massive Argentinosaurus, which may have weighed to kilograms (90 to 110 short tons) and reached lengths of 30 to 40 meters (98 to 131 ft); some of the longest were the 33.5-meter (110 ft) long Diplodocus hallorum (formerly Seismosaurus), the 33-to-34-meter (108 to 112 ft) long Supersaurus, and 37-meter (121 ft) long Patagotitan; and the tallest, the 18-meter (59 ft) tall Sauroposeidon, which could have reached a sixth-floor window. The heaviest and longest dinosaur may have been Maraapunisaurus, known only from a now lost partial vertebral neural arch described in 1878. Extrapolating from the illustration of this bone, the animal may have been 58 meters (190 ft) long and weighed kg ( lb). However, as no further evidence of sauropods of this size has been found, and the discoverer, Cope, had made typographic errors before, it is likely to have been an extreme overestimation.

The largest carnivorous dinosaur was Spinosaurus, reaching a length of 12.6 to 18 meters (41 to 59 ft), and weighing 7 to 20.9 metric tons (7.7 to 23.0 short tons). Other large carnivorous theropods included Giganotosaurus, Carcharodontosaurus and Tyrannosaurus. Therizinosaurus and Deinocheirus were among the tallest of the theropods. The largest ornithischian dinosaur was probably the hadrosaurid Shantungosaurus giganteus which measured 16.6 meters (54 ft). The largest individuals may have weighed as much as 16 metric tons (18 short tons).

The smallest dinosaur known is the bee hummingbird, with a length of only 5 centimeters (2.0 in) and mass of around 1.8 g (0.063 oz). The smallest known non- avialan dinosaurs were about the size of pigeons and were those theropods most closely related to birds. For example, Anchiornis huxleyi is currently the smallest non-avialan dinosaur described from an adult specimen, with an estimated weight of 110 g (3.9 oz) and a total skeletal length of 34 centimeters (1.12 ft). The smallest herbivorous non-avialan dinosaurs included Microceratus and Wannanosaurus, at about 60 centimeters (2.0 ft) long each.

Many modern birds are highly social, often found living in flocks. There is general agreement that some behaviors that are common in birds, as well as in crocodiles (closest living relatives of birds), were also common among extinct dinosaur groups. Interpretations of behavior in fossil species are generally based on the pose of skeletons and their habitat, computer simulations of their biomechanics, and comparisons with modern animals in similar ecological niches.

The first potential evidence for herding or flocking as a widespread behavior common to many dinosaur groups in addition to birds was the 1878 discovery of 31 Iguanodon, ornithischians that were then thought to have perished together in Bernissart, Belgium, after they fell into a deep, flooded sinkhole and drowned. Other mass-death sites have been discovered subsequently. Those, along with multiple trackways, suggest that gregarious behavior was common in many early dinosaur species. Trackways of hundreds or even thousands of herbivores indicate that duck-billed (hadrosaurids) may have moved in great herds, like the American bison or the African springbok. Sauropod tracks document that these animals traveled in groups composed of several different species, at least in Oxfordshire, England, although there is no evidence for specific herd structures. Congregating into herds may have evolved for defense, for migratory purposes, or to provide protection for young. There is evidence that many types of slow-growing dinosaurs, including various theropods, sauropods, ankylosaurians, ornithopods, and ceratopsians, formed aggregations of immature individuals. One example is a site in Inner Mongolia that has yielded remains of over 20 Sinornithomimus, from one to seven years old. This assemblage is interpreted as a social group that was trapped in mud. The interpretation of dinosaurs as gregarious has also extended to depicting carnivorous theropods as pack hunters working together to bring down large prey. However, this lifestyle is uncommon among modern birds, crocodiles, and other reptiles, and the taphonomic evidence suggesting mammal-like pack hunting in such theropods as Deinonychus and Allosaurus can also be interpreted as the results of fatal disputes between feeding animals, as is seen in many modern diapsid predators.

The crests and frills of some dinosaurs, like the marginocephalians, theropods and lambeosaurines, may have been too fragile to be used for active defense, and so they were likely used for sexual or aggressive displays, though little is known about dinosaur mating and territorialism. Head wounds from bites suggest that theropods, at least, engaged in active aggressive confrontations.

From a behavioral standpoint, one of the most valuable dinosaur fossils was discovered in the Gobi Desert in 1971. It included a Velociraptor attacking a Protoceratops, providing evidence that dinosaurs did indeed attack each other. Additional evidence for attacking live prey is the partially healed tail of an Edmontosaurus, a hadrosaurid dinosaur; the tail is damaged in such a way that shows the animal was bitten by a tyrannosaur but survived. Cannibalism amongst some species of dinosaurs was confirmed by tooth marks found in Madagascar in 2003, involving the theropod Majungasaurus.

Comparisons between the scleral rings of dinosaurs and modern birds and reptiles have been used to infer daily activity patterns of dinosaurs. Although it has been suggested that most dinosaurs were active during the day, these comparisons have shown that small predatory dinosaurs such as dromaeosaurids, Juravenator, and Megapnosaurus were likely nocturnal. Large and medium-sized herbivorous and omnivorous dinosaurs such as ceratopsians, sauropodomorphs, hadrosaurids, ornithomimosaurs may have been cathemeral, active during short intervals throughout the day, although the small ornithischian Agilisaurus was inferred to be diurnal.

Based on fossil evidence from dinosaurs such as Oryctodromeus, some ornithischian species seem to have led a partially fossorial (burrowing) lifestyle. Many modern birds are arboreal (tree climbing), and this was also true of many Mesozoic birds, especially the enantiornithines. While some early bird-like species may have already been arboreal as well (including dromaeosaurids) such as Microraptor) most non-avialan dinosaurs seem to have relied on land-based locomotion. A good understanding of how dinosaurs moved on the ground is key to models of dinosaur behavior; the science of biomechanics, pioneered by Robert McNeill Alexander, has provided significant insight in this area. For example, studies of the forces exerted by muscles and gravity on dinosaurs' skeletal structure have investigated how fast dinosaurs could run, whether diplodocids could create sonic booms via whip-like tail snapping, and whether sauropods could float.

Communication

Modern birds are known to communicate using visual and auditory signals, and the wide diversity of visual display structures among fossil dinosaur groups, such as horns, frills, crests, sails, and feathers, suggests that visual communication has always been important in dinosaur biology. Reconstruction of the plumage color of Anchiornis, suggest the importance of color in visual communication in non-avian dinosaurs. Vocalization in non-avian dinosaurs is less certain. In birds, the larynx plays no role in sound production. Instead they vocalize with a novel organ called the syrinx, located further down the trachea. The earliest remains of a syrinx was found in a specimen of the duck-like Vegavis iaai dated 69 -66 million years ago, and this organ is unlikely to have existed in non-avian dinosaurs.

Paleontologist Phil Senter has suggested that since non-avian dinosaurs did not have a syrinx, and their next closest living relatives, crocodilians, use the larynx, they could not vocalize as the common ancestor would have been mute. He states that they mostly on visual displays and possibly non-vocal acoustic sounds like hissing, jaw grinding or clapping, splashing and wing beating (possible in winged maniraptoran dinosaurs). Other researchers have countered that vocalizations also exist in turtles, the closest relatives of archosaurs, suggesting that the trait is ancestral to their lineage. In addition, vocal communication in dinosaurs is indicated by the development of advanced hearing in nearly all major groups. Hence the syrinx may have supplemented and then replaced the larynx as a vocal organ rather than there being a "silent period" in bird evolution.

In 2023, a fossilized larynx was described from a specimen of the ankylosaurid Pinacosaurus. The structure was composed of cricoid and arytenoid cartilages, similar to those of non-avian reptiles. However, the mobile cricoid-arytenoid joint and long arytenoid cartilages would have allowed for air-flow control similar to that of birds, and thus could have made bird-like vocalizations. In addition, the cartilages were ossified, implying that laryngeal ossification is a feature of some non-avian dinosaurs. A 2016 study concludes that some dinosaurs may have produced closed mouth vocalizations like cooing, hooting and booming. These occur in both reptiles and birds and involve inflating the esophagus or tracheal pouches. Such vocalizations evolved independently in extant archosaurs numerous times, following increases in body size. The crests of some hadrosaurids and the nasal chambers of ankylosaurids have been suggested to have functioned in acoustic resonance.

Reproductive biology

All dinosaurs laid amniotic eggs. Dinosaur eggs were usually laid in a nest. Most species create somewhat elaborate nests which can be cups, domes, plates, beds scrapes, mounds, or burrows. Some species of modern bird have no nests; the cliff-nesting common guillemot lays its eggs on bare rock, and male emperor penguins keep eggs between their body and feet. Primitive birds and many non-avialan dinosaurs often lay eggs in communal nests, with males primarily incubating the eggs. While modern birds have only one functional oviduct and lay one egg at a time, more primitive birds and dinosaurs had two oviducts, like crocodiles. Some non-avialan dinosaurs, such as , exhibited iterative laying, where the adult might lay a pair of eggs every one or two days, and then ensured simultaneous hatching by delaying brooding until all eggs were laid.

When laying eggs, females grow a special type of bone between the hard outer bone and the marrow of their limbs. This medullary bone, which is rich in calcium, is used to make eggshells. A discovery of features in a Tyrannosaurus skeleton provided evidence of medullary bone in extinct dinosaurs and, for the first time, allowed paleontologists to establish the sex of a fossil dinosaur specimen. Further research has found medullary bone in the carnosaur Allosaurus and the ornithopod Tenontosaurus. Because the line of dinosaurs that includes Allosaurus and Tyrannosaurus diverged from the line that led to Tenontosaurus very early in the evolution of dinosaurs, this suggests that the production of medullary tissue is a general characteristic of all dinosaurs.

Fossil interpreted as a nesting oviraptorid Citipati at the American Museum of Natural History. Smaller fossil far right showing inside one of the eggs.

Another widespread trait among modern birds (but see below in regards to fossil groups and extant megapodes) is parental care for young after hatching. Jack Horner's 1978 discovery of a ("good mother lizard") nesting ground in Montana demonstrated that parental care continued long after birth among ornithopods. A specimen of the oviraptorid Citipati osmolskae was discovered in a chicken-like brooding position in 1993, which may indicate that they had begun using an insulating layer of feathers to keep the eggs warm. An embryo of the basal sauropodomorph Massospondylus was found without teeth, indicating that some parental care was required to feed the young dinosaurs. Trackways have also confirmed parental behavior among ornithopods from the Isle of Skye in northwestern Scotland.

However, there is ample evidence of precociality or superprecociality among many dinosaur species, particularly theropods. For instance, non- ornithuromorph birds have been abundantly demonstrated to have had slow growth rates, megapode-like egg burying behavior and the ability to fly soon after birth. Both Tyrannosaurus and Troodon had juveniles with clear superprecociality and likely occupying different ecological niches than the adults. Superprecociality has been inferred for sauropods.

Genital structures are unlikely to fossilize as they lack scales that may allow preservation via pigmentation or residual calcium phosphate salts. In 2021, the best preserved specimen of a dinosaur's cloacal vent exterior was described for Psittacosaurus, demonstrating lateral swellings similar to crocodylian musk glands used in social displays by both sexes and pigmented regions which could also reflect a signalling function. However, this specimen on its own does not offer enough information to determine whether this dinosaur had sexual signalling functions; it only supports the possibility. Cloacal visual signalling can occur in either males or females in living birds, making it unlikely to be useful to determine sex for extinct dinosaurs.

Physiology

Because both modern crocodilians and birds have four-chambered hearts (albeit modified in crocodilians), it is likely that this is a trait shared by all archosaurs, including all dinosaurs. While all modern birds have high metabolisms and are endothermic ("warm-blooded"), a vigorous debate has been ongoing since the 1960s regarding how far back in the dinosaur lineage this trait extended. Various researchers have supported dinosaurs as being endothermic, ectothermic ("cold-blooded"), or somewhere in between. An emerging consensus among researchers is that, while different lineages of dinosaurs would have had different metabolisms, most of them had higher metabolic rates than other reptiles but lower than living birds and mammals, which is termed mesothermy by some. Evidence from crocodiles and their extinct relatives suggests that such elevated metabolisms could have developed in the earliest archosaurs, which were the common ancestors of dinosaurs and crocodiles.

After non-avian dinosaurs were discovered, paleontologists first posited that they were ectothermic. This was used to imply that the ancient dinosaurs were relatively slow, sluggish organisms, even though many modern reptiles are fast and light-footed despite relying on external sources of heat to regulate their body temperature. The idea of dinosaurs as ectothermic remained a prevalent view until Robert T. Bakker, an early proponent of dinosaur endothermy, published an influential paper on the topic in 1968. Bakker specifically used anatomical and ecological evidence to argue that sauropods, which had hitherto been depicted as sprawling aquatic animals with their tails dragging on the ground, were endotherms that lived vigorous, terrestrial lives. In 1972, Bakker expanded on his arguments based on energy requirements and predator-prey ratios. This was one of the seminal results that led to the dinosaur renaissance.

One of the greatest contributions to the modern understanding of dinosaur physiology has been paleohistology, the study of microscopic tissue structure in dinosaurs. From the 1960s forward, Armand de Ricqlès suggested that the presence of fibrolamellar bone-bony tissue with an irregular, fibrous texture and filled with blood vessels-was indicative of consistently fast growth and therefore endothermy. Fibrolamellar bone was common in both dinosaurs and pterosaurs, though not universally present. This has led to a significant body of work in reconstructing growth curves and modeling the evolution of growth rates across various dinosaur lineages, which has suggested overall that dinosaurs grew faster than living reptiles. Other lines of evidence suggesting endothermy include the presence of feathers and other types of body coverings in many lineages (see § Feathers); more consistent ratios of the isotope oxygen-18 in bony tissue compared to ectotherms, particularly as latitude and thus air temperature varied, which suggests stable internal temperatures (although these ratios can be altered during fossilization); and the discovery of polar dinosaurs, which lived in Australia, Antarctica, and Alaska when these places would have had cool, temperate climates.

Comparison between the air sacs of an abelisaur and a bird

In saurischian dinosaurs, higher metabolisms were supported by the evolution of the avian respiratory system, characterized by an extensive system of air sacs that extended the lungs and invaded many of the bones in the skeleton, making them hollow. Such respiratory systems, which may have appeared in the earliest saurischians, would have provided them with more oxygen compared to a mammal of similar size, while also having a larger resting tidal volume and requiring a lower breathing frequency, which would have allowed them to sustain higher activity levels. The rapid airflow would also have been an effective cooling mechanism, which in conjunction with a lower metabolic rate would have prevented large sauropods from overheating. These traits may have enabled sauropods to grow quickly to gigantic sizes. Sauropods may also have benefitted from their size-their small surface area to volume ratio meant that they would have been able to thermoregulate more easily, a phenomenon termed gigantothermy.

Like other reptiles, dinosaurs are primarily uricotelic, that is, their kidneys extract nitrogenous wastes from their bloodstream and excrete it as uric acid instead of urea or ammonia via the ureters into the intestine. This would have helped them to conserve water. In most living species, uric acid is excreted along with feces as a semisolid waste. However, at least some modern birds (such as hummingbirds) can be facultatively ammonotelic, excreting most of the nitrogenous wastes as ammonia. This material, as well as the output of the intestines, emerges from the cloaca. In addition, many species regurgitate pellets, and fossil pellets are known as early as the Jurassic from Anchiornis.

The size and shape of the brain can be partly reconstructed based on the surrounding bones. In 1896, Marsh calculated ratios between brain weight and body weight of seven species of dinosaurs, showing that the brain of dinosaurs was proportionally smaller than in today's crocodiles, and that the brain of Stegosaurus was smaller than in any living land vertebrate. This contributed to the widespread public notion of dinosaurs as being sluggish and extraordinarily stupid. Harry Jerison, in 1973, showed that proportionally smaller brains are expected at larger body sizes, and that brain size in dinosaurs was not smaller than expected when compared to living reptiles. Later research showed that relative brain size progressively increased during the evolution of theropods, with the highest intelligence - comparable to that of modern birds - calculated for the troodontid Troodon.

Origin of birds

The possibility that dinosaurs were the ancestors of birds was first suggested in 1868 by Thomas Henry Huxley. After the work of Gerhard Heilmann in the early 20th century, the theory of birds as dinosaur descendants was abandoned in favor of the idea of them being descendants of generalized thecodonts, with the key piece of evidence being the supposed lack of clavicles in dinosaurs. However, as later discoveries showed, clavicles (or a single fused wishbone, which derived from separate clavicles) were not actually absent; they had been found as early as 1924 in , but misidentified as an interclavicle. In the 1970s, Ostrom revived the dinosaur-bird theory, which gained momentum in the coming decades with the advent of cladistic analysis, and a great increase in the discovery of small theropods and early birds. Of particular note have been the fossils of the Jehol Biota, where a variety of theropods and early birds have been found, often with feathers of some type. Birds share over a hundred distinct anatomical features with theropod dinosaurs, which are now generally accepted to have been their closest ancient relatives. They are most closely allied with maniraptoran coelurosaurs. A minority of scientists, most notably Alan Feduccia and Larry Martin, have proposed other evolutionary paths, including revised versions of Heilmann's basal archosaur proposal, or that maniraptoran theropods are the ancestors of birds but themselves are not dinosaurs, only convergent with dinosaurs.

Various feathered non-avian dinosaurs, including Archaeopteryx, Anchiornis, Microraptor and Zhenyuanlong

Feathers are one of the most recognizable characteristics of modern birds, and a trait that was also shared by several non-avian dinosaurs. Based on the current distribution of fossil evidence, it appears that feathers were an ancestral dinosaurian trait, though one that may have been selectively lost in some species. Direct fossil evidence of feathers or feather-like structures has been discovered in a diverse array of species in many non-avian dinosaur groups, both among saurischians and ornithischians. Simple, branched, feather-like structures are known from heterodontosaurids, primitive neornithischians, and theropods, and primitive ceratopsians. Evidence for true, vaned feathers similar to the flight feathers of modern birds has been found only in the theropod subgroup Maniraptora, which includes oviraptorosaurs, troodontids, dromaeosaurids, and birds. Feather-like structures known as pycnofibres have also been found in pterosaurs.

However, researchers do not agree regarding whether these structures share a common origin between lineages (i.e., they are homologous), or if they were the result of widespread experimentation with skin coverings among ornithodirans. If the former is the case, filaments may have been common in the ornithodiran lineage and evolved before the appearance of dinosaurs themselves. Research into the genetics of American alligators has revealed that crocodylian scutes do possess feather-keratins during embryonic development, but these keratins are not expressed by the animals before hatching. The description of feathered dinosaurs has not been without controversy in general; perhaps the most vocal critics have been Alan Feduccia and Theagarten Lingham-Soliar, who have proposed that some purported feather-like fossils are the result of the decomposition of collagenous fiber that underlaid the dinosaurs' skin, and that maniraptoran dinosaurs with vaned feathers were not actually dinosaurs, but convergent with dinosaurs. However, their views have for the most part not been accepted by other researchers, to the point that the scientific nature of Feduccia's proposals has been questioned.

Archaeopteryx was the first fossil found that revealed a potential connection between dinosaurs and birds. It is considered a transitional fossil, in that it displays features of both groups. Brought to light just two years after Charles Darwin's seminal On the Origin of Species (1859), its discovery spurred the nascent debate between proponents of evolutionary biology and creationism. This early bird is so dinosaur-like that, without a clear impression of feathers in the surrounding rock, at least one specimen was mistaken for the small theropod Compsognathus. Since the 1990s, a number of additional feathered dinosaurs have been found, providing even stronger evidence of the close relationship between dinosaurs and modern birds. Many of these specimens were unearthed in the lagerstätten of the Jehol Biota. If feather-like structures were indeed widely present among non-avian dinosaurs, the lack of abundant fossil evidence for them may be due to the fact that delicate features like skin and feathers are seldom preserved by fossilization and thus often absent from the fossil record.

Because feathers are often associated with birds, feathered dinosaurs are often touted as the missing link between birds and dinosaurs. However, the multiple skeletal features also shared by the two groups represent another important line of evidence for paleontologists. Areas of the skeleton with important similarities include the neck, pubis, wrist (semi-lunate carpal), arm and pectoral girdle, furcula (wishbone), and breast bone. Comparison of bird and dinosaur skeletons through cladistic analysis strengthens the case for the link.

Soft anatomy

Large meat-eating dinosaurs had a complex system of air sacs similar to those found in modern birds, according to a 2005 investigation led by Patrick M. O'Connor. The lungs of theropod dinosaurs (carnivores that walked on two legs and had bird-like feet) likely pumped air into hollow sacs in their skeletons, as is the case in birds. "What was once formally considered unique to birds was present in some form in the ancestors of birds", O'Connor said. In 2008, scientists described Aerosteon riocoloradensis, the skeleton of which supplies the strongest evidence to date of a dinosaur with a bird-like breathing system. CT scanning of Aerosteon' s fossil bones revealed evidence for the existence of air sacs within the animal's body cavity.

Behavioral evidence

Fossils of the troodonts and Sinornithoides demonstrate that some dinosaurs slept with their heads tucked under their arms. This behavior, which may have helped to keep the head warm, is also characteristic of modern birds. Several deinonychosaur and oviraptorosaur specimens have also been found preserved on top of their nests, likely brooding in a bird-like manner. The ratio between egg volume and body mass of adults among these dinosaurs suggest that the eggs were primarily brooded by the male, and that the young were highly precocial, similar to many modern ground-dwelling birds.

Extinction of major groups

All non-avian dinosaurs and most lineages of birds became extinct in a mass extinction event, called the Cretaceous-Paleogene (K-Pg) extinction event, at the end of the Cretaceous period. Above the Cretaceous-Paleogene boundary, which has been dated to 66.038 ± 0.025 million years ago, fossils of non-avian dinosaurs disappear abruptly; the absence of dinosaur fossils was historically used to assign rocks to the ensuing Cenozoic. The nature of the event that caused this mass extinction has been extensively studied since the 1970s, leading to the development of two mechanisms that are thought to have played major roles: an extraterrestrial impact event in the Yucatán Peninsula, along with flood basalt volcanism in India. However, the specific mechanisms of the extinction event and the extent of its effects on dinosaurs are still areas of ongoing research. Alongside dinosaurs, many other groups of animals became extinct: pterosaurs, marine reptiles such as mosasaurs and plesiosaurs, several groups of mammals, ammonites ( nautilus-like mollusks), rudists ( reef-building bivalves), and various groups of marine plankton. In all, approximately 47% of genera and 76% of species on Earth became extinct during the K-Pg extinction event. The relatively large size of most dinosaurs and the low diversity of small-bodied dinosaur species at the end of the Cretaceous may have contributed to their extinction; the extinction of the bird lineages that did not survive may also have been caused by a dependence on forest habitats or a lack of adaptations to eating seeds for survival.

Pre-extinction diversity

Just before the K-Pg extinction event, the number of non-avian dinosaur species that existed globally has been estimated at between 628 and 1078. It remains uncertain whether the diversity of dinosaurs was in gradual decline before the K-Pg extinction event, or whether dinosaurs were actually thriving prior to the extinction. Rock formations from the Maastrichtian epoch, which directly preceded the extinction, have been found to have lower diversity than the preceding Campanian epoch, which led to the prevailing view of a long-term decline in diversity. However, these comparisons did not account either for varying preservation potential between rock units or for different extents of exploration and excavation. In 1984, Dale Russell carried out an analysis to account for these biases, and found no evidence of a decline; another analysis by David Fastovsky and colleagues in 2004 even showed that dinosaur diversity continually increased until the extinction, but this analysis has been rebutted. Since then, different approaches based on statistics and mathematical models have variously supported either a sudden extinction or a gradual decline. End-Cretaceous trends in diversity may have varied between dinosaur lineages: it has been suggested that sauropods were not in decline, while ornithischians and theropods were in decline.

Impact event

The bolide impact hypothesis, first brought to wide attention in 1980 by Walter Alvarez, Luis Alvarez, and colleagues, attributes the K-Pg extinction event to a bolide (extraterrestrial projectile) impact. Alvarez and colleagues proposed that a sudden increase in iridium levels, recorded around the world in rock deposits at the Cretaceous-Paleogene boundary, was direct evidence of the impact. Shocked quartz, indicative of a strong shockwave emanating from an impact, was also found worldwide. The actual impact site remained elusive until a crater measuring 180 km (110 mi) wide was discovered in the Yucatán Peninsula of southeastern Mexico, and was publicized in a 1991 paper by Alan Hildebrand and colleagues. Now, the bulk of the evidence suggests that a bolide 5 to 15 kilometers (3 to miles) wide impacted the Yucatán Peninsula 66 million years ago, forming this crater and creating a "kill mechanism" that triggered the extinction event.

Within hours, the Chicxulub impact would have created immediate effects such as earthquakes, tsunamis, and a global firestorm that likely killed unsheltered animals and started wildfires. However, it would also have had longer-term consequences for the environment. Within days, sulfate aerosols released from rocks at the impact site would have contributed to acid rain and ocean acidification. Soot aerosols are thought to have spread around the world over the ensuing months and years; they would have cooled the surface of the Earth by reflecting thermal radiation, and greatly slowed photosynthesis by blocking out sunlight, thus creating an impact winter. (This role was ascribed to sulfate aerosols until experiments demonstrated otherwise.) The cessation of photosynthesis would have led to the collapse of food webs depending on leafy plants, which included all dinosaurs save for grain-eating birds.

Deccan Traps

At the time of the K-Pg extinction, the Deccan Traps flood basalts of India were actively erupting. The eruptions can be separated into three phases around the K-Pg boundary, two prior to the boundary and one after. The second phase, which occurred very close to the boundary, would have extruded 70 to 80% of the volume of these eruptions in intermittent pulses that occurred around 100,000 years apart. Greenhouse gases such as carbon dioxide and sulfur dioxide would have been released by this volcanic activity, resulting in climate change through temperature perturbations of roughly 3 °C (5.4 °F) but possibly as high as 7 °C (13 °F). Like the Chicxulub impact, the eruptions may also have released sulfate aerosols, which would have caused acid rain and global cooling. However, due to large error margins in the dating of the eruptions, the role of the Deccan Traps in the K-Pg extinction remains unclear.

Before 2000, arguments that the Deccan Traps eruptions-as opposed to the Chicxulub impact-caused the extinction were usually linked to the view that the extinction was gradual. Prior to the discovery of the Chicxulub crater, the Deccan Traps were used to explain the global iridium layer; even after the crater's discovery, the impact was still thought to only have had a regional, not global, effect on the extinction event. In response, Luis Alvarez rejected volcanic activity as an explanation for the iridium layer and the extinction as a whole. Since then, however, most researchers have adopted a more moderate position, which identifies the Chicxulub impact as the primary progenitor of the extinction while also recognizing that the Deccan Traps may also have played a role. Walter Alvarez himself has acknowledged that the Deccan Traps and other ecological factors may have contributed to the extinctions in addition to the Chicxulub impact. Some estimates have placed the start of the second phase in the Deccan Traps eruptions within 50,000 years after the Chicxulub impact. Combined with mathematical modelling of the seismic waves that would have been generated by the impact, this has led to the suggestion that the Chicxulub impact may have triggered these eruptions by increasing the permeability of the mantle plume underlying the Deccan Traps.

Whether the Deccan Traps were a major cause of the extinction, on par with the Chicxulub impact, remains uncertain. Proponents consider the climatic impact of the sulfur dioxide released to have been on par with the Chicxulub impact, and also note the role of flood basalt volcanism in other mass extinctions like the Permian-Triassic extinction event. They consider the Chicxulub impact to have worsened the ongoing climate change caused by the eruptions. Meanwhile, detractors point out the sudden nature of the extinction and that other pulses in Deccan Traps activity of comparable magnitude did not appear to have caused extinctions. They also contend that the causes of different mass extinctions should be assessed separately. In 2020, Alfio Chiarenza and colleagues suggested that the Deccan Traps may even have had the opposite effect: they suggested that the long-term warming caused by its carbon dioxide emissions may have dampened the impact winter from the Chicxulub impact.

Possible Paleocene survivors

Non-avian dinosaur remains have occasionally been found above the K-Pg boundary. In 2000, Spencer Lucas and colleagues reported the discovery of a single hadrosaur right femur in the San Juan Basin of New Mexico, and described it as evidence of Paleocene dinosaurs. The rock unit in which the bone was discovered has been dated to the early Paleocene epoch, approximately 64.8 million years ago. If the bone was not re-deposited by weathering action, it would provide evidence that some dinosaur populations survived at least half a million years into the Cenozoic. Other evidence includes the presence of dinosaur remains in the Hell Creek Formation up to 1.3 m (4.3 ft) above the Cretaceous-Paleogene boundary, representing 40,000 years of elapsed time. This has been used to support the view that the K-Pg extinction was gradual. However, these supposed Paleocene dinosaurs are considered by many other researchers to be reworked, that is, washed out of their original locations and then re-buried in younger sediments. The age estimates have also been considered unreliable.

Cultural depictions

Gertie the Dinosaur (1914) by Winsor McCay, featuring the first animated dinosaur

Public enthusiasm for dinosaurs first developed in Victorian England, where in 1854, three decades after the first scientific descriptions of dinosaur remains, a menagerie of lifelike dinosaur sculptures was unveiled in London's Crystal Palace Park. The Crystal Palace dinosaurs proved so popular that a strong market in smaller replicas soon developed. In subsequent decades, dinosaur exhibits opened at parks and museums around the world, ensuring that successive generations would be introduced to the animals in an immersive and exciting way. The enduring popularity of dinosaurs, in its turn, has resulted in significant public funding for dinosaur science, and has frequently spurred new discoveries. In the United States, for example, the competition between museums for public attention led directly to the Bone Wars of the 1880s and 1890s, during which a pair of feuding paleontologists made enormous scientific contributions.

The popular preoccupation with dinosaurs has ensured their appearance in literature, film, and other media. Beginning in 1852 with a passing mention in Charles Dickens' Bleak House, dinosaurs have been featured in large numbers of fictional works. Jules Verne's 1864 novel Journey to the Center of the Earth, Sir Arthur Conan Doyle's 1912 book The Lost World, the 1914 animated film Gertie the Dinosaur (featuring the first animated dinosaur), the iconic 1933 film , the 1954 and its many sequels, the best-selling 1990 novel Jurassic Park by Michael Crichton and its 1993 film adaptation are just a few notable examples of dinosaur appearances in fiction. Authors of general-interest non-fiction works about dinosaurs, including some prominent paleontologists, have often sought to use the animals as a way to educate readers about science in general. Dinosaurs are ubiquitous in advertising; numerous companies have referenced dinosaurs in printed or televised advertisements, either in order to sell their own products or in order to characterize their rivals as slow-moving, dim-witted, or obsolete.

Further reading

University of Southampton (September 29, 2021). "Two New Species of Large Predatory Dinosaur With Crocodile-Like Skulls Discovered on Isle of Wight". SciTechDaily.

Zhou, Zhonghe (October 2004). "The origin and early evolution of birds: discoveries, disputes, and perspectives from fossil evidence" (PDF). Naturwissenschaften. Berlin: Springer Science+Business Media. 91 (10): 455-471. Bibcode:2004NW.....91..455Z. doi:10.1007/s00114-004-0570-4. ISSN 0028-1042. PMID 15365634. S2CID 3329625. Archived from the original (PDF) on July 21, 2011.

Paul, Gregory S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore; London: Johns Hopkins University Press. ISBN 978-0-8018-6763-7. LCCN 2001000242. OCLC 1088130487..

Stewart, Tabori & Chang (1997). The Humongous Book of Dinosaurs. New York: Stewart, Tabori & Chang. ISBN 978-1-55670-596-0. LCCN 97000398. OCLC 1037269801.

Sternberg, Charles Mortram (1966) [Original edition published by E. Cloutier, printer to the King, 1946]. Canadian Dinosaurs. Geological Series. Vol. 54 (2nd ed.). Ottawa: National Museum of Canada. LCCN gs46000214. OCLC 1032865683.

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References

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Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is a subject of active research.

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The UNSEEN Dinosaur Killer | Planet Dinosaur | BBC Earth

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Amazing Facts about Mammals - Science With Kids

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Aquarium 4K VIDEO (ULTRA HD) 🐠 Beautiful Relaxing Coral Reef Fish - Relaxing Sleep Meditation Music

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2 Hours of Beautiful Coral Reef Fish, Relaxing Ocean Fish, & Stunning Aquarium Relax Music

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All About Birds for Children: Animal Learning for Kids - FreeSchool

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Learn about Reptiles || Reptiles Video for Kids

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The 1002 most beautiful EXOTIC BIRDS in the world - Photos and curiosities!

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ExoticBirds.life is born to spread true and quality information about the care, breeding, reproduction and feeding of more than 1000 birds and exotic birds of the world.

Types of exotic birds

In this list you will find the main breeds and types of exotic birds. Click on the bird's name or photo for more information.

Images of birds and exotic birds

Welcome to the best list of pictures of the world's prettiest and strangest exotic birds and birds!

Click on the image or its name to see it in high resolution:

Classification of birds by colour

In this list we show you the tropical birds organized according to the color of the birds.

Green birds

Red birds

Yellow birds

The electrical colours of the feathers of these birds usually combine red and green, as in the case of agapornis, but also blue and yellow, as in macaws. And there are little birds, like Gould's diamond, the rainbow lori or the quetzal, which can have between three and six colors.

As in many other species of nature, males are more striking than females, since their colour is one of the factors that make them choose "the best match" in mating season.

However, there are birds in this classification that do not stand out for their colors. Then why are they considered exotic?

Natural habitats of birds

Here we show you some of the places where exotic birds live in an autochthonous way.

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Among the countries that offer more exotic species to the world, those with tropical forests are the first, since they are the place of origin of large numbers of these birds. For example, the turpial is the national bird of Venezuela, where the beautiful cardinal also lives.

The quetzal is found in the jungles of Guatemala, Mexico and Costa Rica, where we can also find more than forty species of hummingbirds and several species of toucans.

But if we talk about jungle, we can't stop talking about Brazil, as it occupies third place in terms of the number of individuals in South America. It has the Amazon rainforest, where, for example, the pendulum bird lives, a species that also lives in other parts of Central America and the Caribbean islands.

The hyacinth macaw has fewer and fewer trees to nest in due to deforestation caused by cattle ranching, fires and the country's hydrographic plans.

Colombia has become a must for birds migrating in the Americas and, with 20% of the world's bird population, it is a paradise for bird lovers.

A curious and endemic species of the place is the red-billed tucancito. And, without going into detail, we can find countless species, with more types of toucans, macaws, parakeets, etc..

Other parts of the world can also be included, as cockatoos, like parakeets, come from Australia. And there are birds that are part of this classification, as well as pigeons, waders, ducks and some hens, whose habitats are spread around the globe.
WHAT IS AN EXOTIC BIRD?
Exotic birds are generally non-native birds, coming from the rainforests of America, Africa and Australia, their natural habitats.

We can also find some species, such as the non-yellow canaries, which are considered exotic because they are not originally found in nature, but have been created through a selective process.

Classification of tropical birds

The different species of tropical birds we find are:

Psittaciformes: macaws, parrots, parakeets, parrots, parrots, parakeets, parakeets, cockatoos and lories.

Passerines: colourful songbirds (goldfinch, finch, canary, hoopoe, oropendola).

Columbiformes: pigeons, including domestic pigeons.

Galliformes: chickens, quails, turkeys or pheasants.

WHEN YOU DECIDE TO HAVE AN EXOTIC BIRD AS A PET
If you want to have one of these birds as a pet, whether it is a small bird or a large bird with slow movement and long life, you must first find out whether it is legal to own it and what papers are needed to ensure that everything is in order and not contribute to the trafficking of exotic animals, some of which are in danger of extinction. That's why it's a good idea to go to a store, because selling between private individuals is illegal.

Then you must learn the care these birds need and choose the right place to put them at home, the temperature of the room, the cage you use, their diet and how to care for them so that they do not get sick.

When you decide to have a parrot as a pet, you have to really love it and know which species to choose and what you can offer it, because they are birds that demand affectionate treatment. And you also have to be aware of the inconvenience before you buy it, so you won't regret it later, because it's very sad to see birds that have been abandoned by their owners in city parks.

You can also have a breeding partner and then you should know what to do if they have chicks.

All this information can be found on exoticbirds.life, but when you go to the store, find out more so that your bird is happy and lives many years with you, since they are birds that live a long time, in some cases like the macaw, as much as people or more.
BIRD BREEDING
If you want to breed, but with more than one partner, you will have to learn what conditions the place where the birds are should be: how much space you need; the temperature you need; what cages to use; how well the light reaches them; the availability of feeders, drinkers and nests; as well as the time of year you breed each species.

Hybridization (the crossing of individuals of different species) is more complicated than breeding and is difficult to achieve success, as the female, for example, may not incubate the eggs or reject the male or the cage, or the chickens may not thrive, etc.
THREATS
Other threats include global warming, the loss of natural habitats through indiscriminate logging, forest fires, illegal hunting for commercialization and breeding, and the introduction of predatory species.

On the other hand, females born from hybridization are sterile and do not sing, so they are not usually wanted by people.

And as for the introduction of non-endemic species, this often poses another threat, since those that settle in the new site unbalance the ecosystem and the native species are displaced, invaded. This is not always the case, but it does happen too often. We can see it now in the case of the Argentine parrot that, in a short period of time, has settled in city parks as if it had always lived in Spain.
LEGISLATION AND CONSERVATION
Because trade in both animal and plant species is a global business, the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) was created in 1973 to ensure that trade is sustainable and does not endanger the survival of species.

Today, almost every country in the world is a party to the CITES Convention.

Thanks to organizations that work for the conservation of animals and plants, it is being achieved that habitats where these birds live are protected, as well as creating sanctuaries where they can live in peace.

It's funny how the SEO defines an exotic bird, "a species that is outside its area, assuming some kind of human intervention that translates into its transfer. It's longer, but it's the same old story: one part of humanity fixing the other part's disasters...

Last Blog Articles

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▷▷ Discover the World's Most Beautiful Exotic Birds 🐦- 【Exotic Birds】

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ExoticBirds.life is born to spread true and quality information about the care, breeding, reproduction and feeding of more than 1000 birds and exotic birds of the world. In this list you will find the main breeds and types of exotic birds. Click on the bird's name or photo for more information.

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Dinosaur | Definition, Types, Pictures, Videos, & Facts

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Dinosaur, the common name given to a group of reptiles, often very large, that first appeared roughly 245 million years ago and thrived worldwide for nearly 180 million years. Most died out by the end of the Cretaceous Period. Many lines of evidence show that one lineage evolved into birds about 155…

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Raccoon

Procyon lotor

Status: Not Listed

Description

A raccoon's face has several markings that help it stand out. The most noticeable marking is the black "mask"-large black markings around each eye. They extend from the edge of the nose to the lower part of the cheek. In addition, raccoons have whitish patches on top of the eyes and around the nose. Raccoons have grayish-brown fur over most of their body, and their tails have four to six black rings.

Range

Raccoons live throughout the continental United States in woods, wetlands, suburbs, parks, cities, and anywhere there is cover, food, and water. Predators of raccoons include the coyote, fisher, bobcat, red fox, and great horned owl.

Diet

Raccoons are omnivores, meaning they will eat both meat and vegetables. They like grasshoppers, nuts, berries, mice, squirrels, and bird eggs. They are nocturnal and search for food at night. Raccoons are opportunistic feeders and are well known by people for their skillful attempts at stealing food from garbage cans in parks and neighborhoods. Raccoons are able to get food that other animals cannot because they have nimble, almost handlike paws that can grasp at tree branches, nuts, fruits, and even the lids of garbage cans.

Life History

Raccoons are solitary, except during the breeding season, which occurs from January to June. Females usually have one litter a year, with three to seven offspring per litter. The gestation period is roughly two months. Young stay with their mother for their first winter, then venture off on their own in spring. A raccoon can live for 16 years in the wild, but often only live for an average of five years.

Fun Fact

At the National Wildlife Federation, we love raccoons, especially our mascot Ranger Rick! But it can be dangerous to get close to raccoons, because unlike Ranger Rick, wild raccoons may become aggressive if provoked.

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Learn facts about the raccoon's habitat, diet, life history, and more.

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Online bird guide, bird ID help, life history, bird sounds from Cornell

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Use our Bird Guide to identify birds, learn about the life history, listen to the sounds, and watch bird behavior on video--the most comprehensive guide to North American birds

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Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species diversity among all orders of organisms. Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every land habitat. As of August 2022, 50,356 spider species in 132 families have been recorded by taxonomists. However, there has been debate among scientists about how families should be classified, with over 20 different classifications proposed since 1900.

Anatomically, spiders (as with all arachnids) differ from other arthropods in that the usual body segments are fused into two tagmata, the cephalothorax or prosoma, and the opisthosoma, or abdomen, and joined by a small, cylindrical pedicel. However, as there is currently neither paleontological nor embryological evidence that spiders ever had a separate thorax-like division, there exists an argument against the validity of the term cephalothorax, which means fused cephalon (head) and the thorax. Similarly, arguments can be formed against use of the term abdomen, as the opisthosoma of all spiders contains a heart and respiratory organs, organs atypical of an abdomen.

Unlike insects, spiders do not have antennae. In all except the most primitive group, the Mesothelae, spiders have the most centralized nervous systems of all arthropods, as all their ganglia are fused into one mass in the cephalothorax. Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by hydraulic pressure.

Their abdomens bear appendages that have been modified into spinnerets that extrude silk from up to six types of glands. Spider webs vary widely in size, shape and the amount of sticky thread used. It now appears that the spiral orb web may be one of the earliest forms, and spiders that produce tangled cobwebs are more abundant and diverse than orb-weaver spiders. Spider-like arachnids with silk-producing spigots ( Uraraneida) appeared in the Devonian period about 386 million years ago, but these animals apparently lacked spinnerets. True spiders have been found in Carboniferous rocks from 318 to 299 million years ago, and are very similar to the most primitive surviving suborder, the Mesothelae. The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appeared in the Triassic period, before 200 million years ago.

The species Bagheera kiplingi was described as herbivorous in 2008, but all other known species are predators, mostly preying on insects and on other spiders, although a few large species also take birds and lizards. It is estimated that the world's 25 million tons of spiders kill 400-800 million tons of prey per year. Spiders use a wide range of strategies to capture prey: trapping it in sticky webs, lassoing it with sticky bolas, mimicking the prey to avoid detection, or running it down. Most detect prey mainly by sensing vibrations, but the active hunters have acute vision, and hunters of the genus show signs of intelligence in their choice of tactics and ability to develop new ones. Spiders' guts are too narrow to take solids, so they liquefy their food by flooding it with digestive enzymes. They also grind food with the bases of their pedipalps, as arachnids do not have the mandibles that crustaceans and insects have.

To avoid being eaten by the females, which are typically much larger, male spiders identify themselves to potential mates by a variety of complex courtship rituals. Males of most species survive a few matings, limited mainly by their short life spans. Females weave silk egg-cases, each of which may contain hundreds of eggs. Females of many species care for their young, for example by carrying them around or by sharing food with them. A minority of species are social, building communal webs that may house anywhere from a few to 50,000 individuals. Social behavior ranges from precarious toleration, as in the widow spiders, to co-operative hunting and food-sharing. Although most spiders live for at most two years, tarantulas and other mygalomorph spiders can live up to 25 years in captivity.

While the venom of a few species is dangerous to humans, scientists are now researching the use of spider venom in medicine and as non-polluting pesticides. Spider silk provides a combination of lightness, strength and elasticity that is superior to that of synthetic materials, and spider silk genes have been inserted into mammals and plants to see if these can be used as silk factories. As a result of their wide range of behaviors, spiders have become common symbols in art and mythology symbolizing various combinations of patience, cruelty and creative powers. An irrational fear of spiders is called arachnophobia.

Description

Spiders are chelicerates and therefore arthropods. As arthropods they have: segmented bodies with jointed limbs, all covered in a cuticle made of chitin and proteins; heads that are composed of several segments that fuse during the development of the embryo. Being chelicerates, their bodies consist of two tagmata, sets of segments that serve similar functions: the foremost one, called the cephalothorax or prosoma, is a complete fusion of the segments that in an insect would form two separate tagmata, the head and thorax; the rear tagma is called the abdomen or opisthosoma. In spiders, the cephalothorax and abdomen are connected by a small cylindrical section, the pedicel. The pattern of segment fusion that forms chelicerates' heads is unique among arthropods, and what would normally be the first head segment disappears at an early stage of development, so that chelicerates lack the antennae typical of most arthropods. In fact, chelicerates' only appendages ahead of the mouth are a pair of chelicerae, and they lack anything that would function directly as "jaws". The first appendages behind the mouth are called pedipalps, and serve different functions within different groups of chelicerates.

Spiders and scorpions are members of one chelicerate group, the arachnids. Scorpions' chelicerae have three sections and are used in feeding. Spiders' chelicerae have two sections and terminate in fangs that are generally venomous, and fold away behind the upper sections while not in use. The upper sections generally have thick "beards" that filter solid lumps out of their food, as spiders can take only liquid food. Scorpions' pedipalps generally form large claws for capturing prey, while those of spiders are fairly small appendages whose bases also act as an extension of the mouth; in addition, those of male spiders have enlarged last sections used for sperm transfer.

In spiders, the cephalothorax and abdomen are joined by a small, cylindrical pedicel, which enables the abdomen to move independently when producing silk. The upper surface of the cephalothorax is covered by a single, convex carapace, while the underside is covered by two rather flat plates. The abdomen is soft and egg-shaped. It shows no sign of segmentation, except that the primitive Mesothelae, whose living members are the Liphistiidae, have segmented plates on the upper surface.

Circulation and respiration

Like other arthropods, spiders are coelomates in which the coelom is reduced to small areas around the reproductive and excretory systems. Its place is largely taken by a hemocoel, a cavity that runs most of the length of the body and through which blood flows. The heart is a tube in the upper part of the body, with a few ostia that act as non-return valves allowing blood to enter the heart from the hemocoel but prevent it from leaving before it reaches the front end. However, in spiders, it occupies only the upper part of the abdomen, and blood is discharged into the hemocoel by one artery that opens at the rear end of the abdomen and by branching arteries that pass through the pedicle and open into several parts of the cephalothorax. Hence spiders have open circulatory systems. The blood of many spiders that have book lungs contains the respiratory pigment hemocyanin to make oxygen transport more efficient.

Spiders have developed several different respiratory anatomies, based on book lungs, a tracheal system, or both. Mygalomorph and Mesothelae spiders have two pairs of book lungs filled with haemolymph, where openings on the ventral surface of the abdomen allow air to enter and diffuse oxygen. This is also the case for some basal araneomorph spiders, like the family Hypochilidae, but the remaining members of this group have just the anterior pair of book lungs intact while the posterior pair of breathing organs are partly or fully modified into tracheae, through which oxygen is diffused into the haemolymph or directly to the tissue and organs. The tracheal system has most likely evolved in small ancestors to help resist desiccation. The trachea were originally connected to the surroundings through a pair of openings called spiracles, but in the majority of spiders this pair of spiracles has fused into a single one in the middle, and moved backwards close to the spinnerets. Spiders that have tracheae generally have higher metabolic rates and better water conservation. Spiders are ectotherms, so environmental temperatures affect their activity.

Feeding, digestion and excretion

Uniquely among chelicerates, the final sections of spiders' chelicerae are fangs, and the great majority of spiders can use them to inject venom into prey from venom glands in the roots of the chelicerae. The families Uloboridae and Holarchaeidae, and some Liphistiidae spiders, have lost their venom glands, and kill their prey with silk instead. Like most arachnids, including scorpions, spiders have a narrow gut that can only cope with liquid food and two sets of filters to keep solids out. They use one of two different systems of external digestion. Some pump digestive enzymes from the midgut into the prey and then suck the liquified tissues of the prey into the gut, eventually leaving behind the empty husk of the prey. Others grind the prey to pulp using the chelicerae and the bases of the pedipalps, while flooding it with enzymes; in these species, the chelicerae and the bases of the pedipalps form a preoral cavity that holds the food they are processing.

Most spiders convert nitrogenous waste products into uric acid, which can be excreted as a dry material. Malphigian tubules ("little tubes") extract these wastes from the blood in the hemocoel and dump them into the cloacal chamber, from which they are expelled through the anus. Production of uric acid and its removal via Malphigian tubules are a water-conserving feature that has evolved independently in several arthropod lineages that can live far away from water, for example the tubules of insects and arachnids develop from completely different parts of the embryo. However, a few primitive spiders, the suborder Mesothelae and infraorder Mygalomorphae, retain the ancestral arthropod nephridia ("little kidneys"), which use large amounts of water to excrete nitrogenous waste products as ammonia.

Central nervous system

The basic arthropod central nervous system consists of a pair of nerve cords running below the gut, with paired ganglia as local control centers in all segments; a brain formed by fusion of the ganglia for the head segments ahead of and behind the mouth, so that the esophagus is encircled by this conglomeration of ganglia. Except for the primitive Mesothelae, of which the Liphistiidae are the sole surviving family, spiders have the much more centralized nervous system that is typical of arachnids: all the ganglia of all segments behind the esophagus are fused, so that the cephalothorax is largely filled with nervous tissue and there are no ganglia in the abdomen; in the Mesothelae, the ganglia of the abdomen and the rear part of the cephalothorax remain unfused.

Despite the relatively small central nervous system, some spiders (like ) exhibit complex behaviour, including the ability to use a trial-and-error approach.

Sense organs

This jumping spider's main ocelli (center pair) are very acute. The outer pair are "secondary eyes" and there are other pairs of secondary eyes on the sides and top of its head.[25]

Spiders have primarily four pairs of eyes on the top-front area of the cephalothorax, arranged in patterns that vary from one family to another. The principal pair at the front are of the type called pigment-cup ocelli ("little eyes"), which in most arthropods are only capable of detecting the direction from which light is coming, using the shadow cast by the walls of the cup. However, in spiders these eyes are capable of forming images. The other pairs, called secondary eyes, are thought to be derived from the compound eyes of the ancestral chelicerates, but no longer have the separate facets typical of compound eyes. Unlike the principal eyes, in many spiders these secondary eyes detect light reflected from a reflective tapetum lucidum, and wolf spiders can be spotted by torchlight reflected from the tapeta. On the other hand, the secondary eyes of jumping spiders have no tapeta.

The visual acuity of some jumping spiders exceeds by a factor of ten that of dragonflies, which have by far the best vision among insects.[citation needed ] This acuity is achieved by a telephotographic series of lenses, a four-layer retina, and the ability to swivel the eyes and integrate images from different stages in the scan.[citation needed ] The downside is that the scanning and integrating processes are relatively slow.

There are spiders with a reduced number of eyes, the most common having six eyes (example, Periegops suterii) with a pair of eyes absent on the anterior median line. Other species have four eyes and members of the Caponiidae family can have as few as two. Cave dwelling species have no eyes, or possess vestigial eyes incapable of sight.[citation needed ]

Other senses

As with other arthropods, spiders' cuticles would block out information about the outside world, except that they are penetrated by many sensors or connections from sensors to the nervous system. In fact, spiders and other arthropods have modified their cuticles into elaborate arrays of sensors. Various touch sensors, mostly bristles called setae, respond to different levels of force, from strong contact to very weak air currents. Chemical sensors provide equivalents of taste and smell, often by means of setae. An adult may have up to 1,000 such chemosensitive setae, most on the tarsi of the first pair of legs. Males have more chemosensitive bristles on their pedipalps than females. They have been shown to be responsive to sex pheromones produced by females, both contact and air-borne. The jumping spider Evarcha culicivora uses the scent of blood from mammals and other vertebrates, which is obtained by capturing blood-filled mosquitoes, to attract the opposite sex. Because they are able to tell the sexes apart, it is assumed the blood scent is mixed with pheromones. Spiders also have in the joints of their limbs slit sensillae that detect force and vibrations. In web-building spiders, all these mechanical and chemical sensors are more important than the eyes, while the eyes are most important to spiders that hunt actively.

Like most arthropods, spiders lack balance and acceleration sensors and rely on their eyes to tell them which way is up. Arthropods' proprioceptors, sensors that report the force exerted by muscles and the degree of bending in the body and joints, are well-understood. On the other hand, little is known about what other internal sensors spiders or other arthropods may have.

Some spiders use their webs for hearing, where the giant webs function as extended and reconfigurable auditory sensors.

Locomotion

Each of the eight legs of a spider consists of seven distinct parts. The part closest to and attaching the leg to the cephalothorax is the coxa; the next segment is the short trochanter that works as a hinge for the following long segment, the femur; next is the spider's knee, the patella, which acts as the hinge for the tibia; the metatarsus is next, and it connects the tibia to the tarsus (which may be thought of as a foot of sorts); the tarsus ends in a claw made up of either two or three points, depending on the family to which the spider belongs. Although all arthropods use muscles attached to the inside of the exoskeleton to flex their limbs, spiders and a few other groups still use hydraulic pressure to extend them, a system inherited from their pre-arthropod ancestors. The only extensor muscles in spider legs are located in the three hip joints (bordering the coxa and the trochanter). As a result, a spider with a punctured cephalothorax cannot extend its legs, and the legs of dead spiders curl up. Spiders can generate pressures up to eight times their resting level to extend their legs, and jumping spiders can jump up to 50 times their own length by suddenly increasing the blood pressure in the third or fourth pair of legs. Although larger spiders use hydraulics to straighten their legs, unlike smaller jumping spiders they depend on their flexor muscles to generate the propulsive force for their jumps.

Most spiders that hunt actively, rather than relying on webs, have dense tufts of fine bristles between the paired claws at the tips of their legs. These tufts, known as scopulae, consist of bristles whose ends are split into as many as 1,000 branches, and enable spiders with scopulae to walk up vertical glass and upside down on ceilings. It appears that scopulae get their grip from contact with extremely thin layers of water on surfaces. Spiders, like most other arachnids, keep at least four legs on the surface while walking or running.

Silk production

The abdomen has no appendages except those that have been modified to form one to four (usually three) pairs of short, movable spinnerets, which emit silk. Each spinneret has many spigots, each of which is connected to one silk gland. There are at least six types of silk gland, each producing a different type of silk. Spitting spiders also produce silk in modified venom glands.

Silk is mainly composed of a protein very similar to that used in insect silk. It is initially a liquid, and hardens not by exposure to air but as a result of being drawn out, which changes the internal structure of the protein. It is similar in tensile strength to nylon and biological materials such as chitin, collagen and cellulose, but is much more elastic. In other words, it can stretch much further before breaking or losing shape.

Some spiders have a cribellum, a modified spinneret with up to 40,000 spigots, each of which produces a single very fine fiber. The fibers are pulled out by the calamistrum, a comblike set of bristles on the jointed tip of the cribellum, and combined into a composite woolly thread that is very effective in snagging the bristles of insects. The earliest spiders had cribella, which produced the first silk capable of capturing insects, before spiders developed silk coated with sticky droplets. However, most modern groups of spiders have lost the cribellum.

Even species that do not build webs to catch prey use silk in several ways: as wrappers for sperm and for fertilized eggs; as a " safety rope"; for nest-building; and as " parachutes" by the young of some species.

Reproduction and life cycle

Spiders reproduce sexually and fertilization is internal but indirect, in other words the sperm is not inserted into the female's body by the male's genitals but by an intermediate stage. Unlike many land-living arthropods, male spiders do not produce ready-made spermatophores (packages of sperm), but spin small sperm webs onto which they ejaculate and then transfer the sperm to special syringe-styled structures, palpal bulbs or palpal organs, borne on the tips of the pedipalps of mature males. When a male detects signs of a female nearby he checks whether she is of the same species and whether she is ready to mate; for example in species that produce webs or "safety ropes", the male can identify the species and sex of these objects by "smell".

Spiders generally use elaborate courtship rituals to prevent the large females from eating the small males before fertilization, except where the male is so much smaller that he is not worth eating. In web-weaving species, precise patterns of vibrations in the web are a major part of the rituals, while patterns of touches on the female's body are important in many spiders that hunt actively, and may "hypnotize" the female. Gestures and dances by the male are important for jumping spiders, which have excellent eyesight. If courtship is successful, the male injects his sperm from the palpal bulbs into the female via one or two openings on the underside of her abdomen.

Female spiders' reproductive tracts are arranged in one of two ways. The ancestral arrangement ("haplogyne" or "non-entelegyne") consists of a single genital opening, leading to two seminal receptacles (spermathecae) in which females store sperm. In the more advanced arrangement ("entelegyne"), there are two further openings leading directly to the spermathecae, creating a "flow through" system rather than a "first-in first-out" one. Eggs are as a general rule only fertilized during oviposition when the stored sperm is released from its chamber, rather than in the ovarian cavity. A few exceptions exist, such as Parasteatoda tepidariorum. In these species the female appears to be able to activate the dormant sperm before oviposition, allowing them to migrate to the ovarian cavity where fertilization occurs. The only known example of direct fertilization between male and female is an Israeli spider, Harpactea sadistica, which has evolved traumatic insemination. In this species the male will penetrate its pedipalps through the female's body wall and inject his sperm directly into her ovaries, where the embryos inside the fertilized eggs will start to develop before being laid.

Males of the genus amputate one of their palps before maturation and enter adult life with one palp only. The palps are 20% of the male's body mass in this species, and detaching one of the two improves mobility. In the Yemeni species Tidarren argo, the remaining palp is then torn off by the female. The separated palp remains attached to the female's epigynum for about four hours and apparently continues to function independently. In the meantime, the female feeds on the palpless male. In over 60% of cases, the female of the Australian redback spider kills and eats the male after it inserts its second palp into the female's genital opening; in fact, the males co-operate by trying to impale themselves on the females' fangs. Observation shows that most male redbacks never get an opportunity to mate, and the "lucky" ones increase the likely number of offspring by ensuring that the females are well-fed. However, males of most species survive a few matings, limited mainly by their short life spans. Some even live for a while in their mates' webs.

The tiny male of the golden orb weaver (Trichonephila clavipes) (near the top of the leaf) is protected from the female by producing the right vibrations in the web, and may be too small to be worth eating.

Orange spider egg sac hanging from ceiling

Gasteracantha mammosa spiderlings next to their eggs capsule

Wolf spider carrying its young on its abdomen

Females lay up to 3,000 eggs in one or more silk egg sacs, which maintain a fairly constant humidity level. In some species, the females die afterwards, but females of other species protect the sacs by attaching them to their webs, hiding them in nests, carrying them in the chelicerae or attaching them to the spinnerets and dragging them along.

Baby spiders pass all their larval stages inside the egg sac and emerge as spiderlings, very small and sexually immature but similar in shape to adults. Some spiders care for their young, for example a wolf spider's brood clings to rough bristles on the mother's back, and females of some species respond to the "begging" behaviour of their young by giving them their prey, provided it is no longer struggling, or even regurgitate food.

Like other arthropods, spiders have to molt to grow as their cuticle ("skin") cannot stretch. In some species males mate with newly-molted females, which are too weak to be dangerous to the males. Most spiders live for only one to two years, although some tarantulas can live in captivity for over 20 years, and an Australian female trapdoor spider was documented to have lived in the wild for 43 years, dying of a parasitic wasp attack.

Goliath birdeater (Theraphosa blondi), the largest spider by mass

Spiders occur in a large range of sizes. The smallest, from Colombia, are less than 0.37 mm (0.015 in) in body length. The largest and heaviest spiders occur among tarantulas, which can have body lengths up to 90 mm (3.5 in) and leg spans up to 250 mm (9.8 in).

Coloration

Only three classes of pigment ( ommochromes, bilins and guanine) have been identified in spiders, although other pigments have been detected but not yet characterized. Melanins, carotenoids and pterins, very common in other animals, are apparently absent. In some species, the exocuticle of the legs and prosoma is modified by a tanning process, resulting in a brown coloration. Bilins are found, for example, inMicrommata virescens, resulting in its green color. Guanine is responsible for the white markings of the European garden spider Araneus diadematus. It is in many species accumulated in specialized cells called guanocytes. In genera such as Tetragnatha, , or Theridiosoma, guanine creates their silvery appearance. While guanine is originally an end-product of protein metabolism, its excretion can be blocked in spiders, leading to an increase in its storage. Structural colors occur in some species, which are the result of the diffraction, scattering or interference of light, for example by modified setae or scales. The white prosoma of results from bristles reflecting the light, and both have areas of modified cuticle that act as light reflectors. The peacock spiders of Australia (genus ) are notable for their bright structural colours in the males.

While in many spiders color is fixed throughout their lifespan, in some groups, color may be variable in response to environmental and internal conditions. Choice of prey may be able to alter the color of spiders. For example, the abdomen of Theridion grallator will become orange if the spider ingests certain species of Diptera and adult Lepidoptera, but if it consumes Homoptera or larval Lepidoptera, then the abdomen becomes green. Environmentally induced color changes may be morphological (occurring over several days) or physiological (occurring near instantly). Morphological changes require pigment synthesis and degradation. In contrast to this, physiological changes occur by changing the position of pigment-containing cells. An example of morphological color changes is background matching. Misumena vatia for instance can change its body color to match the substrate it lives on which makes it more difficult to be detected by prey. An example of physiological color change is observed in Cyrtophora cicatrosa, which can change its body color from white to brown near instantly.

Ecology and behavior

Non-predatory feeding

A jumping spider seen in Chennai.

Although spiders are generally regarded as predatory, the jumping spider Bagheera kiplingi gets over 90% of its food from fairly solid plant material produced by acacias as part of a mutually beneficial relationship with a species of ant.

Juveniles of some spiders in the families Anyphaenidae, Corinnidae, Clubionidae, Thomisidae and Salticidae feed on plant nectar. Laboratory studies show that they do so deliberately and over extended periods, and periodically clean themselves while feeding. These spiders also prefer sugar solutions to plain water, which indicates that they are seeking nutrients. Since many spiders are nocturnal, the extent of nectar consumption by spiders may have been underestimated. Nectar contains amino acids, lipids, vitamins and minerals in addition to sugars, and studies have shown that other spider species live longer when nectar is available. Feeding on nectar avoids the risks of struggles with prey, and the costs of producing venom and digestive enzymes.

Various species are known to feed on dead arthropods (scavenging), web silk, and their own shed exoskeletons. Pollen caught in webs may also be eaten, and studies have shown that young spiders have a better chance of survival if they have the opportunity to eat pollen. In captivity, several spider species are also known to feed on bananas, marmalade, milk, egg yolk and sausages.

Capturing prey

The Phonognatha graeffei or leaf-curling spider's web serves both as a trap and as a way of making its home in a leaf.

The best-known method of prey capture is by means of sticky webs. Varying placement of webs allows different species of spider to trap different insects in the same area, for example flat horizontal webs trap insects that fly up from vegetation underneath while flat vertical webs trap insects in horizontal flight. Web-building spiders have poor vision, but are extremely sensitive to vibrations.

Females of the water spider Argyroneta aquatica build underwater "diving bell" webs that they fill with air and use for digesting prey, molting, mating and raising offspring. They live almost entirely within the bells, darting out to catch prey animals that touch the bell or the threads that anchor it. A few spiders use the surfaces of lakes and ponds as "webs", detecting trapped insects by the vibrations that these cause while struggling.

Net-casting spiders weave only small webs, but then manipulate them to trap prey. Those of the genus and the family Theridiosomatidae stretch their webs and then release them when prey strike them, but do not actively move their webs. Those of the family Deinopidae weave even smaller webs, hold them outstretched between their first two pairs of legs, and lunge and push the webs as much as twice their own body length to trap prey, and this move may increase the webs' area by a factor of up to ten. Experiments have shown that Deinopis spinosus has two different techniques for trapping prey: backwards strikes to catch flying insects, whose vibrations it detects; and forward strikes to catch ground-walking prey that it sees. These two techniques have also been observed in other deinopids. Walking insects form most of the prey of most deinopids, but one population of Deinopis subrufa appears to live mainly on tipulid flies that they catch with the backwards strike.

Mature female bolas spiders of the genus Mastophora build "webs" that consist of only a single "trapeze line", which they patrol. They also construct a bolas made of a single thread, tipped with a large ball of very wet sticky silk. They emit chemicals that resemble the pheromones of moths, and then swing the bolas at the moths. Although they miss on about 50% of strikes, they catch about the same weight of insects per night as web-weaving spiders of similar size. The spiders eat the bolas if they have not made a kill in about 30 minutes, rest for a while, and then make new bolas. Juveniles and adult males are much smaller and do not make bolas. Instead they release different pheromones that attract moth flies, and catch them with their front pairs of legs.

A trapdoor spider in the genus Cyclocosmia, an ambush predator

The primitive Liphistiidae, the "trapdoor spiders" of the family Ctenizidae and many tarantulas are ambush predators that lurk in burrows, often closed by trapdoors and often surrounded by networks of silk threads that alert these spiders to the presence of prey. Other ambush predators do without such aids, including many crab spiders, and a few species that prey on bees, which see ultraviolet, can adjust their ultraviolet reflectance to match the flowers in which they are lurking. Wolf spiders, jumping spiders, fishing spiders and some crab spiders capture prey by chasing it, and rely mainly on vision to locate prey.

Portia uses both webs and cunning, versatile tactics to overcome prey.[62]

Some jumping spiders of the genus hunt other spiders in ways that seem intelligent, outflanking their victims or luring them from their webs. Laboratory studies show that Portia's instinctive tactics are only starting points for a trial-and-error approach from which these spiders learn very quickly how to overcome new prey species. However, they seem to be relatively slow "thinkers", which is not surprising, as their brains are vastly smaller than those of mammalian predators.

An ant-mimicking jumping spider

Ant-mimicking spiders face several challenges: they generally develop slimmer abdomens and false "waists" in the cephalothorax to mimic the three distinct regions (tagmata) of an ant's body; they wave the first pair of legs in front of their heads to mimic antennae, which spiders lack, and to conceal the fact that they have eight legs rather than six; they develop large color patches round one pair of eyes to disguise the fact that they generally have eight simple eyes, while ants have two compound eyes; they cover their bodies with reflective bristles to resemble the shiny bodies of ants. In some spider species, males and females mimic different ant species, as female spiders are usually much larger than males. Ant-mimicking spiders also modify their behavior to resemble that of the target species of ant; for example, many adopt a zig-zag pattern of movement, ant-mimicking jumping spiders avoid jumping, and spiders of the genus Synemosyna walk on the outer edges of leaves in the same way as Pseudomyrmex. Ant mimicry in many spiders and other arthropods may be for protection from predators that hunt by sight, including birds, lizards and spiders. However, several ant-mimicking spiders prey either on ants or on the ants' " livestock", such as aphids. When at rest, the ant-mimicking crab spider does not closely resemble Oecophylla, but while hunting it imitates the behavior of a dying ant to attract worker ants. After a kill, some ant-mimicking spiders hold their victims between themselves and large groups of ants to avoid being attacked.

Threat display by a Sydney funnel-web spider (Atrax robustus).

There is strong evidence that spiders' coloration is camouflage that helps them to evade their major predators, birds and parasitic wasps, both of which have good color vision. Many spider species are colored so as to merge with their most common backgrounds, and some have disruptive coloration, stripes and blotches that break up their outlines. In a few species, such as the Hawaiian happy-face spider, Theridion grallator, several coloration schemes are present in a ratio that appears to remain constant, and this may make it more difficult for predators to recognize the species. Most spiders are insufficiently dangerous or unpleasant-tasting for warning coloration to offer much benefit. However, a few species with powerful venom, large jaws or irritant bristles have patches of warning colors, and some actively display these colors when threatened.

Many of the family Theraphosidae, which includes tarantulas and baboon spiders, have urticating hairs on their abdomens and use their legs to flick them at attackers. These bristles are fine setae (bristles) with fragile bases and a row of barbs on the tip. The barbs cause intense irritation but there is no evidence that they carry any kind of venom. A few defend themselves against wasps by including networks of very robust threads in their webs, giving the spider time to flee while the wasps are struggling with the obstacles. The golden wheeling spider, Carparachne aureoflava, of the Namibian desert escapes parasitic wasps by flipping onto its side and cartwheeling down sand dunes.

Socialization

A few spider species that build webs live together in large colonies and show social behavior, although not as complex as in social insects. (in the family Theridiidae) can form colonies of up to 50,000 individuals. The genus Anelosimus has a strong tendency towards sociality: all known American species are social, and species in Madagascar are at least somewhat social. Members of other species in the same family but several different genera have independently developed social behavior. For example, although Theridion nigroannulatum belongs to a genus with no other social species, T. nigroannulatum build colonies that may contain several thousand individuals that co-operate in prey capture and share food. Other communal spiders include several Philoponella species (family Uloboridae), Agelena consociata (family Agelenidae) and (family Dictynidae). Social predatory spiders need to defend their prey against kleptoparasites ("thieves"), and larger colonies are more successful in this. The herbivorous spider Bagheera kiplingi lives in small colonies which help to protect eggs and spiderlings. Even widow spiders (genus Latrodectus), which are notoriously cannibalistic, have formed small colonies in captivity, sharing webs and feeding together.

In experiments, spider species like Steatoda grossa, Latrodectus hesperus and Eratigena agrestis stayed away from Myrmica rubra ant colonies. These ants are predators and the pheromones they release for communication have a notable deterrent effect on these spider species.

There is no consistent relationship between the classification of spiders and the types of web they build: species in the same genus may build very similar or significantly different webs. Nor is there much correspondence between spiders' classification and the chemical composition of their silks. Convergent evolution in web construction, in other words use of similar techniques by remotely related species, is rampant. Orb web designs and the spinning behaviors that produce them are the best understood. The basic radial-then-spiral sequence visible in orb webs and the sense of direction required to build them may have been inherited from the common ancestors of most spider groups. However, the majority of spiders build non-orb webs. It used to be thought that the sticky orb web was an evolutionary innovation resulting in the diversification of the Orbiculariae. Now, however, it appears that non-orb spiders are a subgroup that evolved from orb-web spiders, and non-orb spiders have over 40% more species and are four times as abundant as orb-web spiders. Their greater success may be because sphecid wasps, which are often the dominant predators of spiders, much prefer to attack spiders that have flat webs.

The hubs of orb webs, where the spiders lurk, are usually above the center, as the spiders can move downwards faster than upwards. If there is an obvious direction in which the spider can retreat to avoid its own predators, the hub is usually offset towards that direction.

Spiders often attach highly visible silk bands, called decorations or stabilimenta, to their webs. Field research suggests that webs with more decorative bands captured more prey per hour. However, a laboratory study showed that spiders reduce the building of these decorations if they sense the presence of predators.

There are several unusual variants of orb web, many of them convergently evolved, including: attachment of lines to the surface of water, possibly to trap insects in or on the surface; webs with twigs through their centers, possibly to hide the spiders from predators; "ladderlike" webs that appear most effective in catching moths. However, the significance of many variations is unclear. The orb-weaving species, Zygiella x-notata, for example, is known for its characteristic missing sector orb web. The missing sector contains a signal thread used to detect prey vibrations on the female's web.

In 1973, Skylab 3 took two orb-web spiders into space to test their web-spinning capabilities in zero gravity. At first, both produced rather sloppy webs, but they adapted quickly.

Web design in zero gravity

Many experiments have been conducted to study the effect of zero gravity on the design of spider webs. In late 2020, reports of recent experiments were published that indicated that although web design was affected adversely in zero gravity conditions, having access to a light source could orient spiders and enable them to build their normally shaped webs under such conditions.

Fossil record

Spider preserved in amber

Although the fossil record of spiders is considered poor, almost 1000 species have been described from fossils. Because spiders' bodies are quite soft, the vast majority of fossil spiders have been found preserved in amber. The oldest known amber that contains fossil arthropods dates from 130 million years ago in the Early Cretaceous period. In addition to preserving spiders' anatomy in very fine detail, pieces of amber show spiders mating, killing prey, producing silk and possibly caring for their young. In a few cases, amber has preserved spiders' egg sacs and webs, occasionally with prey attached; the oldest fossil web found so far is 100 million years old. Earlier spider fossils come from a few lagerstätten, places where conditions were exceptionally suited to preserving fairly soft tissues.

The oldest known exclusively terrestrial arachnid is the trigonotarbid Palaeotarbus jerami, from about 420 million years ago in the Silurian period, and had a triangular cephalothorax and segmented abdomen, as well as eight legs and a pair of pedipalps. Attercopus fimbriunguis, from 386 million years ago in the Devonian period, bears the earliest known silk-producing spigots, and was therefore hailed as a spider at the time of its discovery. However, these spigots may have been mounted on the underside of the abdomen rather than on spinnerets, which are modified appendages and whose mobility is important in the building of webs. Hence Attercopus and the similar Permian arachnid Permarachne may not have been true spiders, and probably used silk for lining nests or producing egg cases rather than for building webs. The largest known fossil spider as of 2011 is the araneid Nephila jurassica, from about 165 million years ago, recorded from Daohuogo, Inner Mongolia in China. Its body length is almost 25 mm, (i.e., almost one inch).

Several Carboniferous spiders were members of the Mesothelae, a primitive group now represented only by the Liphistiidae. The mesothelidPaleothele montceauensis, from the Late Carboniferous over 299 million years ago, had five spinnerets. Although the Permian period 299 to 251 million years ago saw rapid diversification of flying insects, there are very few fossil spiders from this period.

The main groups of modern spiders, Mygalomorphae and Araneomorphae, first appear in the Triassic well before 200 million years ago. Some Triassic mygalomorphs appear to be members of the family Hexathelidae, whose modern members include the notorious Sydney funnel-web spider, and their spinnerets appear adapted for building funnel-shaped webs to catch jumping insects. Araneomorphae account for the great majority of modern spiders, including those that weave the familiar orb-shaped webs. The Jurassic and Cretaceous periods provide a large number of fossil spiders, including representatives of many modern families.

According to a 2020 study using a molecular clock calibrated with 27 chelicerate fossils, spiders most likely diverged from other chelicerates between 375 and 328 million years ago.

External relationships

The spiders (Araneae) are monophyletic (i.e., a clade, consisting of a last common ancestor and all of its descendants). There has been debate about what their closest evolutionary relatives are, and how all of these evolved from the ancestral chelicerates, which were marine animals. This 2019 cladogram illustrates the spiders' phylogenetic relationships.

Arachnids lack some features of other chelicerates, including backward-pointing mouths and gnathobases ("jaw bases") at the bases of their legs; both of these features are part of the ancestral arthropod feeding system. Instead, they have mouths that point forwards and downwards, and all have some means of breathing air. Spiders (Araneae) are distinguished from other arachnid groups by several characteristics, including spinnerets and, in males, pedipalps that are specially adapted for sperm transfer.

Internal relationships

Spiders are divided into two suborders, Mesothelae and Opisthothelae, of which the latter contains two infraorders, Mygalomorphae and Araneomorphae. Some 50,356 living species of spiders (order Araneae) have been identified, grouped into 132 families and 4,280 genera by arachnologists in 2022.

Mesothelae

Ryuthela sasakii, a member of the Liphistiidae[102]

The only living members of the primitive Mesothelae are the family Liphistiidae, found only in Southeast Asia, China, and Japan. Most of the Liphistiidae construct silk-lined burrows with thin trapdoors, although some species of the genus Liphistius build camouflaged silk tubes with a second trapdoor as an emergency exit. Members of the genus Liphistius run silk " tripwires" outwards from their tunnels to help them detect approaching prey, while those of the genus Heptathela do not and instead rely on their built-in vibration sensors. Spiders of the genus Heptathela have no venom glands, although they do have venom gland outlets on the fang tip.

The extinct families Arthrolycosidae, found in Carboniferous and Permian rocks, and Arthromygalidae, so far found only in Carboniferous rocks, have been classified as members of the Mesothelae.

Mygalomorphae

The Mygalomorphae, which first appeared in the Triassic period, are generally heavily built and ″hairy″, with large, robust chelicerae and fangs (technically, spiders do not have true hairs, but rather setae). Well-known examples include tarantulas, ctenizid trapdoor spiders and the Australasian funnel-web spiders. Most spend the majority of their time in burrows, and some run silk tripwires out from these, but a few build webs to capture prey. However, mygalomorphs cannot produce the pirifom silk that the Araneomorphae use as an instant adhesive to glue silk to surfaces or to other strands of silk, and this makes web construction more difficult for mygalomorphs. Since mygalomorphs rarely " balloon" by using air currents for transport, their populations often form clumps. In addition to arthropods, some mygalomorphs are known to prey on frogs, small mammals, lizards, snakes, snails, and small birds.

Araneomorphae

Leucauge venusta, an orb-web spider

In addition to accounting for over 90% of spider species, the Araneomorphae, also known as the "true spiders", include orb-web spiders, the cursorial wolf spiders, and jumping spiders, as well as the only known herbivorous spider, Bagheera kiplingi. They are distinguished by having fangs that oppose each other and cross in a pinching action, in contrast to the Mygalomorphae, which have fangs that are nearly parallel in alignment.

Human interaction

Media coverage and misconceptions

Information about spiders in the media is often emphasizing how dangerous and unpleasant they are. A recent study investigated the global spread of (mis-)information on spiders using a high-resolution global database of online newspaper articles on spider-human interactions. These reports covered stories of spider-human encounters and bites published from 2010 to 2020. The study found that 47% of articles contained errors and 43% were sensationalist.

Although spiders are widely feared, only a few species are dangerous to people. Spiders will only bite humans in self-defense, and few produce worse effects than a mosquito bite or bee sting. Most of those with medically serious bites, such as recluse spiders (genus Loxosceles) and widow spiders (genus Latrodectus), would rather flee and bite only when trapped, although this can easily arise by accident. The defensive tactics of Australian funnel-web spiders (family Atracidae) include fang display. Their venom, although they rarely inject much, has resulted in 13 attributed human deaths over 50 years. They have been deemed to be the world's most dangerous spiders on clinical and venom toxicity grounds, though this claim has also been attributed to the Brazilian wandering spider (genus Phoneutria).

There were about 100 reliably reported deaths from spider bites in the 20th century, compared to about 1,500 from jellyfish stings. Many alleged cases of spider bites may represent incorrect diagnoses, which would make it more difficult to check the effectiveness of treatments for genuine bites. A review published in 2016 agreed with this conclusion, showing that 78% of 134 published medical case studies of supposed spider bites did not meet the necessary criteria for a spider bite to be verified. In the case of the two genera with the highest reported number of bites, Loxosceles and Latrodectus, spider bites were not verified in over 90% of the reports. Even when verification had occurred, details of the treatment and its effects were often lacking.

Because spider silk is both light and very strong, attempts are being made to produce it in goats' milk and in the leaves of plants, by means of genetic engineering.

Arachnophobia

Arachnophobia is a specific phobia-it is the abnormal fear of spiders or anything reminiscent of spiders, such as webs or spiderlike shapes. It is one of the most common specific phobias, and some statistics show that 50% of women and 10% of men show symptoms. It may be an exaggerated form of an instinctive response that helped early humans to survive, or a cultural phenomenon that is most common in predominantly European societies.

Cooked tarantulas are considered a delicacy in Cambodia.

Spiders are used as food. Cooked tarantulas are considered a delicacy in Cambodia, and by the Piaroa Indians of southern Venezuela - provided the highly irritant bristles, the spiders' main defense system, are removed first.

Spiders in culture

This Moche ceramic depicts a spider, and dates from around 300 CE.

Spiders have been the focus of stories and mythologies of various cultures for centuries. Uttu, the ancient Sumerian goddess of weaving, was envisioned as a spider spinning her web. According to her main myth, she resisted her father Enki's sexual advances by ensconcing herself in her web, but let him in after he promised her fresh produce as a marriage gift, thereby allowing him to intoxicate her with beer and rape her. Enki's wife Ninhursag heard Uttu's screams and rescued her, removing Enki's semen from her vagina and planting it in the ground to produce eight previously-nonexistent plants.

In a story told by the Roman poet Ovid in his Metamorphoses, Arachne (Ancient Greek for "spider") was a Lydian girl who challenged the goddess Athena to a weaving contest. Arachne won, but Athena destroyed her tapestry out of jealousy, causing Arachne to hang herself. In an act of mercy, Athena brought Arachne back to life as the first spider. Stories about the trickster-spider Anansi are prominent in the folktales of West Africa and the Caribbean.

In some cultures, spiders have symbolized patience due to their hunting technique of setting webs and waiting for prey, as well as mischief and malice due to their venomous bites. The Italian tarantella is a dance to rid the young woman of the lustful effects of a spider bite. Web-spinning also caused the association of the spider with creation myths, as they seem to have the ability to produce their own worlds. Dreamcatchers are depictions of spiderwebs. The Moche people of ancient Peru worshipped nature. They placed emphasis on animals and often depicted spiders in their art.

General and cited references

Further reading

Bilger, Burkhard (5 March 2007). "Spider Woman". The New Yorker. pp. 66-73.

Bristowe, W. S. (1976). The World of Spiders. Taplinger Publishing Company. ISBN 978-0-8008-8598-4. OCLC 256272177.

Crompton, John (1950). The Life of the Spider. New York: Mentor. OCLC 1979220.

Hillyard, Paul (1994). The Book of the Spider: From Arachnophobia to the Love of Spiders. New York: Random House. ISBN 978-0-679-40881-9. OCLC 35231232.

Kaston, B. J.; Kaston, Elizabeth (1953). How to Know the Spiders: Pictured-Keys for Determining the More Common Spiders, with Suggestions for Collecting and Studying Them (1st ed.). Dubuque, Iowa: W.C. Brown Company. OCLC 628203833.

Main, Barbara York (1975). Spiders. Sydney: Collins. ISBN 978-0-00-211443-1. OCLC 123151744.

Wise, David A. (1993). Spiders in Ecological Webs. Cambridge Studies in Ecology. Cambridge: Cambridge University Press. ISBN 978-0-521-32547-9. OCLC 25833874.

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Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species diversity among all orders of organisms. Spiders are found worldwide on every continent except for Antarctica, and have become established in nearly every land habitat.

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