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LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 34 The Origin and Evolution of Vertebrates Lectures by Erin Barley Kathleen Fitzpatrick © 2011 Pearson Education, Inc. Figure 34.1 • One lineage of vertebrates colonized land 365 million years ago • There are about 52,000 species of vertebrates, including the largest organisms ever to live on the Earth (how many beetles were there?) • Theme: Vertebrates have great disparity, a wide range of differences within the group – WHY? © 2011 Pearson Education, Inc. Concept 34.1: Chordates have a notochord and a dorsal, hollow nerve cord • Chordates (phylum Chordata) are bilaterian animals that belong to the clade of animals known as Deuterostomia • Chordates comprise all vertebrates and two groups of invertebrates, the urochordates and cephalochordates © 2011 Pearson Education, Inc. Figure 34.2 We will explore 11 clades of the phylum Chordata Echinodermata Chordates Cephalochordata ANCESTRAL DEUTEROSTOME Urochordata Notochord Craniates Vertebrates Gnathostomes Osteichthyans Lobe-fins Myxini Common ancestor of chordates Head Petromyzontida Chondrichthyes Vertebral column Actinopterygii Jaws, mineralized skeleton Actinistia Lungs or lung derivatives Dipnoi Lobed fins Reptilia Limbs with digits Amniotic egg Mammalia Milk What do you notice that is odd? Tetrapods Amniotes Amphibia Figure 34.3 4 Derived Characters of Chordates Dorsal, hollow nerve cord Muscle segments Notochord Mouth Anus Muscular, post-anal tail Pharyngeal slits or clefts Figure 34.4 Cephalochordata Cirri Mouth Pharyngeal slits Atrium Notochord 1 cm Digestive tract Atriopore Dorsal, hollow nerve cord Segmental muscles Anus Tail • Lancelets (Cephalochordata) are named for their bladelike shape • They are marine suspension feeders that retain characteristics of the chordate body plan as adults Figure 34.5 Urochordata Notochord Water flow Dorsal, hollow nerve cord Tail Muscle segments Excurrent siphon Incurrent siphon Intestine Stomach Atrium Pharynx with slits (a) Tunicate larva Incurrent siphon to mouth Excurrent siphon Excurrent siphon Anus Intestine Atrium Pharynx with numerous slits Tunic Esophagus Stomach (b) Adult tunicate (c) Adult tunicate • Tunicates (Urochordata) are more closely related to other chordates than are lancelets • When attacked, tunicates, or “sea squirts,” shoot water through their excurrent siphon Early Chordate Evolution • Ancestral chordates may have resembled lancelets • The same Hox genes that organize the vertebrate brain are expressed in the lancelet’s simple nerve cord tip • Genome sequencing suggests that – Genes associated with the heart and thyroid are common to all chordates – Genes associated with transmission of nerve impulses are unique to vertebrates © 2011 Pearson Education, Inc. Figure 34.8 5 mm Segmented muscles Pharyngeal slits Figure 34.9 Hagfishes Slime glands Hagfishes • The most basal group of craniates is Myxini, the hagfishes • Hagfishes have a cartilaginous skull and axial rod of cartilage derived from the notochord, but lack jaws and vertebrae • They have a small brain, eyes, ears, and tooth-like formations • Hagfishes are marine; most are bottom-dwelling scavengers © 2011 Pearson Education, Inc. Figure 34.10 Lampreys Lampreys • Lampreys (Petromyzontida) represent the oldest living lineage of vertebrates • They are jawless vertebrates that feed by clamping their mouth onto a live fish, suck blood • They inhabit various marine and freshwater habitats • They have cartilaginous segments surrounding the notochord and arching partly over the nerve cord © 2011 Pearson Education, Inc. Figure 34.11 Conodonts were the first vertebrates with mineralized skeletal elements Dental elements (within head) Concept 34.4: Gnathostomes are vertebrates that have jaws • Today, jawed vertebrates, or gnathostomes, outnumber jawless vertebrates • Gnathostomes include sharks and their relatives, ray-finned fishes, lobe-finned fishes, amphibians, reptiles (including birds), and mammals © 2011 Pearson Education, Inc. Figure 34.13 Gill slits Cranium Mouth Skeletal rods Figure 34.14 Fossil of an early gnathostome. 0.5 m Figure 34.15 Dorsal fins Chondrichthyans (Sharks, Rays, and Their Relatives) Pectoral Pelvic fins fins (a) Blacktip reef shark (Carcharhinus melanopterus) (b) Southern stingray (Dasyatis americana) (c) Spotted ratfish (Hydrolagus colliei) Chondrichthyans (Sharks, Rays, and Their Relatives) • Chondrichthyans (Chondrichthyes) have a skeleton composed primarily of cartilage • The largest and most diverse group of chondrichthyans includes the sharks, rays, and skates © 2011 Pearson Education, Inc. • The reproductive tract, excretory system, and digestive tract empty into a common cloaca • Shark eggs are fertilized internally but embryos can develop in different ways – Oviparous: Eggs hatch outside the mother’s body – Ovoviviparous: The embryo develops within the uterus and is nourished by the egg yolk – Viviparous: The embryo develops within the uterus and is nourished through a yolk sac placenta from the mother’s blood © 2011 Pearson Education, Inc. Video: Shark Eating Seal © 2011 Pearson Education, Inc. Video: Manta Ray © 2011 Pearson Education, Inc. Ray-Finned Fishes and Lobe-Fins • The vast majority of vertebrates belong to a clade of gnathostomes called Osteichthyes • Osteichthyans include the bony fish and tetrapods – Aquatic osteichthyans are the vertebrates we informally call fishes © 2011 Pearson Education, Inc. Figure 34.16 Spinal cord Swim bladder Dorsal fin Adipose fin Brain Caudal fin Nostril Cut edge of operculum Anal fin Liver Gills Anus Stomach Gonad Kidney Intestine Heart Pelvic fin Lateral line Urinary bladder Ray-Finned Fishes • Actinopterygii, the ray-finned fishes, include nearly all the familiar aquatic osteichthyans • Ray-finned fishes originated during the Silurian period (444 to 416 million years ago) • The fins, supported mainly by long, flexible rays, are modified for maneuvering, defense, and other functions © 2011 Pearson Education, Inc. Video: Clownfish and Anemone © 2011 Pearson Education, Inc. Video: Coral Reef © 2011 Pearson Education, Inc. Video: Seahorse Camouflage © 2011 Pearson Education, Inc. Figure 34.17 Yellowfin tuna (Thunnus albacares) Red lionfish (Pterois volitans) Common sea horse (Hippocampus ramulosus) Fine-spotted moray eel (Gymnothorax dovii) Figure 34.18 Lobe-Fins 5 cm Lower jaw Scaly covering Dorsal spine • The lobe-fins (Sarcopterygii) have muscular pelvic and pectoral fins • Lobe-fins also originated in the Silurian period Figure 34.19 Concept 34.5: Tetrapods are gnathostomes that have limbs • One of the most significant events in vertebrate history was when the fins of some lobe-fins evolved into the limbs and feet of tetrapods © 2011 Pearson Education, Inc. Derived Characters of Tetrapods • Tetrapods have some specific adaptations – Four limbs, and feet with digits – A neck, which allows separate movement of the head – Fusion of the pelvic girdle to the backbone – The absence of gills (except some aquatic species) – Ears for detecting airborne sounds © 2011 Pearson Education, Inc. Figure 34.20 Fish Characters Scales Fins Gills and lungs Tetrapod Characters Neck Ribs Fin skeleton Flat skull Eyes on top of skull Shoulder bones Ribs Neck Scales Head Eyes on top of skull Humerus Ulna Flat skull Elbow Radius Fin “Wrist” Fin skeleton Figure 34.21 Lungfishes Eusthenopteron Panderichthys Tiktaalik Acanthostega Limbs with digits Tulerpeton Amphibians Amniotes Silurian PALEOZOIC Permian Carboniferous Devonian 415 400 385 370 355 340 325 310 295 280 265 0 Time (millions of years ago) Key to limb bones Ulna Radius Humerus Figure 34.22 Amphibians • Amphibians (class Amphibia) are represented by about 6,150 species • Order Urodela includes salamanders, which have tails (a) Order Urodela (salamanders) (b) Order Anura (frogs) (c) Order Apoda (caecilians) Figure 34.23 (a) Tadpole (b) During metamorphosis (c) Mating adults • Amphibian means “both ways of life,” referring to the metamorphosis of an aquatic larva into a terrestrial adult • Most amphibians have moist skin that complements the lungs in gas exchange • Amphibian populations have been declining in recent decades…why are they so vulnerable? © 2011 Pearson Education, Inc. • Fertilization is external in most species, and the eggs require a moist environment • In some species, males or females care for the eggs on their back, in their mouth, or in their stomach © 2011 Pearson Education, Inc. Concept 34.6: Amniotes are tetrapods that have a terrestrially adapted egg • Amniotes are a group of tetrapods whose living members are the reptiles, including birds, and mammals © 2011 Pearson Education, Inc. Figure 34.25 Parareptiles Turtles Reptiles Archosaurs Crocodilians Pterosaurs Saurischians Dinosaurs Diapsids Ornithischian dinosaurs Saurischian dinosaurs other than birds Birds Plesiosaurs ANCESTRAL AMNIOTE Ichthyosaurs Synapsids Lepidosaurs Tuataras Squamates Mammals Figure 34.26 Extraembryonic membranes Allantois Amnion Chorion Yolk sac Embryo Amniotic cavity with amniotic fluid Shell Yolk (nutrients) Albumen Reptiles • The reptile clade includes the tuataras, lizards, snakes, turtles, crocodilians, birds, and some extinct groups • Reptiles have scales that create a waterproof barrier • Most reptiles lay shelled eggs on land © 2011 Pearson Education, Inc. • Most reptiles are ectothermic, absorbing external heat as the main source of body heat • Birds are endothermic, capable of keeping the body warm through metabolism © 2011 Pearson Education, Inc. Figure 34.29 (a) Tuatara (Sphenodon punctatus) (b) Australian thorny devil lizard (Moloch horridus) (c) Wagler’s pit viper (Tropidolaemus wagleri) (e) American alligator (Alligator mississippiensis) (d) Eastern box turtle (Terrapene carolina carolina) Birds • Birds are archosaurs, but almost every feature of their reptilian anatomy has undergone modification in their adaptation to flight © 2011 Pearson Education, Inc. Derived Characters of Birds • Many characters of birds are adaptations that facilitate flight • The major adaptation is wings with keratin feathers • Other adaptations include lack of a urinary bladder, females with only one ovary, small gonads, and loss of teeth © 2011 Pearson Education, Inc. Figure 34.30 Finger 1 (b) Bone structure Palm Finger 2 (a) Wing Forearm Shaft Vane Wrist Finger 3 Shaft Barb Barbule Hook (c) Feather structure Figure 34.31 Toothed beak Airfoil wing with contour feathers Wing claw Long tail with many vertebrae • The demands of flight have rendered the general body form of many flying birds similar to one another Video: Flapping Geese © 2011 Pearson Education, Inc. Video: Soaring Hawk © 2011 Pearson Education, Inc. Video: Swans Taking Flight © 2011 Pearson Education, Inc. Figure 34.33 Behavior and morphology has adapted to fulfill distinct niches Figure 34.34 Figure 34.35 Figure 34.36 Concept 34.7: Mammals are amniotes that have hair and produce milk • Mammals, class Mammalia, are represented by more than 5,300 species • Derived characters of mammals: – – – – Mammary glands, which produce milk Hair A high metabolic rate, due to endothermy A larger brain than other vertebrates of equivalent size – Differentiated teeth • © 2011 Pearson Education, Inc. Figure 34.37 Biarmosuchus, a synapsid Key Temporal fenestra Articular Quadrate Dentary Squamosal Jaw joint (a) Articular and quadrate bones in the jaw Middle ear Stapes Eardrum Inner ear Eardrum Middle ear Inner ear Stapes Sound Sound Incus (quadrate) Malleus (articular) Present-day reptile Present-day mammal (b) Articular and quadrate bones in the middle ear Figure 34.38 Monotremes • Monotremes are a small group of egg-laying mammals consisting of echidnas and the platypus Marsupials • Marsupials include opossums, kangaroos, and koalas • The embryo develops within a placenta in the mother’s uterus • A marsupial is born very early in its development • It completes its embryonic development while nursing in a maternal pouch called a marsupium © 2011 Pearson Education, Inc. Figure 34.39 (a) A young brushtail possum (b) Long-nosed bandicoot Marsupial mammals Figure 34.40 Convergent evolution of marsupials and eutherians (placental mammals). Plantigale Eutherian mammals Deer mouse Mole Marsupial mole Sugar glider Flying squirrel Wombat Tasmanian devil Kangaroo Woodchuck Wolverine Patagonian cavy ANCESTRAL MAMMAL Monotremes Marsupials (324 species) (5 species) Figure 34.41a Monotremata Marsupialia Eutherians (5,010 species) Proboscidea Sirenia Tubulidentata Hyracoidea Afrosoricida Macroscelidea Xenarthra Rodentia Lagomorpha Primates Dermoptera Scandentia Carnivora Cetartiodactyla Perissodactyla Chiroptera Eulipotyphia Pholidota Figure 34.41b Orders and Examples Main Characteristics Lay eggs; no nipples; young suck milk from fur of mother Monotremata Platypuses, echidnas Orders and Examples Completes embryonic development in pouch on mother’s body Marsupialia Kangaroos, opossums, koalas Echidna Proboscidea Elephants Koala Long, muscular trunk; thick, loose skin; upper incisors elongated as tusks Tubulidentata Aardvarks Teeth consisting of many thin tubes cemented together; eats ants and termites Aardvark African elephant Sirenia Manatees, dugongs Aquatic; finlike forelimbs and no hind limbs; herbivorous Hyracoidea Hyraxes 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 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 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 Star-nosed mole Video: Bat Licking Nectar © 2011 Pearson Education, Inc. Video: Bat Pollinating Agave Plant © 2011 Pearson Education, Inc. Video: Galápagos Sea Lion © 2011 Pearson Education, Inc. Video: Wolf Agonistic Behavior © 2011 Pearson Education, Inc. Derived Characters of Primates – – – – Hands, feet for grasping Flat nails A large brain and short jaws Forward-looking eyes close together on the face, providing depth perception – Complex social behavior and parental care – A fully opposable thumb (in monkeys and apes) © 2011 Pearson Education, Inc. Living Primates • There are three main groups of living primates – Lemurs, lorises, and pottos – Tarsiers – Anthropoids (monkeys and apes) © 2011 Pearson Education, Inc. Figure 34.42 Figure 34.43 Lemurs, lorises, and bush babies Tarsiers ANCESTRAL PRIMATE Old World monkeys Gibbons Orangutans Gorillas Chimpanzees and bonobos Humans 60 50 20 30 40 Time (millions of years ago) 10 0 Anthropoids New World monkeys Figure 34.44 (a) New World monkey: spider monkey with prehensile tail (b) Old World monkey: macaque Video: Gibbons Brachiating © 2011 Pearson Education, Inc. Video: Chimp Agonistic Behavior © 2011 Pearson Education, Inc. Video: Chimp Cracking Nut © 2011 Pearson Education, Inc. Figure 34.45 (a) Gibbon (b) Orangutan (c) Gorilla (d) Chimpanzees (e) Bonobos Concept 34.8: Humans are mammals that have a large brain and bipedal locomotion • The species Homo sapiens is about 200,000 years old, which is very young, considering that life has existed on Earth for at least 3.5 billion years © 2011 Pearson Education, Inc. Derived Characters of Humans • A number of characters distinguish humans from other apes – Upright posture and bipedal locomotion – Larger brains capable of language, symbolic thought, artistic expression, the manufacture and use of complex tools – Reduced jawbones and jaw muscles – Shorter digestive tract © 2011 Pearson Education, Inc. • The human and chimpanzee genomes are 99% identical • How can we be this close, yet so different? © 2011 Pearson Education, Inc. Figure 34.46 Paranthropus robustus 0 ? Paranthropus boisei 0.5 Homo Homo neanderthalensis sapiens Homo ergaster 1.0 Australopithecus africanus Millions of years ago 1.5 2.0 2.5 Kenyanthropus platyops Australopithecus garhi Australo3.0 pithecus anamensis 3.5 Homo habilis 4.0 4.5 6.0 6.5 7.0 Ardipithecus ramidus Orrorin tugensis Sahelanthropus tchadensis Homo rudolfensis • Hominins originated in Africa about 6–7 million years ago • Early hominins show evidence of small brains and increasing bipedalism Australopithecus afarensis 5.0 5.5 Homo erectus Figure 34.47: Ardi, 4.4 million years old • Misconception: Early hominins were chimpanzees – Correction: Hominins and chimpanzees shared a common ancestor • Misconception: Human evolution is like a ladder leading directly to Homo sapiens – Correction: Hominin evolution included many branches or coexisting species, though only humans survive today © 2011 Pearson Education, Inc. Australopiths • Australopiths are a paraphyletic assemblage of hominins living between 4 and 2 million years ago • Some species, such as Australopithecus afarensis walked fully erect © 2011 Pearson Education, Inc. Figure 34.48 Evidence that hominins walked upright 3.5 million years ago. (a) The Laetoli footprints (b) Artist’s reconstruction of A. afarensis • Homo erectus originated in Africa by 1.8 million years ago • It was the first hominin to leave Africa © 2011 Pearson Education, Inc. Neanderthals • Neanderthals, Homo neanderthalensis, lived in Europe and the Near East from 350,000 to 28,000 years ago • They were thick-boned with a larger brain, they buried their dead, and they made hunting tools • Debate is ongoing about the extent to which genetic material was exchanged between neanderthals and Homo sapiens © 2011 Pearson Education, Inc. Figure 34.50 EXPERIMENT Hypothesis: Neanderthals gave rise to European humans. Expected phylogeny: Chimpanzees Neanderthals Living Europeans Other living humans RESULTS Chimpanzees Neanderthal 1 Neanderthal 2 European and other living humans Homo Sapiens • Homo sapiens appeared in Africa by 195,000 years ago • All living humans are descended from these African ancestors © 2011 Pearson Education, Inc. Figure 34.51 • The oldest fossils of Homo sapiens outside Africa date back about 115,000 years and are from the Middle East • Humans first arrived in the New World sometime before 15,000 years ago • In 2004, 18,000-year-old fossils were found in Indonesia, and a new small hominin was named: Homo floresiensis © 2011 Pearson Education, Inc. • Homo sapiens were the first group to show evidence of symbolic and sophisticated thought • In 2002, a 77,000-year-old artistic carving was found in South Africa © 2011 Pearson Education, Inc. Amniotes: amniotic egg, rib cage ventilation Lobe-fins: muscular fins or limbs Tetrapods: four limbs, neck, fused pelvic girdle Osteichthyans: bony skeleton Gnathostomes: hinged jaws, four sets of Hox genes Vertebrates: Dix genes duplication, backbone of vertebrae Craniates: two sets of Hox genes, neural crest Chordates: notochord; dorsal, hollow nerve cord; pharyngeal slits; post-anal tail Figure 34.UN10 Clade Description Cephalochordata (lancelets) Basal chordates; marine suspension feeders that exhibit four key derived characters of chordates Urochordata (tunicates) Marine suspension feeders; larvae display the derived traits of chordates Myxini (hagfishes and relatives) Jawless marine organisms; have head that includes a skull and brain, eyes, and other sensory organs Petromyzontida (lampreys) Jawless vertebrates; typically feed by attaching to a live fish and ingesting its blood Chondrichthyes (sharks, rays, skates, ratfishes) Actinopterygii (ray-finned fishes) Aquatic gnathostomes; have cartilaginous skeleton, a derived trait formed by the reduction of an ancestral mineralized skeleton Aquatic gnathostomes; have bony skeleton and maneuverable fins supported by rays Actinistia (coelacanths) Dipnoi (lungfishes) Ancient lineage of aquatic lobe-fins still surviving in Indian Ocean Freshwater lobe-fins with both lungs and gills; sister group of tetrapods Amphibia (salamanders, frogs, caecilians) Have four limbs descended from modified fins; most have moist skin that functions in gas exchange; many live both in water (as larvae) and on land (as adults) Reptilia (tuataras, lizards and snakes, turtles, crocodilians, birds) One of two groups of living amniotes; have amniotic eggs and rib cage ventilation, key adaptations for life on land Mammalia (monotremes, marsupials, eutherians) Evolved from synapsid ancestors; include egg-laying monotremes (echidnas, platypus); pouched marsupials (such as kangaroos, opossums); and eutherians (placental mammals, such as rodents, primates)