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Amphibians to reptiles … Origins and evolution of the major groups… 1 2 Origins… Devonian 410 – 360 mya Carboniferous 360 – 286 mya Permian 286 – 245 mya ‘The Age of Fishes’ (& First Tetrapods) ‘The Age of the Giant Amphibians’ End of the Age of the Giant Amphibians 3 Vertebrate Cladogram 340 mya (Amniotic egg) (Four-limbed vertebrates) (Animals with backbones) Credit: Stanislav Traykov Jan 28, 2005 4 The amniotic egg Shared derived character of amniotes Possible only with internal fertilization Shell protects against desiccation Laying in terrestrial environments Allantois (waste mgmt, gas exchange) Reduces predation Chorion (encloses embryo, interface with mother) Amnion Amnion (protection) 5 Early amniotes Diverged ~ 340 mya Middle of Age of Amphibs Small and lizard-like Nails/claws for burrowing Skin relatively impervious to reduce water loss Enlarged lungs Amniotic egg Exploited the opportunity of aridity 6 Amniote Evolution Rhipidistian crossopterygian lobe-finned fish Synapsids Lepospondyls “Anapsids” Anthracosaurs Diapsids Euryapsids 7 Fundamental splits among the Amniotes Skull fenestrae are the synapomorphies that distinguish these groups Synapsids Diapsids Anapsids Euryapsids 8 Amniote Evolution Rhipidistian crossopterygian lobe-finned fish Synapsids Lepospondyls “Anapsids” Anthracosaurs Diapsids Lepidosaurs (tuatara & squamates) Mammalia Chelonia Euryapsids ? Archosaurs (birds & crocs) 9 Class Reptilia – 4 Orders Anapsids: Testudines (Turtles) (~300 spp) Diapsids: Squamata (Lizards and Snakes) (~7,900 spp) Crocodylia (Crocodylians) (23 spp) Rhynchocephalia (Tuatara)(2-3 spp) 10 Unifying characteristics Tetrapods - a few have secondarily lost their limbs (snakes and glass lizards) Shelled amniotic egg encases the embryo in a fluid-filled container frees reptiles from locating and using a moist habitat for egg laying Scales of epidermal origin made from keratin prevents water loss and slows dehydration rates more widespread than the amphibians, which are limited by temperature and moisture 11 Reptile skin …scales, scutes, and osteoderms Three-chambered heart Atria Ventricle All reptiles except crocodilians have a three-chambered heart Ridges still help oxygenated and deoxygenated blood stay separate = thermal flexibility can adjust the proportion of oxygenated blood that goes to the body versus the lungs (intracardiac blood shunt) Aids in heating and cooling 13 14 Other unifying characteristics All have internal fertilization accomplished by a copulatory organ Penis (turtles and crocs) Hemipenes (lizards and snakes): paired outpocketings of the cloaca Tuatara lack a copulatory organ - mate by repressing the cloaca Why? Timing with egg shelling 15 Amniote Evolution Rhipidistian crossopterygian lobe-finned fish Synapsids Lepospondyls “Anapsids” Anthracosaurs Diapsids Euryapsids 16 Subclass Anapsida stem reptiles Lack fenestrae - Ancestral condition Developed shells and attained huge sizes (one turtle reached 6 m in length) Eventually lead to the modern chelonians — ancient reptiles that have not changed much since the Triassic 251-199MYBP) Includes turtles and their extinct relatives 17 What are turtles? Because the anapsid condition is ancestral, it gives no clue to their relationship with extant reptiles Both morphological and molecular analyses now suggest that turtles may be nested within Diapsida This would mean that the anapsid skulls of turtles may be secondarily derived ? ? 18 Three hypotheses of turtle origins Lyson et al. (2013) Evolutionary origin of the turtle shell. Current Biology 23:1113-1119. Gastralia = dermal bones of ventral body wall still found in crocs – not associated with vertebra = became plastron? 20 Archelon, Late Cretaceous 21 Turtles (Order Chelonia or Testudines) = ~300 species Only living “anapsids” = one of earliest reptile lineages (little changed since the Triassic) Protostega from ~ 60 mya 22 Figure 1.5 The body forms of turtles reflect their habits and habitats 24 Turtles characteristics A bony shell completely encloses internal organs Characterized by a carapace, fused to the vertebrae and ribs, and joined to the plastron. Only head and tail free Outer layer = keratin / “living” 25 Shell-associated Demography Limbs, head, and neck can be drawn into shell Extraordinary armor Implications: great longevity, high survival. 26 Other characteristics No teeth, replaced by a keratinous beak as in birds. 27 Other characteristics All turtles are oviparous with longterm sperm storage. No parental care. Dig nests with hind feet on land 28 Other characteristics Highly mobile neck follows one of two patterns Hidden necks: Cryptodira (suborder) Side necks: Pleurodira (suborder) 29 Fang sheng – Buddhist practice of releasing captive animals, motivated by compassion Turtle remedies Distinct in public appreciation = birds Freshwater turtles and land tortoises are used for food and ancient medicinal remedies in China Thought to cure cancer and prevent aging Turtles in Native American mythology “…in the ancient days of creation, the world was all water and the only living creatures dwelt upon the back of a huge tortoise…Then crayfish obtained mud from the bottom of the ocean….From the mud the earth was built upon the back of the tortoise.” Turtles “ A well-known scientist (some say it was Bertrand Russell) once gave a public lecture on astronomy. He described how the earth orbits around the sun and how the sun, in turn, orbits around the center of a vast collection of stars called our galaxy. At the end of the lecture, a little old lady at the back of the room got up and said: "What you have told us is rubbish. The world is really a flat plate supported on the back of a giant tortoise." The scientist gave a superior smile before replying, "What is the tortoise standing on?" "You're very clever, young man, very clever," said the old lady. "But it's turtles all the way down!“ Stephen Hawking, A Brief History of Time Amniote Evolution Rhipidistian crossopterygian lobe-finned fish Synapsids Lepospondyls “Anapsids” Anthracosaurs Diapsids Euryapsids 34 Subclass Diapsida Most species-rich group of amniotes (>16,000 spp.) Includes most extant reptiles Enormous radiation of diapsids in the Mesozoic (245 – 65 mya) Only two groups are extant: Lepidosaurs: Lizards, snakes, tuatara and their extinct relatives Archosaurs: Crocodylians, birds, and their extinct relatives 35 Diapsida Subclass Lepidosauria (scaly lizards) (versus Archosauria – crocs and birds) Two living orders: Rhynchocephalia (Tuatara) Squamata (Lizards, snakes, amphisbaenids) 36 Subclass Diapsida Lepidosauria Defining characters Transverse cloacal slit Hemipenes Ecdysis 37 Hemipenes of squamates Reptile skin (vs amphibian skin) and shedding Subclass Diapsida / Lepidosauria Order: Rhynchocephalia Rhynchocephalians ("beak head”) or Tuatara (“spines on back”) Sister group to the Squamates 49 Why are Tuatara “so special”? Order “Rhynchocephalia” 51 Rhynchocephalians: Characteristics Lizard-like in body form Series of spines on nape and back tuatara = "spines on back” in Maori Chisel-beaked upper jaw overhanging the lower jaw. Teeth fused to the jaw and not replaced throughout life 52 Rhynchocephalians: Characteristics Vestigial eye (parietal eye) Has its own lens, cornea, retina with rod-like structures and degenerated nerve connection to the brain Possibly evolved from a real eye Used for navigation, thermoregulation, tracking photoperiod? 53 Tuatara Long-lived, incubation of a year or more, sexual maturity of a decade or more Do not copulate – rub cloacas together 54 Rhynchocephalians Once a widespread group, but now just 2-3 species of the genus Sphenodon on small islands off New Zealand. Most Rhynchocephalians went extinct 70 MYA 55 Figure 4.6 Rhynchocephalia Tuatara, the Keeper of the Knowledge, guards the trails of the mind and spirit 57 To here 58 Order “Rhynchocephalia” Subclass Lepidosauria Order “Squamata” Subclass Archosauria 59 Squamates; the “scaly reptiles” (Order Squamata) Most diverse group of reptiles (with > 7,900 species) Lizards [Suborder Lacertilia] Snakes [Suborders Serpentes or Ophidia] Amphisbaenians [Suborder Amphisbaenia] 60 Squamates – True lizards (Suborder: Lacertilia) • ~3,000 species, 25 families • Range in size from 0.03-3 m • Most are small (80% are < 2 g) and insectivorous, larger species are often herbivorous Size range: Sphaerodactylus ariasae Squamates – true lizards (Suborder: Lacertilia) External ear openings Moveable eyelids (unlike snakes) 63 Four limbs, but limb reduction or loss in some groups…all Lerista spp. (Aust. skinks) Squamates - Lizards (Suborder: Lacertilia) Most are oviparous, but some are viviparous Two species are venomous (Gila monster, Mexican beaded lizard) 65 caudal autotomy Intravertebral tail autotomy (> 50% families) 68 Unreliability: “She’s a chameleon…” Squamates – Snakes (Suborder: Ophidia, Serpentes) Squamates – Snakes (Suborder: Ophidia, Serpentes) ~2,300 species No limbs, eyelids or external ear (inner ear detects vibrations) Have lost the sternum – ribs extend the entire length of the vertebral column Left lung is reduced or absent, organs elongate 71 72 Snakes and feeding All are carnivorous Most take relatively large prey Subdue by venom and/or constriction Heat sensitive pits Skull and jaws exceedingly mobile 73 Python eating a wallaby 74 75 Many snakes have infrared-sensitive organs Many snakes have infraredsensitive organs When God learned of the Serpent’s role, he placed a curse on the snake… God cursed the snake "above every beast of the field" and commanded that "upon your belly you shall crawl, and dust you shall eat all the days of your life." (Gen 3:14) Roots in evolution In honor of Asclepius, a particular type of non-venomous snake was often used in ancient Greek healing rituals… https://en.wikipedia.org/wiki/Rod_of_Ascle pius Gadsden flag – revolutionary war origins Originally had 13 rattles Jefferson’s original Squamates – Amphisbaenids (Suborder: Amphisbaenia) Amphisbaena = “to go both ways” named after a mythical ant-eating serpent with a head at each end Move like accordions – backward or forward ~ 180 species Oviparous Morphology for burrowing Mostly legless – burrow, eat inverts Skin loose, scales in annuli 83 Prolific burrowers: 84 Bipes spp. – “Mexican mole lizards” or “ajolotes” 85 Subclass Diapsida Archosauria The ‘Ruling Reptiles’ Originated ~ 250 mya in the Permian Dominant terrestrial vertebrates in the Mesozoic Era Descendants include crocodiles, birds and dinosaurs 86 Subclass Diapsida Archosauria Modifications of the skeleton allowing for diverse locomotor specializations Adaptations associated with increased predatory efficiency Teeth set in sockets in the jaw bone Forelimbs with sharp claws 87 The Age of the Reptiles - Mesozoic 245 – 65 mya (Triassic, Jurassic, Cretaceous periods) Reptile-dominated trophic pyramid Explosive radiation of flowering plants, and the insects and small reptiles that fed on them Many medium- to large-sized semiaquatic herbivorous reptiles arose Immense biomass of insect- and plant-eating reptiles Dominated by carnivorous species 88 89 90 91 End of the age of Reptiles (~65 mya): The KT (Cretaceous-Tertiary) event Mass extinctions correlate with the occurrence of a 310-kmwide impact crater on the Yucatan Peninsula 70% of all species on earth went extinct 92 Legacies of the archosaurs…. The birds (~10,000 extant species: Class Aves or Neornithes) 93 Microraptor discovered 2003 Origins of flight Archaeopteryx First known bird ~250 mya 130 mya 94 Class Aves 95 Archosaurs also gave rise to the Crocodylia Originated ~200 mya in the Triassic Survive today in small numbers Only ~ 23 species 96 Order Crocodylia (crocodiles = 23 species – most are endangered) Survivors (along with birds) of the once-prevalent group Archosauria thrived during the Mesozoic 97 Figure 1.6 Crocodylians’ jaws provide information about their food habits Crocodylia Thecodont teeth (teeth set in bony sockets) Teeth are replaced as often as once a month 99 Crocodylia... Bodies armored by sheets of abutting osteoderms (plates of bone under the skin) and covered by thick non-overlapping scales 100 Crocodylia A very advanced Four-chambered heart Cogged valve allows blood to be shunted from the right side of the heart to the systemic circulation, introducing deoxygenated blood Can re-route blood usually pumped into the lungs into other parts of the body Allows crocodiles to dive for several hours without surfacing to breathe When blood rich with carbon dioxide goes to the stomach instead of the lungs, it can aid digestion 101 Secondary palate Allows crocodiles to breathe even when submerged under water Their internal nostrils open in the back of their throat, where a part of the tongue called the "palatal valve" closes off their respiratory system. This way they can open their mouths underwater. Most reptiles lack a secondary palate 102 Crocodylia... All build nests, either as mounds of rotting vegetation (alligators, caimans, and a few crocodiles) or in friable soils (most crocodiles) Parents guard nests and assist young 103 Crocodiles in mythology In Egypt the reptile was equated with the crocodile-headed "Sebek" who symbolized viscous passions, deceit, treachery and hypocrisy “crocodile tears” Amniote Evolution Rhipidistian crossopterygian lobe-finned fish Synapsids Mammalia Lepospondyls Anapsids Anthracosaurs Diapsids Euryapsids Lepidosaurs Chelonia? (tuatara &squamates) Archosaurs (birds & crocs) ? 105 Euryapsids With a single dorsal opening on skull Modified from the diapsid condition Polyphyletic group Highly successful and included many highly derived aquatic forms, but none extant Or are they? 106 Actually, one Euryapsid plesiosaur may still be extant… 107 Legend of Silver Lake 108 Subclass Synapsida Subclass Synapsida, with a single opening on the side of the skull The ‘proto-mammals’ or mammal-like reptiles (technically not reptiles) Originated 315 mya in the Carboniferous Eventually lead to the modern mammals Tetraceratops Oldest known therapsid (Permian) 109 110 First mammals The first mammals appeared only shortly after the early archosaurs (late Triassic). 1st 2/3 of their history, mammals were numerous but relatively insignificant Mostly nocturnal or arboreal shrew-like animals Did not expand into a wide variety of environments until after the extinction of the early archosaurs 111 112 113 Definitions of Reptilia Origins of reptiles End… 115 crocodile complaint # 303068 Alligator in the park might eat the kids Hey mister city manager city parks are for people Parks guy tells the manager No alligator in the park I looked myself It's a crocodile Crocodile in the park might eat the kids Call Channel 7 then tell the Mayor State biologist tells the city crocodiles are protected Here's a useful brochure "Living with crocodiles" - Philip Stoddardit's