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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 • Overview: Half a Billion Years of Backbones • By the end of the Cambrian period, some 540 million years ago – An astonishing variety of animals inhabited Earth’s oceans • One of these types of animals – Gave rise to vertebrates, one of the most successful groups of animals 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 Four Characteristics of Chordates • Notochord • Dorsal Hollow Nerve Cord, • Pharyngeal Slits or Clefts, and • Muscular post-anal tail Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 Early Chordate Evolution • The current life history of tunicates – Probably does not reflect that of the ancestral chordate Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Gene expression in lancelets – Holds clues to the evolution of the vertebrate form BF1 Otx Hox3 Nerve cord of lancelet embryo BF1 Hox3 Otx Brain of vertebrate embryo (shown straightened) Midbrain Figure 34.6 Forebrain Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Hindbrain • 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 The Origin of Craniates • Craniates evolved at least 530 million years ago – During the Cambrian explosion Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The most primitive of the fossils – Are those of the 3-cm-long Haikouella (a) Haikouella. Discovered in 1999 in southern China, Haikouella had eyes and a brain but lacked a skull, a derived trait of craniates. Figure 34.8a Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In other Cambrian rocks – Paleontologists have found fossils of even more advanced chordates, such as Haikouichthys 5 mm (b) Haikouichthys. Haikouichthys had a skull and thus is considered a true craniate. Figure 34.8b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Division Myxini Characteristics Jawless marine organisms have head that includes a skull and brain; eyes and sensory organs Petromyzontida Jawless vertebrates --typically feed attaching to a live fish and ingesting its blood Chondrichthyes Aquatic gnathostomes have cartilaginous skeleton secondarily evolved from an ancestral mineralized skelton Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Examples Hagfish lamprey Sharks, rays, skates Actinopterygii Aquatic gnathostomes Yellowfin tuna Bony skeleton Clownfish Maneuverable fins supported by fins Sea Horse Actinista Ancient lineage of aquatic lobe fins still surviving in the Indian Ocean Latimeria Dipnoi Freshwater lobe-fins with both lungs and gills Sister to the tetrapods Have four limbs descended from modified fins Moist skin that function for gas exchange Live both water (as larvae) and on land (as adults) Lungfish Amphibia Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Frogs Salamanders Caecilians Reptilia One of two groups of living amniotes have amniotic eggs and rib cage ventilation – which is the key adaptation to living on land Mammalia Mammary glands Hair Well developed circulatory and respiratory system Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Lizards Snakes Turtles Crocodilians Birds Humans Kangaroos Duckbill Platypus Sponges Zebras 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 Fossils of Early Vertebrates • Conodonts were the first vertebrates – With mineralized skeletal elements in their mouth and pharynx Dorsal view of head Figure 34.11 Dental elements Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Armored, jawless vertebrates called ostracoderms – Had defensive plates of bone on their skin Pteraspis Pharyngolepis Figure 34.12 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Origins of Bone and Teeth • Mineralization – Appears to have originated with vertebrate mouthparts • The vertebrate endoskeleton – Became fully mineralized much later 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 Fossil Gnathostomes • The earliest gnathostomes in the fossil record – Are an extinct lineage of armored vertebrates called placoderms (a) Coccosteus, a placoderm Figure 34.14a Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Another group of jawed vertebrates called acanthodians – Radiated during the Devonian period – Were closely related to the ancestors of osteichthyans (b) Climatius, an acanthodian Figure 34.14b 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 • A second subclass – Is composed of a few dozen species of ratfishes (c) Spotted ratfish (Hydrolagus colliei). Ratfishes, or chimaeras, typically live at depths greater than 80 m and feed on shrimps, molluscs, and sea urchins. Some species have a poisonous spine at the front of their dorsal fin. Figure 34.15c Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Most sharks – Have a streamlined body and are swift swimmers – Have acute senses Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 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 The Origin of Tetrapods • In one lineage of lobe-fins – The fins became progressively more limb-like while the rest of the body retained adaptations for aquatic life Bones supporting gills Figure 34.19 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Tetrapod limb skeleton • Extraordinary fossil discoveries over the past 20 years – Have allowed paleontologists to reconstruct the origin of tetrapods Millions of years ago 420 415 400 385 370 355 340 325 310 295 280 265 Silurian Devonian Carboniferous Paleozoic Permian Ray-finned fishes Coelacanths Lungfishes Eusthenopteron Panderichthys Elginerpeton Metaxygnathus Acanthostega lchthyostega Hynerpeton Greerpeton Amphibians Amniotes Figure 34.20 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings To present 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 • Order Urodela – Includes salamanders, which have tails (a) Order Urodela. Urodeles (salamanders) retain their tail as adults. Figure 34.21a Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Order Anura – Includes frogs and toads, which lack tails (b) Order Anura. Anurans, such as this poison arrow frog, lack a tail as adults. Figure 34.21b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Order Apoda – Includes caecilians, which are legless and resemble worms (c) Order Apoda. Apodans, or caecilians, are legless, mainly burrowing amphibians. Figure 34.21c 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 • A phylogeny of amniotes Saurischians Dinosaurs Lepidosaurs Archosaurs Synapsids Diapsids Reptiles Ancestral amniote Figure 34.23 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 • The extraembryonic membranes – Have various functions Extraembryonic membranes Allantois. The allantois is a disposal sac for certain metabolic wastes produced by the embryo. The membrane of the allantois also functions with the chorion as a respiratory organ. Amnion. The amnion protects the embryo in a fluid-filled cavity that cushions against mechanical shock. Chorion. The chorion and the membrane of the allantois exchange gases between the embryo and the air. Oxygen and carbon dioxide diffuse freely across the shell. Yolk sac. The yolk sac contains the yolk, a stockpile of nutrients. Blood vessels in the yolk sac membrane transport nutrients from the yolk into the embryo. Other nutrients are stored in the albumen (“egg white”). Embryo Amniotic cavity with amniotic fluid Yolk (nutrients) Albumen Shell Figure 34.24 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 Early Amniotes • Early amniotes – Appeared in the Carboniferous period – Included large herbivores and predators 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 Origin and Evolutionary Radiation of Reptiles • The oldest reptilian fossils – Date to about 300 million years ago • The first major group of reptiles to emerge – Were the parareptiles, which were mostly large, stocky herbivores Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • As parareptiles were dwindling – The diapsids were diversifying • The diapsids are composed of two main lineages – The lepidosaurs and the archosaurs 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 • Traditionally, dinosaurs were considered slow, sluggish creatures – But fossil discoveries and research have led to the conclusion that dinosaurs were agile and fast moving • Paleontologists have also discovered signs of parental care among dinosaurs Figure 34.26 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Lepidosaurs • One surviving lineage of lepidosaurs – Is represented by two species of lizard-like reptiles called tuatara Figure 34.27a (a) Tuatara (Sphenodon punctatus) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The other major living lineage of lepidosaurs – Are the squamates, the lizards and snakes • Lizards – Are the most numerous and diverse reptiles, apart from birds Figure 34.27b (b) Australian thorny devil lizard (Moloch horridus) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Snakes are legless lepidosaurs – That evolved from lizards (c) Wagler’s pit viper (Tropidolaemus wagleri), a snake Figure 34.27c 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 • All turtles have a boxlike shell – Made of upper and lower shields that are fused to the vertebrae, clavicles, and ribs Figure 34.27d (d) Eastern box turtle (Terrapene carolina carolina) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Alligators and Crocodiles • Crocodilians – Belong to an archosaur lineage that dates back to the late Triassic Figure 34.27e (e) American alligator (Alligator mississipiensis) 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 The Origin of Birds • Birds probably descended from theropods – A group of small, carnivorous dinosaurs Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 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 Living Birds • The ratites, order Struthioniformes – Are all flightless (a) Emu. This ratite lives in Australia. Figure 34.30a Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 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 Early Evolution of Mammals • Mammals evolved from synapsids – In the late Triassic period 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 • Living lineages of mammals originated in the Jurassic – But did not undergo a significant adaptive radiation until after the Cretaceous 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 some species of marsupials, such as the bandicoot – The marsupium opens to the rear of the mother’s body as opposed to the front, as in other marsupials (b) Long-nosed bandicoot. Most bandicoots are diggers and burrowers that eat mainly insects but also some small vertebrates and plant material. Their rear-opening pouch helps protect the young from dirt as the mother digs. Other marsupials, such as kangaroos, have a pouch that opens to the front. Figure 34.34b 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 • Phylogenetic relationships of mammals This clade of eutherians evolved in Africa when the continent was isolated from other landmasses. It includes Earth’s largest living land animal (the African elephant), as well as species that weigh less than 10 g. All members of this clade, which underwent an adaptive radiation in South America, belong to the order Xenarthra. One species, the nine-banded armadillo, is found in the southern United States. Monotremata Marsupialia Monotremes Marsupials This is the largest eutherian clade. It includes the rodents, which make up the largest mammalian order by far, with about 1,770 species. Humans belong to the order Primates. Proboscidea Sirenia Tubulidentata Hyracoidea Afrosoricida (golden moles and tenrecs) Macroscelidea (elephant shrews) Ancestral mammal Figure 34.36 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Xenarthra Rodentia Lagomorpha Primates Dermoptera (flying lemurs) Scandentia (tree shrews) This diverse clade includes terrestrial and marine mammals as well as bats, the only flying mammals. A growing body of evidence, including Eocene fossils of whales with feet, supports putting whales in the same order (Cetartiodactyla) as pigs, cows, and hippos. Carnivora Cetartiodactyla Perissodactyla Chiroptera Eulipotyphla Pholidota (pangolins) Eutherians Possible phylogenetic tree of mammals. All 20 extant orders of mammals are listed at the top of the tree. Boldfaced orders are explored on the facing page. • The major eutherian orders MAIN CHARACTERISTICS ORDERS AND EXAMPLES Lay eggs; no nipples; young suck milk from fur of mother Monotremata Platypuses, echidnas ORDERS AND EXAMPLES Proboscidea Elephants Koala Long, muscular trunk; thick, loose skin; upper incisors elongated as tusks Tubulidentata Aardvark African elephant Teeth consisting of many thin tubes cemented together; eats ants and termites Aardvark Aquatic; finlike forelimbs and no hind limbs; herbivorous Sirenia Manatees, dugongs Embryo completes development in pouch on mother Marsupialia Kangaroos, opossums, koalas Echidna MAIN CHARACTERISTICS Hyracoidea Hyraxes Rock hyrax Short legs; stumpy tail; herbivorous; complex, multichambered stomach Manatee Xenarthra Sloths, anteaters, armadillos Reduced teeth or no teeth; herbivorous (sloths) or carnivorous (anteaters, armadillos) Rodentia Squirrels, beavers, rats, porcupines, mice Chisel-like, continuously growing incisors worn down by gnawing; herbivorous Red squirrel Tamandua Lagomorpha Rabbits, hares, picas Chisel-like incisors; hind legs longer than forelegs and adapted for running and jumping Primates Lemurs, monkeys, apes, humans Golden lion tamarin Jackrabbit Carnivora Dogs, wolves, bears, cats, weasels, otters, seals, walruses Sharp, pointed canine teeth and molars for shearing; carnivorous Perissodactyla Horses, zebras, tapirs, rhinoceroses Hooves with an even number of toes on each foot; herbivorous Chiroptera Bats Frog-eating bat Bighorn sheep Cetaceans Whales, dolphins, porpoises Figure 34.36 Pacific whitesided porpoise Hooves with an odd number of toes on each foot; herbivorous Indian rhinoceros Coyote Cetartiodactyla Artiodactyls Sheep, pigs cattle, deer, giraffes Opposable thumbs; forward-facing eyes; well-developed cerebral cortex; omnivorous Aquatic; streamlined body; paddle-like forelimbs and no hind limbs; thick layer of insulating blubber; carnivorous Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Eulipotyphla “Core insectivores”: some moles, some shrews Adapted for flight; broad skinfold that extends from elongated fingers to body and legs; carnivorous or herbivorous Diet consists mainly of insects and other small invertebrates Star-nosed mole Primates • The mammalian order Primates include – Lemurs, tarsiers, monkeys, and apes • Humans are members of the ape group Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Derived Characters of Primates • Most primates – Have hands and feet adapted for grasping • Primates also have – A large brain and short jaws – Forward-looking eyes close together on the face, providing depth perception – Well-developed parental care and complex social behavior – A fully opposable thumb Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Living Primates • There are three main groups of living primates – The lemurs of Madagascar and the lorises and pottos of tropical Africa and southern Asia Figure 34.37 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings – The tarsiers of Southeast Asia – The anthropoids, which include monkeys and hominids worldwide Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The oldest known anthropoid fossils, about 45 million years old – Indicate that tarsiers are more closely related to anthropoids Millions of years ago 20 30 40 50 Figure 34.38 60 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Ancestral primate Humans Chimpanzees Gorillas Orangutans Gibbons Old World monkeys New World monkeys 10 Tarsiers 0 Lemurs, lorises, and pottos Anthropoids • The fossil record indicates that monkeys – First appeared in the New World (South America) during the Oligocene • The first monkeys – Evolved in the Old World (Africa and Asia) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • New World and Old World monkeys – Underwent separate adaptive radiations during their many millions of years of separation Figure 34.39a, (a) New World monkeys, such as spider (b) Old World monkeys lack a prehensile tail, and their nostrils monkeys (shown here), squirrel monkeys, and open downward. This group includes macaques (shown here), capuchins, have a prehensile tail and nostrils mandrills, baboons, and rhesus monkeys. that open to the sides. b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The other group of anthropoids, the hominoids – Consists of primates informally called apes (a) Gibbons, such as this Muller's gibbon, are found only in southeastern Asia. Their very long arms and fingers are adaptations for brachiation. (b) Orangutans are shy, solitary apes that live in the rain forests of Sumatra and Borneo. They spend most of their time in trees; note the foot adapted for grasping and the opposable thumb. Figure 34.40a–e (d) Chimpanzees live in tropical Africa. They feed and sleep in trees but also spend a great deal of time on the ground. Chimpanzees are intelligent, communicative, and social. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (e) Bonobos are closely related to chimpanzees but are smaller. They survive today only in the African nation of Congo. (c) Gorillas are the largest apes: some males are almost 2 m tall and weigh about 200 kg. Found only in Africa, these herbivores usually live in groups of up to about 20 individuals. • Hominoids – Diverged from Old World monkeys about 20– 25 million years ago Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 34.8: Humans are bipedal hominoids with a large brain • Homo sapiens is about 160,000 years old – Which is very young considering that life has existed on Earth for at least 3.5 billion years Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Derived Characters of Hominids • A number of characters distinguish humans from other hominoids – Upright posture and bipedal locomotion – Larger brains – Language capabilities – Symbolic thought – The manufacture and use of complex tools – Shortened jaw Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings