Reptiles: Nonavian Diapsid Amniotes Learning Outcomes: • Justify the statement that “the amniotic egg provided solutions that made development apart from external watery environments possible.” • Compare amniote taxonomy before and after the application of cladistic methods. Evolutionary Perspective • Amniota – Monophyletic lineage including reptiles, birds, and mammals – Amniotic eggs • Extraembryonic membranes protect embryo from desiccation; cushion embryo; promote gas transfer, and store waste. – Impervious skin, nails, waterconserving kidneys, large lungs Figure 20.2 The amniotic egg. Cladistic Interpretation of the Amniotic Lineage • Mammals – Most closely related to ancestral amniotes – Reptilian lineage • Birds • Dinosaurs • Other reptiles – Traditional class “Reptilia” is paraphyletic. Early Amniote Evolution and Skull Structure • Amniote lineages – Synapsida • Mammals • Single opening (fenestra) in temporal region of skull – Anapsida • Skull without fenestra • Entirely extinct – Diapsida • Two fenestrae • All living reptiles (including birds) and numerous extinct lineages Figure 20.4 Amniote kull characteristics. Survey of the Reptiles • Order Testudines – Turtles – Bony shell, limbs articulate internally to ribs, keratinized beak rather than teeth – Shell • Carapace (dorsal) and plastron (ventral) – oviparous Figure 20.5 Skeleton of a turtle. Survey of the Reptiles • Order Crocodylia – Archosaur lineage – Alligators, crocodiles, gavials, caimans – Openings in front of eyes, triangular eye orbits, laterally compressed teeth – Secondary palate – Oviparous with parental care Survey of the Reptiles • Order Sphenodontida – Tuataras – Two rows of teeth on upper jaw and single row of teeth in lower jaw – New Zealand – Oviparous Figure 20.7 Order Sphenodontia (Sphenodon punctatus). Survey of the Reptiles • Order Squamata – Kinetic skull • Moveable quadrate bones and other skull modifications • Increases skull flexibility – Suborder Sauria—lizards • • • • Usually two pairs of legs Jaws unite anteriorly. Oviparous, ovoviviparous, or viviparous Includes legless amphisbaenias – Suborder Serpentes—snakes • Legless • Skull adaptions for swallowing large prey • Most oviparous Figure 20.8 Order Squamata (Heloderma suspectum). Figure 20.9 Order Squamata, an amphisbaenian “worm lizard” (Amphisbaenia alba). Evolutionary Pressures • External structure and locomotion – Skin • Thick, dry, and keratinized • Scales may be modified. • Epidermal layers shed through ecdysis. – Support and movement • Skeleton highly ossified • Skull with secondary palate • Additional cervical vertebrae (including atlas and axis) • Two or more sacral vertebrae Figure 20.11 The secondary palate. Evolutionary Pressures • Nutrition and the digestive system – Most carnivores • Turtles (carnivores, herbivores, or omnivores) – Tongues • Nonprotrusible – Turtles and crocodilians • Protrusibile – Lizards and snakes Figure 20.12 Order Squamata. A chameleon (Chameleo chameleon) using its tonge to capture prey. Evolutionary Pressures • Nutrition and the digestive system – Feeding adaptations of snakes • Bones of upper jaw are moveable on skull. • Ligaments loosely join the halves of upper and lower jaws anteriorly. • Posterior pointing teeth • Glottis opens forward in mouth. • Hinged maxillary bone in vipers • Venom glands in some – Modified salivary glands Figure 20.13 Feeding adaptations of snakes. (a) A copperhead (Askistrodon) ingesting prey. (b) The skull of a viper. (c) Note the hinged maxillary bone into which the fang is embedded swings forward when the mouth opens. Evolutionary Pressures • Circulation, gas exchange, and temperature regulation – Heart • Sinus venosus reduced to pacemaker (except turtles) • Two atria and incompletely divided ventricle (except crocodilians) – Permits shunting blood away from pulmonary circuit during intermittent breathing • Conus arteriosus and ventral aorta divided embryologically into three major arteries leaving heart Figure 20.14 Heart and major arteries of a lizard. Evolutionary Pressures • Gas exchange – Internal respiratory surfaces • Lungs spongelike – Large surface area for gas exchange – Negative pressure ventilation • Posterior movement of ribs and body wall expands body cavity. – Modified in turtles • Decreases body cavity pressure and draws air into lungs Evolutionary Pressures • Temperature Regulation – Most ectotherms • Exception brooding Indian pythons • Behavioral regulation – Orientation to sun’s rays – Warming by conduction from warm surfaces – Cooling by seeking shade or burrows, assuming erect posture, and nocturnal habits • Physiological regulation – Panting – Diverting blood to skin while basking (marine iguana) • Torpor – Temperate climates – Enter hibernacula » Not hibernation—body temperature is not regulated. Evolutionary Pressures • Nervous and Sensory Functions – Brain similar to other vertebrates • Cerebral hemispheres, optic lobes, and cerebellum enlarged – Sensory functions • Vision – Dominant sense – Accommodation by lens movement (snakes) or changing lens shape (other reptiles) – Upper and lower eyelids and nictitating membrane • Median (parietal eye) – Outgrowth of roof of forebrain • Hearing – Airborne and substrate borne vibrations Evolutionary Pressures • Nervous and Sensory Functions – Sensory functions • Olfaction – Jacobson’s (vomeronasal) organs » Diapsids, especially squamates • Temperature sense – Pit organs » Pit vipers • Magnetic sense – Sea turtles Evolutionary Pressures • Excretion and osmoregulation – Metanephric kidneys – Uric acid – Internal respiratory surfaces reduce water loss. – Water conserving behaviors • Nocturnal habits • Avoidance by burrowing – Water storage • Lymphatic spaces • Urinary bladder Evolutionary Pressures • Reproduction and development – Internal fertilization and amniotic egg makes development apart from external water sources possible. – Intromittent organ • Hemipenes – Parthenogesis • A few lizards and snakes – Courtship behaviors common – Eggs usually develop unattended by a parent. • American alligator and a few others are exceptions. Figure 20.15 Reptile eggs and young. The Madagascar day gecko (Phelsuma madagascariensis). Figure 20.16 Parental care in reptiles is not common. The American alligator (Alligator mississippiensis) is an exception. Further Phylogenetic Considerations • Archosaur branch – Diverged 200 mya – Branches • Pterosaurs • Dinosaur lineages including birds • Synapsid branch – Diverged 320 mya – Mammals