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Early Tetrapods & Modern Amphibians Chapter 25 I. Movement Onto Land • Amphibians are vertebrate transition to land; other organisms like plants, gastropods, and arthropods made transition earlier • Since organisms made mostly of water, dangerous transition • Also had to adapt to different oxygen content, density, temperature regulation, and habitat diversity II. Evolution of Terrestrial Vertebrates • A. Devonian Origin • Mild temperatures, and periods of flooding or droughts creating unstable sources of freshwater • Fish with lungs were better able to survive • Early fish fossils that could crawl along mud with “walking fins:” Eusthenopteron, Acanthostega, and Ichthyostega. • Land adaptations include: skull, teeth, stronger pectoral and pelvic girdles, jointed limbs, stronger backbone, muscles to support body in air and elevate head, more protective rib cage, ear structure, and longer snout • Many fossils have more than 5 digits B. Carboniferous Radiation • Uniformly warm and wet • Tetrapods radiated in swampy, fern areas eating insects, larvae, and invertebrates • Temnospondyls form lineage from which modern amphibians are derived; they have 4 digits on forelimbs • Became better adapted to aquatic life; bodies flattened, some like salamanders developed weaker limbs and stronger tails, and frogs developed webbing on hind limbs for better swimming III. Modern Amphibians (Class Amphibia) • • • • A. Diversity Over 4200 species Ears redesigned Remain tied to water since eggs are laid in water and larvae have gills • Thin skin loses moisture rapidly restricting them to moist habitats • Ectothermic which also restricts habitat and range B. Caecilians: Order Gymnophiona • 160 species live in tropical rainforests of South America, Africa, and Southeast Asia • Elongate, limbless, and burrowing • Some larvae develop in folds of body and in others develop in oviduct, eating it for nourishment C. Salamanders: Order Caudata • 1. Characteristics • 360 species living in northern temperate areas • Most are small, under 15 cm, but Japanese giant salamander is 1.5 m long • Limbs are at right angles of body, with fore and hind limbs of equal length • Burrowing and some aquatic species have lost limbs • Carnivorous, eating high fat and protein foods so do not store much fat or glycogen Salamanders 2. Breeding Behavior • Fertilize eggs internally • Terrestrial species deposit egg clusters under logs or in soft earth; eggs hatch out to mini adults • Most salamanders lay strings of eggs in water; larvae hatch with gills, and then may turn into terrestrial or aquatic adults • Newts have “red eft” stage with a terrestrial juvenile, that then turns into aquatic, breeding adult • Some newts stay entirely aquatic 3. Respiration • Thin skin easily exchanges gases; also use mouth cavity • At various stages may also have gills, lungs, both gills and lungs, or neither • If lungs are used, they are present from birth, but only become functional following metamorphosis; hold nostrils above water to breathe • Larvae hatch with gills, and lose them following metamorphosis, along with fin-like tail; if a lineage does not undergo metamorphosis, it retains these characteristics • Those in terrestrial family Plethodontidae have no lungs and use only skin 4. Paedomorphosis • Preservation of larval features into adulthood • Trend found in salamanders • Some never metamorphose, like the mudpuppy • Others typically may not, but can change if conditions change (typically in dry conditions) • Another example is Ambystoma tigrinum, which stays in an axolotyl stage; can metamorphose if treated with thyroid hormone • Some have partial paedomorphsis, retaining larval characteristics but switching to lungs D. Frogs and Toads: Order Anura • • • • • • • • 1. Characteristics 3450 species Evolved during Jurassic Period, 150 mya All have tailed larval stage but are tailless as adults; none retain larval characteristics as adults 21 families Family Ranidae, larger frogs of North America Family Hylidae, tree frogs Family Bufonidae, toads 2. Habitat and Distribution • 260 species of genus Rana found in temperate and tropical areas • Frogs and toads are found in damp forested floors, although in tropical rainforests may live entire life in 1 tree • Swamps, ponds, streams • Frogs and toads are declining worldwide and becoming patchy in distribution; cause is unknown 3. Life Cycle • Solitary except during breeding season • Spend breeding season swimming around in water • During winter, burrow into mud to hibernate, using energy from stored fat and glycogen • Also accumulate glucose and glycerol in tissues to create an “antifreeze” to prevent tissues from forming ice crystals • Easy prey; protects themselves by concealment, poison glands, and aggression 4. Integument and Coloration • Thin, moist, and attached to body in several points • Epidermis contains keratin; this is thicker in more terrestrial ones • Inner dermal layer has mucous glands, which secrete waterproofing agents, and serous glands, which secrete poisons • Dendrobatid frogs from South America secrete highly toxic poisons • Chromatophores are pigment cells that produce skin color • Xanthophores are upper pigments with yellow, orange, and red • Iridophores are middle layer with silvery light-reflecting pigment that gives iridescent quality • Melanophores are deepest layer with brown or black melanin • Green color is produced by interactions among all of these pigments • Frogs can adjust color to camouflage themselves Skin Pigmentation 5. Skeletal and Muscular Systems • Well developed endoskeleton of bone and cartilage with changes to allow for jumping and swimming • Front of skull is light weight and flattened • Limbs have 3 joints: hip, knee, ankle • Foot has 5 rays on hind limb, and 4 on front limb; digits jointed 6. Respiration • Use skin, mouth, and lungs with skin being critical during hibernation • CO2 is mainly lost across skin whereas O2 is mainly absorbed through lungs • The movement of air into lungs is somewhat passive, depending on movement of throat 7. Vocalization • As air enters and leaves lungs, passes over vocal cords, on way to vocal sacs • Both males and females have vocal cords but males have a more developed larynx • Songs are unique and characteristic of the species http://www.naturesound.com/frogs/frogs.html 8. Circulation • 6th aortic arch present in gills was converted into pulmonary artery to lungs • Frog heart has 2 atria and 1 single ventricle • Blood from body enters right atrium and blood from lungs enters left atrium; both deoxygenated and oxygenated blood mix in ventricle • Valves do control blood though so mainly deoxygenated blood goes to lungs and oxygenated goes to body 9. Feeding and Digestion • Carnivorous • Catch prey with tongue that is hinged at front of mouth • Free end is glandular with sticky secretions that cause prey to adhere • Teeth are used to hold prey, not to chew or bite • Digestive tract produces enzymes to break down carbohydrates, proteins, and fats • Tadpoles are herbivorous and have longer tracts to digest plant material 10. Nervous System and Senses • a. Brain • Consists of 3 parts: forebrain (sense of smell), midbrain (vision), and hindbrain (hearing and balance) • Forebrain has cerebrum, but not really used. Olfactory lobe most important part • Midbrain has optic lobe • Hindbrain has anterior cerebellum (not used) and posterior medulla which controls auditory reflexes, respiration, swallowing, and circulatory system b. Vision • Dominant sense in many amphibians (except caecilians) • Eyelids keep eyes moist, protected, and free of dust; upper is fixed and lower has clear nictitating membrane • Cornea and lens bend light to focus image on retina, with both rods and cones for color vision • Iris changes to adjust to different light levels • At rest, frog focuses on distant objects c. Other Senses • Pressure sensitive lateral line is only found in larvae and aquatic adults • Ear is sensitive to airborne sounds; tympanic membrane passes vibrations to structure similar to cochlea • Chemical receptors in skin, on tongue, and in nasal cavity 11. Reproduction and Development • a. Egg • In spring, males migrate to same pond or stream and call females • This is dependent on temperature, humidity, and hormonal changes • Male clasps female in water and as she releases eggs, male discharges sperm over them • Eggs usually laid in masses • Eggs absorb water and swell; development begins immediately • Tadpoles hatch in 6-9 days Frog Life Cycle b. Tadpoles • Tadpole has horny jaws for grazing and a ventral adhesive disc for clinging to objects • 3 pairs of external gills develop into internal gills covered by flap on right side fused to body wall and a spiracle on left side • Water flows through mouth, over gills, and out spiracle c. Metamorphosis • Hind legs appear first; forelimbs are temporarily hidden by operculum • Tail is resorbed • Intestine becomes shorter • Mouth transforms into adult version • Lungs develop and gills resorbed Metamorphosis