Though there is diversity in snout and tooth shape, all crocodilian species have essentially the same body morphology. Their limbs are reduced in size; the front feet have five mostly non-webbed digits, and the hind feet have four webbed digits and an extra fifth. The pelvis and ribs of crocodilians are modified; the
cartilaginous processes of the ribs allow the
thorax to collapse when submerging and the structure of the pelvis can accommodate large amounts of food, The crocodilian penis is permanently erect; it relies on cloacal muscles to protrude it, and elastic ligaments and a tendon to retract it. The
gonads are located near the
kidneys. Crocodilians range in size from the
dwarf caimans and
African dwarf crocodiles, which reach , to the
saltwater crocodile and Nile crocodile, which reach and weigh up to . Some prehistoric species such as the
Miocene caiman Purussaurus were even larger, with some estimates putting it over . Crocodilians tend to be
sexually dimorphic; males are much larger than females.
Locomotion swimming. Crocodilians are excellent swimmers. During
aquatic locomotion, the muscular tail undulates from side to side to drive the animal through the water while the limbs are held close to the body to reduce
drag. When the animal needs to stop or change direction, the limbs are splayed out. Crocodilians are less well-adapted for moving on land, and are unusual among vertebrates in having two means of terrestrial locomotion: the "high walk" and the "low walk". showing the crocodilian "high walk", with lower limb portions held almost vertically, unlike other reptiles. The limbs move much the same as those of other
quadrupeds; the left forelimb moves first, followed by the right hindlimb, then right forelimb, and finally left hindlimb. The high walk of crocodilians, with the belly and most of the tail held off the ground and the limbs held directly under the bodies, resembles that of mammals and birds. Though they are typically slow on land, crocodilians can produce brief bursts of speed; some can run at for short distances. In some small species, such as the
freshwater crocodile, running can progress to galloping, which involves the hind limbs launching the body forward and the fore limbs subsequently taking the weight. Next, the hind limbs swing forward as the spine flexes
dorso-ventrally, and this sequence of movements is repeated. During terrestrial locomotion, a crocodilian can keep its back and tail straight because muscles attach the scales to the vertebrae. A fast entry into water from a muddy bank can be effected by plunging to the ground, twisting the body from side to side and splaying out the limbs. The muscles that close the jaws are larger and more powerful than the ones that open them, The powerful closing muscles attach at the middle of the lower jaw. The jaw hinge attaches behind the
atlanto-occipital joint, giving the animal a wide gape. A folded membrane holds the tongue stationary. Crocodilians have some of the strongest
bite forces in the animal kingdom. In a study published in 2003, an American alligator's bite force was measured at up to ; and in a 2012 study, a saltwater crocodile's bite force was measured at . This study found no correlation between bite force and snout shape, though the gharial's extremely slender jaws are relatively weak and are built for quick jaw closure. Crocodilian teeth vary from dull and rounded to sharp and pointed. Crocodilians are
homodonts, meaning each of their teeth are of the same type; they do not have different tooth types, such as canines and molars. Crocodilians are
polyphyodonts; they are able to replace each of their approximately 80 teeth up to 50 times in their 35-to-75-year lifespan. Crocodilians are the only non-mammalian vertebrates with
tooth sockets. Next to each full-grown tooth is a small replacement tooth and an
odontogenic stem cell in the
dental lamina that can be activated when required. Tooth replacement slows and eventually stops as the animal ages. When in bright light, the pupils of a crocodilian contract into narrow slits, whereas in darkness they become large circles, as is typical for animals that hunt at night. Crocodilians' eyes have a
tapetum lucidum that enhances vision in low light. When the animal completely submerges, the
nictitating membranes cover its eyes. Glands on the nictitating membrane secrete a salty lubricant that keeps the eye clean. When a crocodilian leaves the water and dries off, this substance is visible as "tears". Crocodilians appear to have undergone a "nocturnal bottleneck" early in their history, during which their eyes lost traits like
scleral rings, an annular pad of the lens and coloured cone
oil droplets, giving them
dichromatic vision (red-green colourblindness). Since then, some crocodilians appear to have re-evolved
full-colour vision. The ears are adapted for hearing both in air and underwater, and the
eardrums are protected by flaps that can be opened or closed by muscles. Crocodilians have a wide
hearing range, with sensitivity comparable to most birds and many mammals. Hearing in crocodilians does not degrade as the animal ages because they can regrow and replace
hair cells. The well-developed
trigeminal nerve allows them to detect vibrations in water, such as those made by potential prey. Crocodilians have a single
olfactory chamber and the
vomeronasal organ disappears when they reach adulthood. Behavioural and olfactometer experiments indicate crocodiles detect both air-borne and water-soluble chemicals, and use their olfactory system for hunting. When above water, crocodiles enhance their ability to detect volatile odorants by gular pumping, a rhythmic movement of the floor of the
pharynx. Crocodiles appear to have lost their
pineal organ but still show signs of
melatonin rhythms.
Skin and scales The skin of crocodilians is clad in non-overlapping scales known as
scutes that are covered by
beta-keratin. Many of the scutes are strengthened by bony plates known as
osteoderms. Scutes are most numerous on the back and neck of the animal. The belly and underside of the tail have rows of broad, flat, square-shaped scales. Underneath the surface, the
dermis is thick with
collagen. Both the head and jaws lack scales and are instead covered in tight, keratinised skin that is fused directly to the bones of the skull and which, over time, develop a pattern of cracks as the skull develops. The skin on the neck and sides is loose. The scutes contain blood vessels and may act to absorb or release heat during
thermoregulation. Some scutes contain a single pore known as an integumentary sense organ. Crocodiles and gharials have these on large parts of their bodies, while alligators and caimans only have them on the head. Their exact function is not fully understood, but it has been suggested they may be
mechanosensory organs. There are prominent, paired integumentary glands in skin folds on the throat, and others in the side walls of the cloaca. Various functions for these have been suggested; they may play a part in communication—indirect evidence suggests they secrete
pheromones used in courtship or nesting. In the genus
Crocodylus, the skin contains
chromatophores, allowing animals to change colour from dark to light and
vice versa.
Circulation Crocodilians may have the most-complex vertebrate
circulatory system with a four-chambered
heart and two
ventricles, an unusual trait among extant reptiles. Both have left and right
aorta are connected by a hole called the
Foramen of Panizza. This system may allow the animals to remain submerged for a lengthy period, but this explanation has been questioned. Other possible reasons for the peculiar circulatory system include assistance with thermoregulatory needs, prevention of
pulmonary oedema, and quick recovery from
metabolic acidosis. Retention of carbon dioxide within the body permits an increase in the rate of gastric acid secretion and thus the efficiency of digestion, and other gastrointestinal organs such as the
pancreas,
spleen,
small intestine, and
liver also function more efficiently. When submerged, a crocodilian's heart may beat at only once or twice a minute, with little blood flow to the muscle. When it rises and takes a breath, its heart rate almost immediately increases and the muscles receive newly oxygenated blood. Unlike many
marine mammals, crocodilians have little
myoglobin to store oxygen in their muscles. While diving, an increasing concentration of
bicarbonate ions causes
haemoglobin in the blood to release oxygen for the muscles.
Respiration Crocodilians were traditionally thought to breathe like mammals, with airflow tidally moving in and out, but studies published in 2010 and 2013 conclude respiration in crocodilians is more
bird-like, with airflow moving in a unidirectional loop within the lungs. During inhalation, air flows through the trachea and into two primary
bronchi (airways) that divide into narrower secondary passageways. The air continues to move through these, then into even narrower tertiary airways, and then into other secondary airways that were bypassed the first time. The air then flows back into the primary airways and is exhaled. In crocodilians, the
diaphragmaticus muscle, which is
analogous to the
diaphragm in mammals, attaches the lungs to the liver and pelvis. During inhalation, the
external intercostal muscles expand the ribs, allowing the animal to take in more air, while the
ischiopubis muscle causes the hips to swing downwards and push the belly outward, while the diaphragmaticus pulls the liver back. When exhaling, the
internal intercostal muscles push the ribs inwards while the
rectus abdominis pulls the hips and liver forwards and the belly inward. Crocodilians can also use these muscles to adjust the position of their lungs, controlling their buoyancy in the water. An animal sinks when the lungs are pulled towards the tail and floats when they move back towards the head. This allows them to move through the water without creating disturbances that could alert potential prey. They can also spin and twist by moving their lungs laterally. When swimming and diving, crocodilians appear to rely on
lung volume for buoyancy more than for oxygen storage. When diving, the nostrils of a crocodilian shut tight. The depth to which crocodilians can dive is unknown, but crocodiles can dive to at least . Crocodilians vocalize by vibrating
vocal folds in the larynx. The folds of the American alligator have a complex morphology consisting of
epithelium,
lamina propria and muscle, and according to Riede et al. (2015): "it is reasonable to expect species-specific morphologies in vocal folds/analogues as far back as basal reptiles".
Digestion Crocodilian teeth can only hold onto prey, and food is swallowed unchewed. The stomach consists of a grinding
gizzard and a digestive chamber. Indigestible items are regurgitated as pellets. The stomach is more acidic than that of any other vertebrate and contains ridges for
gastroliths, which play a role in the crushing of food. Digestion takes place more quickly at higher temperatures. Compared to crocodiles, alligators digest more
carbohydrates relative to
protein. Crocodilians have a very low metabolic rate and thus low energy requirements. They can withstand extended fasting by living on
stored fat. Even recently hatched crocodiles are able to survive 58 days without food, losing 23% of their bodyweight during this time.
Thermoregulation basking and gaping Crocodilians are
ectotherms ('cold-blooded'), relying mostly on their environment to control their body temperature. The main means of warming is sun's heat, while immersion in water may either raise its temperature via
thermal conduction or cool the animal in hot weather. The main method for regulating its temperature is behavioural; temperate-living alligators may start the day by basking in the sun on land and move into water for the afternoon, with parts of the back breaking the surface so it can still be warmed by the sun. At night, it remains submerged and its temperature slowly falls. The basking period is longer in winter. Tropical crocodiles bask briefly in the morning and move into water for rest of the day. They may return to land at nightfall when the air cools. Animals also cool themselves by gaping the mouth, which cools by
evaporation from the mouth lining. By these means, the temperature range of crocodilians is usually maintained between , and mainly stays in the range . Both the
American and
Chinese alligator can be found in areas that sometimes experience periods of frost in winter. In cold weather, alligators remain submerged with their tails in deeper, less-cold water and their nostrils projecting just above the surface. If ice forms on the water, they maintain ice-free breathing holes, and there have been occasions when their snouts have become frozen into ice. Temperature-sensing probes implanted in wild American alligators have found their core body temperatures can fall to around , but as long as they remain able to breathe, they show no ill effects when the weather warms. Water is lost during breathing, and salts and water are lost in the urine and faeces, through the skin, and in crocodiles and gharials via
salt-excreting glands on the tongue. In fresh water, the
osmolality (the concentration of solutes that contribute to a solution's
osmotic pressure) in the
plasma is much higher than that of the surrounding water. The animals are well-hydrated, the urine in the cloaca is abundant and dilute, and nitrogen is excreted as
ammonium bicarbonate. Sodium loss is low and mainly occurs through the skin in freshwater conditions. In seawater, the opposite is true; the osmolality in the plasma is lower than that of the surrounding water, causing the animal to dehydrate. The cloacal urine is much more concentrated, white, and opaque, and nitrogenous waste is mostly excreted as insoluble
uric acid. ==Distribution and habitat==