Reptile

? Reptiles
Eastern Hermann's Tortoise
Scientific classification
Kingdom: Animalia Phylum: Chordata Subphylum: Vertebrata Class: Sauropsida Goodrich, 1916
Orders
See text.

Reptiles (also called 'herps') are tetrapods and amniotes, animals whose embryos are surrounded by an amniotic membrane. Today they are represented by four surviving orders:

• Crocodilia (crocodiles, caimans and alligators): 23 species
• Rhynchocephalia ( tuataras from New Zealand): 2 species
• Squamata (lizards, snakes and amphisbaenids ("worm-lizards")): approximately 7,600 species
• Testudines (turtles): approximately 300 species

Reptiles are found on every continent except for Antarctica, although their main distribution comprises the tropics and subtropics. Though all cellular metabolism produces some heat, modern species of reptiles do not generate enough to maintain a constant body temperature and are thus referred to as " cold-blooded" (ectothermic). (See the Leatherback Sea Turtle for an exception: a reptile that elevates its body temperature well above that of its surroundings.) Instead they rely on gathering and losing heat from the environment to regulate their internal temperature, e.g, by moving between sun and shade, or by preferential circulation — moving warmed blood into the body core, while pushing cool blood to the periphery. In their natural habitats, most species are adept at this, and can maintain core body temperatures within a fairly narrow range, comparable to that of mammals and birds, the two surviving groups of " warm-blooded" animals. While this lack of adequate internal heating imposes costs relative to temperature regulation through behaviour, it also provides a large benefit by allowing reptiles to survive on much less food than comparably-sized mammals and birds, who burn much of their food for warmth. While warm-blooded animals move faster in general, an attacking lizard, snake or crocodile moves very quickly.

Except for some members of the Testudines, all reptiles are covered by scales.

Most reptile species are oviparous (egg-laying). Many species of squamates, however, are capable of giving live birth. This is achieved, either through ovoviviparity (egg retention), or viviparity (babies born without use of calcified eggs). Many of the viviparous species feed their fetuses through various forms of placenta, just like mammals (Pianka & Vitt, 2003 pgs: 116-118). They often provide considerable initial care for their hatchlings.

However, note the taxonomy issues described below; mammals and birds can also be viewed as descendants of reptiles.

Classification of reptiles

From the classical standpoint, reptiles included all the amniotes except birds and mammals. Thus reptiles were defined as the set of animals that includes crocodiles, alligators, tuatara, lizards, snakes, amphisbaenians and turtles, grouped together as the class Reptilia (Latin repere, "to creep"). This is still the usual definition of the term.

However, in recent years, many taxonomists have begun to insist that taxa should be monophyletic, that is, groups should include all descendants of a particular form. The reptiles as defined above would be paraphyletic, since they exclude both birds and mammals, although these also developed from the original reptile. Colin Tudge writes:

Mammals are a clade, and therefore the cladists are happy to acknowledge the traditional taxon Mammalia; and birds, too, are a clade, universally ascribed to the formal taxon Aves. Mammalia and Aves are, in fact, subclades within the grand clade of the Amniota. But the traditional class reptilia is not a clade. It is just a section of the clade Amniota: the section that is left after the Mammalia and Aves have been hived off. It cannot be defined by synamorphies, as is the proper way. It is instead defined by a combination of the features it has and the features it lacks: reptiles are the amniotes that lack fur or feathers. At best, the cladists suggest, we could say that the traditional Reptila are 'non-avian, non-mammalian amniotes'. (Tudge, p.85)

Some cladists thus redefine Reptilia as a monophyletic group, including both the classic reptiles as well as the birds and perhaps the mammals (depending on ideas about their relationships). Others abandon it as a formal taxon altogether, dividing it into several different classes. However, other biologists believe that the common characters of the standard four orders are more important than the exact relationships, or feel that redefining the Reptilia to include birds and mammals would be a confusing break with tradition. A number of biologists have adopted a compromise system, marking paraphyletic groups with an asterisk, e.g. class Reptilia*. Colin Tudge notes other uses of this compromise system:

By the same token, the traditional class Amphibia becomes Amphibia*, because some ancient amphibian or other gave rise to all the amniotes; and the phylum Crustacea becomes Crustacea*, because it may have given rise to the insects and myriapods (centipedes and millipedes). If we believe, as some (but not all) zoologists do, that myriapods gave rise to insects, then they should be called Myriapoda*....by this convention Reptilia without an asterisk is synonymous with Amniota, and includes birds and mammals, whereas Reptilia* means non-avian, non-mammalian amniotes. (Tudge, p.85)

Recent college-level references, such as Benton (2004) [1], offer another compromise by applying traditional ranks to accepted phylogenetic relationships. In this case, reptiles belong to the Class Sauropsida, and mammal-like reptiles to the Class Synapsida, with birds and mammals separated into their own traditional classes.

Reptile Groups

• Class Sauropsida

• Family Protorothyrididae - Hylonomus (extinct)
• Subclass Anapsida

Family Mesosauridae (extinct)

• Order Procolophonida - Pareiasaurs (extinct)
  
• Order Testudines - Turtles
  

• Subclass Diapsida

• Superorder Ichthyopterygia - Ichthyosaurs (extinct)
  
• Superorder Sauropterygia - Plesiosaurs (extinct)
• Superorder Lepidosauria

• Order Rhynchocephalia - Tuataras
  
• Order Squamata
  

Suborder Lacertilia- Lizards

Suborder Serpentes - Snakes

Suborder Amphisbaenia - Worm lizards

• Order Crocodilia - Crocodilians
  
• Order Pterosauria - Pterodactyls (extinct)
  
• Superorder Dinosauria - Dinosaurs
  

• Order Ornithischia (extinct)
  
• Order Saurischia
  

• Class Aves - Birds

Evolution of the reptiles

Several thousand fossil species showing a clear smooth transition from the ancestors of reptiles to present-day reptiles exist.

Hylonomus is the oldest-known reptile, and was about 8 to 12 inches (20 to 30 cm) long. Westlothiana has been suggested as the oldest reptile, but is for the moment considered to be more related to amphibians than amniotes. Petrolacosaurus, Araeoscelis, Paleothyris, Hylonomus, Ophiacodontidae, Archaeothyris, mesosaurs and Ophiacodon are other examples. The first true "reptile" or Amniotes are categorized as Anapsids, having a solid skull with holes only for nose, eyes, spinal cord, etc. Turtles are believed by some to be surviving Anapsids, as they also share this skull structure; but this point has become contentious lately, with some arguing that turtles reverted to this primitive state in order to improve their armor. Both sides have strong evidence, and the conflict has yet to be resolved.

Shortly after the first reptiles, two branches split off, one leading to the Anapsids, which did not develop holes in their skulls. The other group, Diapsida, possessed a pair of holes in their skulls behind the eyes, along with a second pair located higher on the skull. The Diapsida split yet again into two lineages, the lepidosaurs (which contain modern snakes, lizards and tuataras, as well as (in debate) the extinct sea reptiles of the Mesozoic) and the archosaurs (modernly represented by only crocodiles and birds, but containing pterosaurs and dinosaurs).

The earliest, solid-skulled amniotes also gave rise to a separate line, the Synapsida. Synapsids developed a pair of holes in their skulls behind the eyes (similar to the diapsids), which were used to both lighten the skull and increase the space for jaw muscles. The synapsids eventually evolved into mammals, and are often refered to as mammal-like reptiles, though they are not true members of the class Sauropsida.

Systems

Circulatory

Most reptiles have closed circulation via a three-chamber heart consisting of two atria and one, variably-partitioned ventricle. There is usually one pair of aortic arches. In spite of this, due to the fluid dynamics of blood flow through the heart, there is little mixing of oxygenated and deoxygenated blood in the three-chamber heart. Furthermore, the blood flow can be altered to shunt either deoxygenated blood to the body or oxygenated blood to the lungs, which gives the animal greater control over its blood flow, allowing more effective thermoregulation and longer diving times for aquatic species. There are some interesting exceptions among reptiles. For instance, crocodilians have an incredibly complicated four-chamber heart that is capable of becoming a functionally three-chamber heart during dives (Mazzotti, 1989 pg 47). Also, it has been discovered that some snake and lizard species (e.g., monitor lizards and pythons) have three-chamber hearts that become functional four-chamber hearts during contraction. This is made possible by a muscular ridge that subdivides the ventricle during ventricular diastole and completely divides it during ventricular systole. Because of this ridge, some of these squamates are capable of producing ventricular pressure differentials that are equivalent to those seen in mammalian and avian hearts (Wang et al, 2003).

Respiratory

All reptiles breathe using lungs. Exceptions to this are in aquatic turtles. These animals have developed more permeable skin, and even gills in their anal region, for some species (Orenstein, 2001). Even with these adaptations, breathing is never fully accomplished without lungs. Lung ventilation is accomplished differently in each main reptile group. In squamates the lungs are ventilated almost exclusively by the axial musculature. This is also the same musculature that is used during locomotion. Because of this constraint, most squamates are forced to hold their breath during intense runs. Some, however, have found a way around it. Varanids, and a few other lizard species, employ buccal pumping as a complement to their normal "axial breathing." This allows the animals to completely fill their lungs during intense locomotion, and thus remain aerobically active for a long time. Tegu lizards are known to possess a proto- diaphragm, which separates the pulmonary cavity from the visceral cavity. While not actually capable of movement, it does allow for greater lung inflation, by taking the weight of the viscera off the lungs (Klein et al, 2003). Crocodilians actually have a muscular diaphragm that is analogous to the mammalian diaphragm. The difference is that the muscles for the crocodilian diaphragm pull the pubis (part of the pelvis, which is movable in crocodilians) back, which brings the liver down, thus freeing space for the lungs to expand. This type of diaphragmatic setup has been referred to as the "hepatic piston."

Also, there are the Turtles & Tortoises. How these animals breathe has been the subject of much study. To date, only a few species have been studied thoroughly enough to get an idea of how turtles do it. The results indicate that turtles & tortoises have found a variety of solutions to this problem. The problem is that most turtle shells are rigid and do not allow for the type of expansion and contraction that other amniotes use to ventilate their lungs. Some turtles such as the Indian flapshell (Lissemys punctata) have a sheet of muscle that envelopes the lungs. When it contracts, the turtle can exhale. When at rest, the turtle can retract the limbs into the body cavity and force air out of the lungs. When the turtle protracts its limbs, the pressure inside the lungs is reduced, and the turtle can suck air in. Turtle lungs are attached to the inside of the top of the shell (carapace), with the bottom of the lungs attached (via connective tissue) to the rest of the viscera. By using a series of special muscles (roughly equivalent to a[diaphragm]), turtles are capable of pushing their viscera up and down, resulting in effective respiration, since many of these muscles have attachment points in conjunction with their forelimbs (indeed, many of the muscles expand into the limb pockets during contraction). Breathing during locomotion has been studied in three species, and they show different patterns. Adult female green sea turtles do not breathe as they crutch along their nesting beaches. They hold their breath during terrestrial locomotion and breathe in bouts as they rest. North American box turtles breathe continuously during locomotion, and the ventilation cycle is not coordinated with the limb movements (Landberg et al., 2003). They are probably using their abdominal muscles to breathe during locomotion. The last species to have been studied is red-eared sliders, which also breathe during locomotion, but they had smaller breaths during locomotion than during small pauses between locomotor bouts, indicating that there may be mechanical interference between the limb movements and the breathing apparatus. Box turtles have also been observed to breathe while completely sealed up inside their shells (ibid).

Most reptiles lack a secondary palate, meaning that they must hold their breath while swallowing. Crocodylians have evolved a bony secondary palate that allows them to continue breathing while remaining submerged (and protect their brains from getting kicked in by struggling prey). Skinks (family Scincidae) also have evolved a bony secondary palate, to varying degrees. Snakes took a different approach and extended their trachea instead. Their tracheal extension sticks out like a fleshy straw, and allows these animals to swallow large prey without suffering from asphyxiation.

Excretion

Excretion with two small kidneys. Uric acid is the main nitrogenous waste product.

Nervous

Advanced nervous system compared to amphibians. They have twelve pairs of cranial nerves.

Sexual

Most reptiles reproduce sexually. Asexual reproduction has been identified in squamates in six families of lizards and one snake. In some species of squamates, a population of females are able to produce a unisexual diploid clone of the mother. This asexual reproduction called parthenogenesis occurs in several species of gecko, and is particularly widespread in the teiids (expecially Aspidocelis) and lacertids ( Lacerta). Parthenogentic species are also suspected to occur among chameleons, agamids, xantusiids, and typhlopids.

Amniotic eggs covered with leathery or calcareous shells: Amnion, chorion, and allantois present during embryonic life. No larval stages.