Evolution and Diversity of Amniotes: The Transition from Aquatic to Terrestrial Life

1. CHAPTER 26 Amniote Origins and Nonavian Reptiles

3. Diversity Amniotic “Pond” • Any animal with a shell-less egg remains tied to water • Development of a shelled egg freed the reptilian groups to exploit land • Extraembryonic membranes from previous evolutionary aquatic stages are maintained • Allantois serves as a respiratory surface and a chamber to store nitrogenous wastes • Chorion allows O2 and CO2 to freely pass • Surrounding the organism is a porous, parchment-like or leathery shell

4. Diversity Amniotes • Amniota • Nonavian reptiles, birds, and mammals • Developing young enclosed by extraembryonic membrane called the amnion • Secretes fluid, amniotic fluid, in which embryo/fetus floats

5. Origin and Early Evolution of Amniotes History • separated into groups based on three patterns of fenestra in the temporal region • Anapsids • Have a skull with no temporal opening behind the orbits • Modern turtles • Diapsids • Gave rise to all other traditional “reptiles” (except turtles) and to birds • Skull has two temporal openings • One pair below the cheeks and another above

7. Origin and Early Evolution of Amniotes • Synapsids • Mammal-like reptiles with a single pair of temporal openings low on the cheeks • These openings are associated with large muscles that elevate the lower jaw • Changes in jaw musculature • Might reflect a shift from suction feeding in aquatic vertebrates to terrestrial feeding • Required larger muscles to produce static pressure • Functional significance of the evolution of temporal openings in amniotes is not fully understood

9. Origin and Early Evolution of Amniotes Derived Characters of Amniotes Amniotic egg • All amniotes have eggs with four extraembryonic membranes • Amnion, allantois, chorion, and yolk sac • Amnion • Encloses the embryo in fluid-filled space • Cushions the embryo and provides an aqueous medium for growth • Allantois • Stores metabolic wastes

11. Origin and Early Evolution of Amniotes • Chorion • Surrounds embryo and all other extraembryonic membranes • Lies just beneath shell • Highly vascularized • Respiratory surface • Allantois and chorion sometimes fuse to form respiratory structure: chorioallantoic membrane • Yolk sac • Nutrient storage

12. Origin and Early Evolution of Amniotes • Amniotic egg permitted development of a larger, faster-growing embryo • One hypothesis suggests that one step in the evolution of the amniotic egg was replacement of the jelly layer with a shell • Provided better support and movement of oxygen • Shell could also be broken down to provide calcium for growing skeletal structures • All amniotes • Lack gilled larvae and have internal fertilization • Eliminated the need for aquatic environments • copulatory organ • Derived from cloacal wall, appears to be an amniote innovation

13. Origin and Early Evolution of Amniotes Thicker and more waterproof skin • Amniote skin is thick and tends to be more keratinized and less permeable to water • Variety of structures composed of keratin such as scales, hair, feathers, and claws project from the skin • Keratin protects the skin from physical trauma, and lipids prevent water loss • Few amniotes use skin as a primary respiratory organ because keratin and lipids limit exchange of gases

14. Characteristics of Nonavian Reptiles • Resistance to desiccation • Provided primarily hydrophobic lipids in epidermis • Scales of nonavian reptiles • Formed mostly of beta keratin and provide protection against wear in terrestrial environments • In crocodilians, scales remain throughout life • In other nonavian reptiles such as lizards and snakes, new keratinized epidermis replaces old epidermis which is shed • Turtles have scutes • Platelike modified scales • Crocodiles and many lizards possess osteoderms • Bony plates located the dermis beneath scales

15. Characteristics of Nonavian Reptiles • Nonavian reptiles have better developed lungs than amphibians • Nonavian lungs have more surface area than amphibians’ lungs • Nonavian ventilation occurs by drawing air into the lungs rather than pushing air into the lungs • Nonavian reptiles expand the thoracic cavity thus expanding the rib cage in order to draw air in • Pulmonary respiration is supplemented by respiration in the cloaca or pharynx in many aquatic turtles and by cutaneous respiration in sea snakes and turtles

16. Characteristics of Nonavian Reptiles • Jaws of nonavian reptile • Efficiently designed for applying crushing or gripping force to prey • Larger jaw muscles have mechanical advantage over the jaws of fishes which are designed for suction feeding or for quick closure • Tongue is muscular and mobile • Functions to move food in mouth for mastication and swallowing

17. Characteristics of Nonavian Reptiles • Nonavian reptiles have an efficient and versatile circulatory system and higher blood pressure than amphibians • Right atrium receives deoxygenated blood and is partitioned from the left atrium which receives oxygenated blood • Crocodilians have separated ventricles • In other nonavian reptiles, ventricle is incompletely partitioned into multiple chambers • All nonavian reptiles have two functionally separate circulations

19. Characteristics of Nonavian Reptiles • Nonavian reptiles haveefficient strategies for water conservation • secrete nitrogenous wastes as uric acid Relatively non-toxic • All amniotes have metanephric kidneys • Nonavian reptiles cannot concentrate urine in the kidneys • Urinary bladder receives undiluted urine • Water and most salts • Resorbed in the bladder • “urine” voided as a semisolid mass of uric acid • Salt is removed by salt glands near the nose, eyes, or tongue

20. Characteristics of Nonavian Reptiles • The nervous system of nonavian reptiles is more complex than that of amphibians • Brain of nonavian reptiles small compared to brain of other amniotes • However, cerebrum of all amniotes is relatively enlarged • Enlargement of cerebrum correlated with integration of sensory information and muscle control during locomotion • Nonavian reptiles have good vision • Snakes and many lizards use a highly sensitive sense of smell to find prey and mates • Olfaction assisted by a Jacobson’s organ, a specialized olfactory chamber in the roof of the mouth

21. Video Links • Reptiles of Africa • The four monophyletic nonavian reptilian groups are: • Turtles • Crocodilians • Squamates • tuataras

22. Characteristics and Natural History of Reptilian Orders Turtles (Chelonia) • Shells consist of a dorsal carapace and a ventral plastron • Unique among vertebrates, limbs and limb girdles are located deep to the ribs • Lack teeth and use tough, horny plates for gripping food • Consequence of having a rigid shell • Turtle cannot expand chest to breathe • Solve problem by using abdominal and pectoral muscles as a “diaphragm” • Air is drawn in by contraction of the limb flank muscles, increasing abdominal cavity volume

24. Characteristics and Natural History of Reptilian Orders • Snapping Turtles • Have a reduced shell that does not permit full withdrawal of the body • Jaws are adequate defense • Entirely carnivorous and can eat fish, frogs, waterfowl, etc. • Aquatic but must come ashore to lay eggs • Loggerhead Turtles

26. Characteristics and Natural History of Reptilian Orders Squamates: Lizards and Snakes • Account for up 95% of living nonavian reptiles • Allowed evolution in lizards of a mobile skull with movable joints, a kinetic skull • Exceptional skull mobility of snakes is considered a major factor in their diversification • Snake Fakes • Jesus Christ Lizard

29. Characteristics and Natural History of Reptilian Orders Suborder Sauria: Lizards • Diverse group with terrestrial, burrowing, aquatic, arboreal, and some aerial members • Lizards • Geckos • Small, agile, nocturnal forms • Adhesive toe pads allow them to walk on ceilings • Iguanids • Include many New World lizards as well as the marine iguana of the Galápagos • Chameleons • Arboreal lizards of Africa and Madagascar • Many have an extendible tongue

32. Characteristics and Natural History of Reptilian Orders • Some have degenerate limbs • Glass lizards are nearly limbless • Movable eyelids whereas snakes have a transparent covering • Amphisbaenians or “worm lizards” highly specialized for a fossorial (burrowing life

34. Characteristics and Natural History of Reptilian Orders • Conserve water by producing semisolid urine with a high content of crystalline uric acid • Some can store fat in tails to provide energy and metabolic water during drought • Gila monster and beaded lizard are the only lizards capable of a venomous bite • Lizards keep body temperature relatively constant by behavioral thermoregulation, although they are ectotherms • Successful strategy in ecosystems with low productivity and warm climates, such as tropical deserts and grasslands

37. Characteristics and Natural History of Reptilian Orders Suborder Serpentes: Snakes • Limbless and have lost pectoral and pelvic girdles (except in pythons) • Feeding apparatus allows them to eat prey several times their own diameter • Two halves of lower jaw are loosely joined, allowing them to spread apart • Skull bones also loosely articulated so mouth can accommodate large prey • To allow breathing during the slow process of swallowing, the tracheal opening is extended • Sidewinding snake

39. Characteristics and Natural History of Reptilian Orders • Eyeballs have reduced mobility and a permanent corneal membrane for protection • Most have poor vision • Arboreal snakes in tropical forests have highly developed vision • Lack external ears and do not respond to most aerial sounds • Can feel vibrations at low frequencies, especially vibrations carried in the ground • Chemical senses rather than vision or hearing are main senses used to hunt prey

41. Characteristics and Natural History of Reptilian Orders • Jacobson’s organs • Pair of pits in the roof of the mouth • Lined with olfactory epithelium • Forked tongue picks up scent particles and conveys them to this organ • Nonvenomous snakes • Kill prey by constriction or by biting and swallowing • Skin is infolded between scales • When stretched by a large meal, the skin is unfolded

45. Characteristics and Natural History of Reptilian Orders • Snakebite and Toxicity • Neurotoxins act on the nervous system, causing blindness or inhibit respiration • Hemorrhagin type venoms break down blood vessels Blood leaks into tissue spaces • Sea snakes and the Australian tiger snake have the most deadly venom per unit volume • Reproduction • Most are oviparous and lay shelled eggs, pit vipers are ovoviviparous, A few snakes are viviparous, having a primitive placenta to exchange nutrients with the young • Female snakes can store sperm and lay several clutches of fertile eggs long after a single mating

47. Characteristics and Natural History of Reptilian Orders The Tuatara • Only 2 living species in New Zealand represent this ancient lineage • Loss of the tuatara populations caused by human introduction of nonnative species which preyed upon the tuatara • Tuatara are vulnerable because they have slow growth and reproductive rates • Lizard-like and live in burrows • Slow growing and may live to 77 years of age • Well-developed median parietal eye buried beneath skin

49. Characteristics and Natural History of Reptilian Orders Crocodilia: Crocodiles and Alligators • Modern crocodilians differ little from primitive crocodilians of the early Mesozoic • There are 3 families of modern crocodilians • Alligators and caimans are found primarily in the New World and have a broader snout • Crocodiles are widely distributed • One species of gavial occurs in India and Burma and has a very narrow snout