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Chapter 34 Vertebrates PowerPoint Lectures for Biology, Seventh Edition Neil Campbell and Jane Reece Lectures by Chris Romero Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The animals called vertebrates – Get their name from vertebrae, the series of bones that make up the backbone Figure 34.1 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • There are approximately 52,000 species of vertebrates – Which include the largest organisms ever to live on the Earth Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 34.1: Chordates have a notochord and a dorsal, hollow nerve cord • Vertebrates are a subphylum of the phylum Chordata • Chordates are bilaterian animals – That belong to the clade of animals known as Deuterostomia • Two groups of invertebrate deuterostomes, the urochordates and cephalochordates – Are more closely related to vertebrates than to invertebrates Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mammalia (mammals) Reptilia (turtles, snakes, crocodiles, birds) Amphibia (frogs, salamanders) Dipnoi (lungfishes) Actinistia (coelacanths) Actinopterygii (ray-finned fishes) Chordates Craniates Vertebrates Gnathostomes Osteichthyans Lobe-fins Tetrapods Amniotes Chondrichthyes (sharks, rays, chimaeras) Cephalaspidomorphi (lampreys) Myxini (hagfishes) Cephalochordata (lancelets) Urochordata (tunicates) Echinodermata (sister group to chordates) • A hypothetical phylogeny of chordates Milk Amniotic egg Legs Lobed fins Lungs or lung derivatives Jaws, mineralized skeleton Vertebral column Head Brain Notochord Figure 34.2 Ancestral deuterostome Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Derived Characters of Chordates • All chordates share a set of derived characters – Although some species possess some of these traits only during embryonic development Dorsal, hollow nerve cord Muscle segments Brain Notochord Mouth Anus Muscular, post-anal tail Figure 34.3 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Pharyngeal slits or clefts Notochord • The notochord – Is a longitudinal, flexible rod located between the digestive tube and the nerve cord – Provides skeletal support throughout most of the length of a chordate • In most vertebrates, a more complex, jointed skeleton develops – And the adult retains only remnants of the embryonic notochord Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Dorsal, Hollow Nerve Cord • The nerve cord of a chordate embryo – Develops from a plate of ectoderm that rolls into a tube dorsal to the notochord – Develops into the central nervous system: the brain and the spinal cord Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Pharyngeal Slits or Clefts • In most chordates, grooves in the pharynx called pharyngeal clefts – Develop into slits that open to the outside of the body • These pharyngeal slits – Function as suspension-feeding structures in many invertebrate chordates – Are modified for gas exchange in aquatic vertebrates – Develop into parts of the ear, head, and neck in terrestrial vertebrates Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Muscular, Post-Anal Tail • Chordates have a tail extending posterior to the anus – Although in many species it is lost during embryonic development • The chordate tail contains skeletal elements and muscles – And it provides much of the propelling force in many aquatic species Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Tunicates • Tunicates, subphylum Urochordata – Belong to the deepest-branching lineage of chordates – Are marine suspension feeders commonly called sea squirts Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Tunicates most resemble chordates during their larval stage – Which may be as brief as a few minutes Notochord Dorsal, hollow nerve cord Tail Excurrent siphon Incurrent siphon Muscle segments Intestine Stomach Atrium Pharynx with slits Figure 34.4c (c) A tunicate larva is a free-swimming but nonfeeding “tadpole” in which all four chief characters of chordates are evident. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • As an adult – A tunicate draws in water through an incurrent siphon, filtering food particles Incurrent siphon to mouth Excurrent siphon Excurrent siphon Atrium Pharynx with numerous slits Tunic Anus Intestine Esophagus Stomach Figure 34.4a, b (a) An adult tunicate, or sea squirt, is a sessile animal (photo is approximately life-sized). Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (b) In the adult, prominent pharyngeal slits function in suspension feeding, but other chordate characters are not obvious. Lancelets • Lancelets, subphylum Cephalochordata – Are named for their bladelike shape Tentacle 2 cm Mouth Pharyngeal slits Atrium Notochord Digestive tract Atriopore Dorsal, hollow nerve cord Segmental muscles Anus Tail Figure 34.5 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Lancelets are marine suspension feeders – That retain the characteristics of the chordate body plan as adults Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 34.2: Craniates are chordates that have a head • The origin of a head – Opened up a completely new way of feeding for chordates: active predation • Craniates share some common characteristics – A skull, brain, eyes, and other sensory organs Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Derived Characters of Craniates • One feature unique to craniates – Is the neural crest, a collection of cells that appears near the dorsal margins of the closing neural tube in an embryo Dorsal edges of neural plate Neural crest Neural tube Ectoderm Ectoderm Notochord (a) The neural crest consists of bilateral bands of cells near the margins of the embryonic Figure 34.7a, b folds that form the neural tube. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Migrating neural crest cells (b) Neural crest cells migrate to distant sites in the embryo. • Neural crest cells – Give rise to a variety of structures, including some of the bones and cartilage of the skull (c) The cells give rise to some of the anatomical structures unique to vertebrates, including some of the bones and cartilage of the skull. Figure 34.7c Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Hagfishes • The least derived craniate lineage that still survives – Is class Myxini, the hagfishes Slime glands Figure 34.9 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Hagfishes are jawless marine craniates – That have a cartilaginous skull and axial rod of cartilage derived from the notochord – That lack vertebrae Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 34.3: Vertebrates are craniates that have a backbone • During the Cambrian period – A lineage of craniates evolved into vertebrates Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Derived Characters of Vertebrates • Vertebrates have – Vertebrae enclosing a spinal cord – An elaborate skull – Fin rays, in aquatic forms Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Lampreys • Lampreys, class Cephalaspidomorphi – Represent the oldest living lineage of vertebrates – Have cartilaginous segments surrounding the notochord and arching partly over the nerve cord Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Lampreys are jawless vertebrates – Inhabiting various marine and freshwater habitats Figure 34.10 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 34.4: Gnathostomes are vertebrates that have jaws • Today, jawless vertebrates – Are far outnumbered by those with jaws Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Derived Characters of Gnathostomes • Gnathostomes have jaws – That evolved from skeletal supports of the Gill slits pharyngeal slits Cranium Mouth Skeletal rods Figure 34.13 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Other characters common to gnathostomes include – Enhanced sensory systems, including the lateral line system – An extensively mineralized endoskeleton – Paired appendages Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Chondrichthyans (Sharks, Rays, and Their Relatives) • Members of class Chondrichthyes – Have a skeleton that is composed primarily of cartilage • The cartilaginous skeleton – Evolved secondarily from an ancestral mineralized skeleton Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The largest and most diverse subclass of Chondrichthyes – Includes the sharks and rays (a) Blacktip reef shark (Carcharhinus melanopterus). Fast swimmers with acute senses, sharks have paired pectoral and pelvic fins. (b) Southern stingray (Dasyatis americana). Most rays are flattened bottom-dwellers that crush molluscs and crustaceans for food. Some rays cruise in open water and scoop food into Figure 34.15a, b their gaping mouth. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Pectoral fins Pelvic fins Ray-Finned Fishes and Lobe-Fins • The vast majority of vertebrates – Belong to a clade of gnathostomes called Osteichthyes Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Nearly all living osteichthyans – Have a bony endoskeleton • Aquatic osteichthyans – Are the vertebrates we informally call fishes – Control their buoyancy with an air sac known as a swim bladder Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Fishes breathe by drawing water over four or five pairs of gills – Located in chambers covered by a protective bony flap called the operculum Adipose fin Dorsal fin (characteristic of trout) Swim bladder Nostril Spinal cord Brain Cut edge of operculum Gills Heart Figure 34.16 Gonad Urinary Anus bladder Liver Kidney Stomach Pelvic fin Intestine Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Caudal fin Anal fin Lateral line Ray-Finned Fishes • Class Actinopterygii, the ray-finned fishes – Includes nearly all the familiar aquatic osteichthyans (a) Yellowfin tuna (Thunnus albacares), a fast-swimming, schooling fish that is an important commercial fish worldwide (b) Clownfish (Amphiprion ocellaris), a mutualistic symbiont of sea anemones Figure 34.17a–d (c) Sea horse (Hippocampus ramulosus), unusual in the animal kingdom in that the male carries the young during their embryonic development Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (d) Fine-spotted moray eel (Gymnothorax dovii), a predator that ambushes prey from crevices in its coral reef habitat • The fins, supported mainly by long, flexible rays – Are modified for maneuvering, defense, and other functions Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Lobe-Fins • The lobe-fins, class Sarcopterygii – Have muscular and pectoral fins – Include coelacanths, lungfishes, and tetrapods Figure 34.18 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 34.5: Tetrapods are gnathostomes that have limbs and feet • 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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Derived Characters of Tetrapods • Tetrapods have some specific adaptations – Four limbs and feet with digits – Ears for detecting airborne sounds Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Amphibians • Class Amphibia – Is represented by about 4,800 species of organisms • Most amphibians – Have moist skin that complements the lungs in gas exchange Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Amphibian means “two lives” – A reference to the metamorphosis of an aquatic larva into a terrestrial adult (b) The tadpole is an aquatic herbivore with a fishlike tail and internal gills. (a) The male grasps the female, stimulating her to release eggs. The eggs are laid and fertilized in water. They have a jelly coat but lack a shell and Figure 34.22a–c would desiccate in air. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (c) During metamorphosis, the gills and tail are resorbed, and walking legs develop. • 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 the mammals Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Derived Characters of Amniotes • Amniotes are named for the major derived character of the clade, the amniotic egg – Which contains specialized membranes that protect the embryo Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Amniotes also have other terrestrial adaptations – Such as relatively impermeable skin and the ability to use the rib cage to ventilate the lungs Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Reptiles • The reptile clade includes – The tuatara, lizards, snakes, turtles, crocodilians, birds, and the extinct dinosaurs Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Reptiles – Have scales that create a waterproof barrier – Lay shelled eggs on land Figure 34.25 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The dinosaurs – Diversified into a vast range of shapes and sizes – Included the long-necked giants called the theropods Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Turtles • Turtles – Are the most distinctive group of reptiles alive today • Some turtles have adapted to deserts – And others live entirely in ponds and rivers Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Birds • Birds are archosaurs – But almost every feature of their reptilian anatomy has undergone modification in their adaptation to flight Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Derived Characters of Birds • Many of the characters of birds – Are adaptations that facilitate flight Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • A bird’s most obvious adaptations for flight – Are its wings and feathers Finger 1 (b) Bone structure Palm (a) wing Finger 2 Forearm Wrist Finger 3 Shaft Vane Shaft Figure 34.28a–c Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Barb Barbule Hook (c) Feather structure • By 150 million years ago – Feathered theropods had evolved into birds • Archaeopteryx – Remains the oldest bird known Wing claw Toothed beak Airfoil wing with contour feathers Figure 34.29 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Long tail with many vertebrae • The demands of flight – Have rendered the general body form of many flying birds similar to one another (b) Mallards. Like many bird species, the mallard exhibits pronounced color differences between the sexes. (c) Laysan albatrosses. Like most birds, Laysan albatrosses have specific mating behaviors, such as this courtship ritual. (d) Barn swallows. The barn swallow is a member of the order Passeriformes. Species in this order are called perching birds because the toes of their feet can lock around a branch or wire, enabling the bird Figure 34.30b–d to rest in place for long periods. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Foot structure in bird feet – Shows considerable variation Perching bird (such as a cardinal) Grasping bird (such as a woodpecker) Figure 34.31 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Raptor (such as a bald eagle) Swimming bird (such as a duck) • Concept 34.7: Mammals are amniotes that have hair and produce milk • Mammals, class Mammalia – Are represented by more than 5,000 species Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Derived Characters of Mammals • Mammary glands, which produce milk – Are a distinctively mammalian character • Hair is another mammalian characteristic • Mammals generally have a larger brain – Than other vertebrates of equivalent size Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The jaw was remodeled during the evolution of mammals from nonmammalian synapsids – And two of the bones that formerly made of the jaw joint were incorporated into the mammalian middle ear Jaw joint Key Jaw joint Dentary Angular Squamosal Articular Quadrate Dimetrodon Morganucodon (a) The lower jaw of Dimetrodon is composed of several fused bones; two small bones, the quadrate and articular, form part of the jaw joint. In Morganucodon, the lower jaw is reduced to a single bone, the dentary, and the location of the jaw joint has shifted. Middle ear Stapes Eardrum Inner ear Middle ear Inner ear Eardrum Stapes Sound Sound Incus (evolved from quadrate) Malleus (evolved from articular) Figure 34.32a, b Morganucodon Dimetrodon (b) During the evolutionary remodeling of the mammalian skull, the quadrate and articular bones became incorporated into the middle ear as two of the three bones that transmit sound from the eardrum to the inner ear. The steps in this evolutionary remodeling are evident in a succession of fossils. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Monotremes • Monotremes – Are a small group of egg-laying mammals consisting of echidnas and the platypus Figure 34.33 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Marsupials • Marsupials – Include opossums, kangaroos, and koalas Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • A marsupial is born very early in its development – And completes its embryonic development while nursing within a maternal pouch called a marsupium (a) A young brushtail possum. The young of marsupials are born very early in their development. They finish their growth while nursing from a nipple (in their mother’s pouch in most species). Figure 34.34a Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In Australia, convergent evolution – Has resulted in a diversity of marsupials that resemble eutherians in other parts of the world Marsupial mammals Plantigale Marsupial mole Eutherian mammals Deer mouse Mole Sugar glider Flying squirrel Wombat Woodchuck Tasmanian devil Kangaroo Figure 34.35 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Wolverine Patagonian cavy Eutherians (Placental Mammals) • Compared to marsupials – Eutherians have a longer period of pregnancy • Young eutherians – Complete their embryonic development within a uterus, joined to the mother by the placenta Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings