Crocodilia

Crocodilians Temporal range: Cretaceous - Recent
American Alligators, Alligator mississippiensis
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Sauropsida
Superorder:	Crocodylomorpha
(unranked):	Archosauria
Order:	Crocodilia Owen, 1842
Families
Gavialidae Alligatoridae Crocodylidae See below for prehistoric taxa
Aquatic and terrestrial range of crocodilians

Crocodilia is an order of large reptiles that appeared about 84 million years ago in the late Cretaceous Period (Campanian stage). They are the closest living relatives of birds (the only known living dinosaurs), as the two groups are the only known survivors of the Archosauria. Members of the crocodilian stem group, the clade Crurotarsi, appeared about 220 million years ago in the Triassic Period and exhibited a wide diversity of forms during the Mesozoic Era.

The correct vernacular term for this group is "crocodilians", not "crocodiles," although the latter term is sometimes used incorrectly to refer to alligators and caiman, or even their distant prehistoric relatives, "marine crocodiles".

Spelling

The group is often spelled 'Crocodylia' for consistency with the genus Crocodylus (Laurenti, 1768). However, Richard Owen used the -i- spelling when he published the name in 1842, so it is generally preferred in the scientific literature . The -i- spelling is also a more accurate Latinization of the Greek κροκόδειλος (krokodeilos, literally "pebble-worm", referring to the shape and texture of the animal).

Description

Size comparison of the largest recorded examples (typical sizes in many cases are much smaller) of several crocodilian species (one of them of prehistorical times) and a prehistoric pholidosaurid with a human.

The basic crocodilian body plan is a very successful one; modern species closely resemble their Cretaceous ancestors of 84 million years ago.DONKEY!. Mammals, too, have adapted to this body plan at least once in history. One ancestral whale family, the Ambulocetidae, were aquatic predators living in rivers and lakes, and they filled an ecological niche similar to the crocodilians.

Crocodilians have a flexible semi-erect (semi-sprawled) posture. They can walk in low, sprawled "belly walk," or hold their legs more directly underneath them to perform the "high walk." Most other reptiles can only walk in a sprawled position, and chameleons are the only modern reptiles with a more erect posture than crocodilians. The semi-erect posture makes it possible for some species to gallop on land if necessary. An Australian species can reach a speed of over 16 km/h while galloping on an irregular forest floor. Crocodilian ancestors, fast-moving terrestrial predators like the rauisuchians, actually had a fully erect posture, indicating that the sprawling and semi-erect posture of crocodilians evolved after they adapted to as semi-aquatic ambush predators. Their ankle bones, or tarsi are highly modified. Modern crocodilian locomotion is not a primitive trait, but a specialization for their semi-aquatic lifestyle.

Teeth and jaws

All crocodilians have, like Homo sapiens (humans), thecodont dentition (teeth set in bony sockets) but unlike mammals, they replace their teeth throughout life (though not in 'extreme' old-age). Juvenile crocodilians replace teeth with larger ones at a rate as high as 1 new tooth per socket every month. After reaching adult size in a few years, however, tooth replacement rates can slow to two years and even longer. Very old members of some species have been seen in an almost "edentulous" (toothless) state, after teeth have been broken and replacement slowed or ceased. The result of this is that a single crocodile can go through at least 3,000 teeth in its lifetime. Each tooth is hollow, and the new one is growing inside the old. In this way, a new tooth is ready once the old is lost.

From the top: Head of an American alligator (Alligator mississippiensis), a Nile crocodile (Crocodylus niloticus), and an Indian gharial (Gavialis gangeticus).

Crocodilians have a secondary bony palate that enables them to breathe when partially submerged, even if the mouth is full of water. Their internal nostrils open in the back of their throat, where a special part of the tongue called the "palatal valve" closes off their respiratory system when they are underwater. This way they can open their mouths underwater without choking. Most reptiles lack a secondary palate, but some skinks (family Scincidae) have evolved a bony secondary palate too, to varying degrees.

Crocodiles and gharials have modified salivary glands on their tongue (salt glands), which are used for excreting excess salt ions from their body. Alligators and caimans have them too, but here they are non-functioning. This indicates that at some point the common origin of the Crocodylia were adapted to saline/marine environments. This also explains their wide disribution across the continents (i.e. marine dispersal). Species like the saltwater crocodile (C. porosus) can survive protracted periods of time in the sea, and can hunt prey within this environment.

Crocodilians are often seen lying with their mouths open, a behavior called gaping. One of its functions is probably to cool them down, but since they also do this at night and when it is raining, it is possible that gaping has a social function too.

Internal organs

Crocodilians lack a vomeronasal organ (yet it is detectable in the embryo) and a urinary bladder.

Like mammals and birds and unlike other reptiles, crocodiles have a four-chambered heart; however, unlike mammals, oxygenated and deoxygenated blood can be mixed because of the presence of the left aortic arch. The right ventricle has two arteries leaving it; a pulmonary artery, which goes to the lungs, and the left aortic arch, which goes to the body, or systemic circulation. There is also a hole, the foramen of Panizza, between the left and right aortic arches. Because the left aortic arch goes directly to the gut, the shunting of oxygen depleted blood which is high in CO₂ may serve to aid in creating stomach acid to assist in digesting bones from its prey. Their blood has been shown to have strong antibacterial properties.

They have alveoli in their lungs and a unique muscular attachment to the liver and viscera that acts as a piston to breathing, separating the thoracic and abdominal cavities (similar to the diaphragm of mammals). Although tegu lizards have a primitive proto-diaphragm, separating the pulmonary cavity from the visceral cavity and allowing greater lung inflation, this has a different evolutionary history.

Crocodylians are known to swallow stones, gastroliths ("stomach-stones"), which act as a ballast in addition to aiding post-digestion processing of their prey. The crocodiylian stomach is divided into two chambers, the first one is described as being powerful and muscular, like a bird gizzard. This is where the gastroliths are found. The other stomach has the most acidic digestive system of any animal, and it can digest mostly everything from their prey; bones, feathers and horns.

The sex of developing crocodilians is determined by the incubation temperature of the eggs. This means crocodilians do not have genetic sex determination, but instead have a form of environmental sex determination which is based upon the temperature embryos are subjected to early in their development.

Sensory organs

Like all reptiles, crocodilians have a relatively small brain, but it is more advanced than in other reptiles. Among other things it has a true cerebral cortex.

As in many other aquatic or amphibian tetrapods, the eyes, ears, and nostrils are all located on the same plane. They see well during the day and may even have color vision, plus the eyes have a vertical, cat-like pupil which also gives them excellent night vision. The iris is silvery and a light reflecting layer of tapetum behind the retina greatly increases their ability to see in weak light and also makes their eyes glow in the dark. A third transparent eyelid, the nictitating membrane, protects their eyes underwater. However, they cannot focus under water, meaning other senses are more important when submerged under water.

While birds and most reptiles have a ring of bones around each eye which supports the eyeball (the sclerotic ring), the crocodiles lack these bones, just like mammals and snakes. The eardrums are located behind the eyes and are covered by a movable flap of skin. This flap closes, along with the nostrils and eyes, when they dive, preventing water from entering their external head openings. The middle ear cavity has a complex of bony air-filled passages and a branching eustachian tube. There is also a small muscle (which is also seen in gecko) next to or upon the stapes, the stapedius, which probably functions in the same way as the mammalian stapedius muscle does, dampening strong vibrations.

The upper and lower jaws are covered with sensory pits, seen as small, black speckles on the skin, the crocodile version of the lateral organ we see in fish and many amphibians. But they have a completely different origin. These pigmented nodules encase bundles of nerve fibers that respond to the slightest disturbance in surface water, detecting vibrations and small pressure changes in water, making it possible for them to detect prey, danger and intruders even in total darkness. These sense organs are known as DPRs (Dermal Pressure Receptors). While alligators and caimans only have them on their jaws, crocodiles have similar organs on almost every scale on their body. The function of the DPRs on the jaws are clear, but it is still not quite clear what the organs on the rest of the body in crocodiles actually do. They are probably doing the same as the organs on their jaws, but it seems like they can do more than that, like assisting in chemical reception or even salinity detection.

Skin and skeleton

West African dwarf crocodile from the forests of West and West Central Africa

The skin is covered with non-overlapping scales composed of the protein keratin (the same protein that forms hooves, skin, horns, feathers, hair, claws and nails in other tetrapods), which are shed individually. On the head the skin is actually fused to the bones of the skull. There are small plates of bone, called osteoderms or scutes, under the scales. Like the scales comprising the shell of a turtle, or the cross-section of a tree trunk, crocodile osteoderms have annual growth rings, and by counting them it is possible to tell their age. Osteoderms are found especially on the back, and in some species also on the belly. The rows of scutes cover the crocodile's body from head to tail, forming a tough protective armor. Beneath the scales and osteoderms is another layer of armor, both strong and flexible and built of rows of bony overlapping shingles called osteoscutes, which are embedded in the animal's back tissue. The blood-rich bumpy scales seen on their backs act as solar panels.

Their spool-shaped vertebrae in their ancestors went from being biconcave to having a concave front and a convex back in the modern forms. This made the vertebral column more flexible and strong.

They possess ribs of dermal origin restricted to the sides of the ventral body wall. The collar bone (clavicle) is absent.

Differences between alligators and crocodiles

File:Young crocodile displaying fourth tooth.jpg

A crocodile showing how, unlike an alligator, its fourth lower tooth sits outside the mouth when the jaw is shut.

Crocodiles are generally much lighter in color.

While alligators are often confused with crocodiles, they belong to two quite separate taxonomic families. Every type of crocodile, except the American crocodile, is usually larger and heavier than alligators, has fewer teeth, and has a shorter life-span.

The most obvious external differences are visible in the head–alligators have wider and shorter heads, and a more U-shaped than V-shaped snout. The alligator's upper jaw is wider than its lower jaw, and the teeth in the lower jaw fit into small depressions in the upper jaw. The upper and lower jaws of the crocodile are the same width, and teeth in the lower jaw fit fall along the edge or outside the upper jaw when the mouth is closed. When the crocodile's mouth is closed, the large fourth tooth in the lower jaw fits into a constriction in the upper jaw. For indistinguishable specimens, the protruding tooth is the most reliable feature to define a species.

Alligators lack the jagged fringe which appears on the hind legs and feet of the crocodile and have the toes of the hind feet webbed, not more than half way to the tips. Alligators strongly prefer freshwater, while crocodiles can better tolerate saltwater due to specialized glands for filtering out salt. However, both taxa can survive in either.

Both species of alligator also tend to be darker in color than crocodiles–often nearly black (but color is very dependent on water quality). Algae-laden waters produce greener skin, while tannic acid from overhanging trees can produce often darker skin.

Alligators tend to be clumsier on both land and water than crocodiles. Some zookeepers have reportedly been able to take more liberties with alligators than with crocodiles. When cleaning alligator pools, some zoo keepers can tread on alligators without soliciting a response, though crocodiles almost invariably react aggressively.

Evolution

Eusuchia, a modern clade which includes the crown group Crocodilia, first appeared in the Lower Cretaceous of Europe. Isisfordia duncani lived approximately 95 to 98 million years ago, during the Cenomanian epoch of the Upper Cretaceous. Isisfordia is the second oldest known eusuchian, and the earliest crocodylomorph yet found in Australia. Eusuchians underwent a mass radiation during the Late Cretaceous and the Paleogene, in which they evolved into numerous forms, such as semi-aquatic dinosaur-eating species (Deinosuchus); hooved, terrestrial carnivores (Pristichampsus), and 'hatchet'-shaped skulled forms (Baru).

Classification

• Superorder Crocodylomorpha
  • Order Crocodilia
    • Superfamily Gavialoidea
      • Family Gavialidae - Gharials & False Gharials
    • Superfamily Alligatoroidea
      • Family Alligatoridae
        • Subfamily Diplocynodontinae (extinct)
        • Subfamily Alligatorinae - Alligators
        • Subfamily Caimaninae - Caimans
    • Superfamily Crocodyloidea
      • Family Crocodylidae
        • Subfamily Mekosuchinae (extinct)
        • Subfamily Crocodylinae - Crocodiles

Phylogeny

Cladogram after Brochu (1997).

Eusuchia
  ├──Hylaeochampsa
  └──┬──Allodaposuchus
     └──Crocodilia
          ├──Gavialoidea
          │     ├──Eothoracosaurus
          │     └──┬──Thoracosaurus
          │        └──┬──Argochampsa
          │           ├──Eosuchus
          │           └──Gavialidae
          └──┬──Borealosuchus
             └──┬──Pristichampsus
                └──Brevirostres
                    ├──Alligatoroidea
                    │   ├──Leidyosuchus
                    │   ├─?Deinosuchus
                    │   └──Globidonta
                    │       ├──Stangerochampsa
                    │       ├──Brachychampsa
                    │       └──Alligatoridae
                    └──Crocodyloidea
                        ├──Prodiplocynodon
                        └──┬──Asiatosuchus
                           └──┬──Brachyuranochampsa
                              └──┬──Harpacochampsa
                                 └──Crocodylidae

In popular culture

• Several science fiction movies have giant crocodilians as their stars, such as Lake Placid, DinoCroc, and the crocodile series.

External links

• Before Crocodiles, the Hairless Coyote
• Florida's Museum of Natural History's Crocodilians Natural History & Conservation site
• International Crocodile Rescue
• Mikko's Phylogeny Archive Crocodyliformes