Gnathostome Evolution

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Origins of Bones Mineralization originated with mouthparts The entire skeleton was mineralized much later Gnathostomes are vertebrates that have jaws Jaws evolved from skeletal supports of the pharyngeal slits Also have enhanced sensory systems, mineralized endoskeleton, and paired appendages 34.14 0.5 m Gill slits Cranium Mouth Skeletal rods 34.13 1 Gnathostome Evolution Placoderms - extinct lineage of armored vertebrates Acanthodians - radiated during the Devonian period (closely related to osteichthyans) (a) Coccosteus, a placoderm (b) Climatius, an acanthodian 2 Class Chondrichthyans Skeleton composed primarily of cartilage Cartilaginous skeleton evolved secondarily from an ancestral mineralized skeleton Reproductive tract excretory system and digestive tract empty into a common cloaca Includes: sharks, rays, and ratfish Dorsal fins Pectoral fins Pelvic fins (a) Blacktip reef shark (Carcharhinus melanopterus) (b) Southern stingray (Dasyatis americana) (c) Spotted ratfish (Hydrolagus colliei) 34.15 3

Ostiechthyes Nearly all have a bony skeleton We informally call them fish Control their buoyancy with an air sac known as a swim bladder Breath by drawing water over gills Located in chambers covered by a protective bony flap called the operculum Brain Spinal cord Swim bladder Dorsal fin Caudal fin Adipose fin Nostril Cut edge of operculum Anal fin Liver Gills Kidney Gonad Stomach Heart Anus Lateral line Pelvic fin Intestine 34.16 Urinary bladder 4 Ostiechthyes Yellowfin tuna (Thunnus albacares) Red lionfish (Pterois volitans) Class Common sea horse (Hippocampus ramulosus) Ray-finned fish fins supported by long flexible rays modified for maneuvering, defense, and other functions Class Sarcopterygii - have muscular, pectoral fins (ex. Coelacanths, lungfish, tetrapods) Fine-spotted moray eel (Gymnothorax dovii) 34.17 & 34.19 5 cm Lower jaw Tetrapods Scaly covering Dorsal spine 5 Fish Characters Scales Fins Gills and lungs Gnathostomes that have limbs and feet Tetrapod Characters Neck Ribs Fin skeleton Flat skull Eyes on top of skull Shoulder bones Ribs Neck Scales Head Eyes on top of skull One of the most significant events in vertebrate history was when the fin of lobefins evolved into the limbs and feet of tetrapods Ulna Elbow Have ears for detecting airborne sounds Wrist Radius Fin Fin skeleton Lungfishes Eusthenopteron Panderichthys Tiktaalik Fins became more limb-like Have four limbs and feet with digits Humerus Flat skull Acanthostega Limbs with digits Tulerpeton ns Amniotes Key to limb bones Ulna Radius Humerus 34.20 & 34.21 Silurian PALEOZOIC Permian Carboniferous Devonian 415 400 385 370 355 340 325 310 295 280 265 0 Time (millions of years ago) 6

Class 4,800 species Most have moist skin that complements the lungs in gas exchange (a) Order Urodela (salamanders) (b) Order Anura (frogs) Name means two lives - undergo metamorphosis from aquatic larva to terrestrial adult (c) (a) Tadpole Order Apoda (caecilians) Order Urodela (salamanders) - have tails (b) During metamorphosis Order Anura (frogs, toads) - lack tails Order Apoda (caecilians) - legless and resemble worms (c) Mating adults 34.20-34.21 7 Parareptiles Turtles ANCESTRAL AMNIOTE Reptiles Synapsids Diapsids Archosaurs Lepidosaurs Dinosaurs Saurischians Crocodilians Pterosaurs Ornithischian dinosaurs Saurischian dinosaurs other than birds Birds Plesiosaurs Ichthyosaurs Tuataras Squamates 34.25 8 Amniotes Tetrapods that have a terrestrially adapted egg Living members are reptiles, birds, and some mammals Named for the major derived character (amniotic egg) Amniotic egg contains specialized membranes that protect the embryo Also have relatively impermeable skin and the ability to use the rib cage to ventilate the lungs Appeared in the Carboniferous period Embryo Amniotic cavity with amniotic fluid Shell Amnion Extraembryonic membranes Allantois Chorion Yolk sac Yolk (nutrients) Albumen 34.26 9

Reptiles Includes tuatara, lizards, snakes, turtles, crocodilians, birds, and dinosaurs Have scales that create a waterproof barrier Lay shelled eggs on land Most are ectothermic - absorb external heat as main source of body heat (Birds are endothermic - keep body warm through metabolism) Date back to about 300 million years ago First major group were parareptiles (large, stocky herbivores) As parareptiles dwindled, diapsids were diversifying (lepidosaurs and archosaurs) 34.28 One surviving lineage of lepidosaurs is the t wo species lizard-like reptiles called tuatara The other lineage are lizards and snakes 10 Reptiles Snakes - legless lepidosaurs (a) Tuatara (Sphenodon punctatus) Turtles - have boxlike shells with upper and lower shields that fuse the the vertebrae, clavicles, and ribs 34.27 (b) Australian thorny devil lizard (Moloch horridus) Crocodilians - belong to the archosaur lineage that dates back to the Triassic (c) Wagler s pit viper (Tropidolaemus wagleri) (e) American alligator (Alligator mississippiensis) (d) Eastern box turtle (Terrapene carolina carolina) Birds - archosaurs (almost every reptilian feature has undergone modification in adaptation to flight 11 Birds Finger 1 Most obvious adaptations are wings and feathers (b) Bone structure Forearm Probably descended from theropods (group of small, carnivorous dinosaurs) 150 millions years ago - Arhaeopteryx (oldest known bird) Order Struthioniforms (flightless birds) Palm Finger 2 (a) Wing Vane Toothed beak Airfoil wing with contour feathers Wrist Finger 3 Shaft Barb Barbule Hook (c) Feather structure Shaft Wing claw Long tail with many vertebrae Demands of flight have caused a general body form to evolve Foot structure also shows variation 34.30-34.34 12

Amniotes that have hair and produce milk Generally have larger brains than other vertebrates of equivalent size 34.32 Evolved from synapsids in the late Triassic Biarmosuchus, a synapsid Jaw was remodeled (t wo bones that were part of the jaw joint were incorporated into the middle ear Key Temporal fenestra Articular Quadrate Dentary Squamosal Jaw joint (a) Articular and quadrate bones in the jaw Middle ear Stapes Eardrum Living lineages originated in the Jurassic but did not undergo significant adaptive radiation until after the Cretaceous Inner ear Eardrum Middle ear Inner ear Stapes Sound Incus (quadrate) Sound Malleus (articular) Present-day reptile Present-day mammal 13 (b) Articular and quadrate bones in the middle ear Monotremes - small group of egg-laying mammals (echidnas and platypus) Marsupials opossums, kangaroos, and koalas Born very early in its development Eutherian mammals Marsupial mammals Plantigale Deer mouse Mole Marsupial mole Sugar glider Flying squirrel (a) A young brushtail possum Wombat In some the marsupium opens in the rear Woodchuck Tasmanian devil Kangaroo Wolverine Patagonian cavy (b) Long-nosed bandicoot 34.3834.40 14 Eutherians placental mammals Have a longer period of pregnancy (compared to marsupials) Young eutherians complete their embryonic development within a uterus joined to the mother by a placenta 34.41 Orders and Examples Monotremata Platypuses, echidnas Main Characteristics Orders and Examples Lay eggs; no nipples; young suck milk from fur of mother Marsupialia Kangaroos, opossums, koalas Long, muscular trunk; thick, loose skin; upper incisors elongated as tusks Tubulidentata Aardvarks Aquatic; finlike forelimbs and no hind limbs; herbivorous Hyracoidea Hyraxes Echidna Proboscidea Elephants Koala Teeth consisting of many thin tubes cemented together; eats ants and termites Aardvark African elephant Sirenia Manatees, dugongs Manatee Xenarthra Sloths, anteaters, armadillos Tamandua Lagomorpha Rabbits, hares, picas Jackrabbit Carnivora Dogs, wolves, bears, cats, weasels, otters, seals, walruses Rock hyrax Short legs; stumpy tail; herbivorous; complex, multichambered stomach Reduced teeth or no teeth; herbivorous (sloths) or carnivorous (anteaters, armadillos) Rodentia Squirrels, beavers, rats, porcupines, mice Chisel-like incisors; hind legs longer than forelegs and adapted for running and jumping; herbivorous Primates Lemurs, monkeys, chimpanzees, gorillas, Golden lion humans tamarin Sharp, pointed canine teeth and molars for shearing; carnivorous Perissodactyla Hooves with an odd Horses, zebras, number of toes on tapirs, each foot; herbivorous rhinoceroses Indian rhinoceros Hooves with an even number of toes on each foot; herbivorous Chiroptera Bats Coyote Cetartiodactyla Artiodactyls Sheep, pigs, cattle, deer, Bighorn sheep giraffes Cetaceans Whales, dolphins, porpoises Pacific whitesided porpoise Main Characteristics Completes embryonic development in pouch on mother s body Red squirrel Frog-eating bat Aquatic; streamlined body; paddle-like fore-limbs and no hind limbs; thick layer of insulating blubber; carnivorous Eulipotyphla Core insectivores : some moles, some shrews Star-nosed mole Chisel-like, continuously growing incisors worn down by gnawing; herbivorous Opposable thumbs; forward-facing eyes; well-developed cerebral cortex; omnivorous Adapted for flight; broad skinfold that extends from elongated fingers to body and legs; carnivorous or herbivorous Eat mainly insects and other small invertebrates 15

34.43-34.44 Primates - include lemurs, tarsiers, monkeys, and apes Hands and feet adapted for grasping Large brain and short jaw Forward-looking eyes close together on the face providing depth perception Well-developed parental care and complex social behavior Opposable thumbs Oldest known anthropoid fossils are from about 45 million years ago ANCESTRAL PRIMATE 60 50 40 30 20 10 0 (a) New World monkey: spider monkey Time (millions of years ago) Lemurs, lorises, and bush babies Tarsiers New World monkeys Old World monkeys Gibbons Orangutans Gorillas Chimpanzees and bonobos Humans (b) Old World monkey: macaque Anthropoids 16 Evolution of Primates New and Old World monkeys underwent separate adaptive radiations during millions of years of separation The other group of anthropoids are hominoids (consist of primates called apes) Hominoids diverged from Old World monkeys about 20-25 million years ago (a) Gibbon (d) Chimpanzees 34.39-34.40 (b) Orangutan (c) Gorilla (e) Bonobos 17 Derived Characters of Hominins Bipedal hominoids with a large brain Language capabilities Symbolic thought Manufacture and use of complex tools Shortened jaw Humans and chimpanzee genomes are more than 99% identical Millions of years ago 0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0 4.5 5.0 5.5 6.0 6.5 7.0 anamensis Kenyanthropus platyops africanus Orrorin tugensis Paranthropus boisei garhi afarensis Ardipithecus ramidus Sahelanthropus tchadensis Paranthropus robustus habilis ergaster rudolfensis? neanderthalensis erectus sapiens 34.46 18

Hominids Study of human origins is paleoanthropology 20 species of distinct hominins have been discovered Originated in Africa 6-7 million years ago Australopiths lived between 4 and 2 million years ago 34.47 & 34.48 Early hominids had a small brain Two common misconceptions: Often thought of as chimpanzees Thinking of human evolution as a ladder leading to sapiens Began to walk on two legs for long distances about 1.9 million years ago Began to use tools (cut marks on animal bones) about 2.5 million years ago (a) The Laetoli footprints (b) Artist s reconstruction of A. afarensis 19 Hominids habilis - 2.4 to 1.6 million years ago Stone tools have been found with them ergaster - 1.9 to 1.6 million years ago First fully bipedal, large brained hominid erectus - about 1.8 million years ago First hominid to leave Africa neanderthalensis - 200,000 to 30,000 years ago Lived in Europe and the Near East Large, thick browed Became extinct a few thousand years after sapiens arrived in Europe sapiens - 160,000 years ago Oldest fossils outside of Africa about 50,000 years ago 34.49 & 34.50 20