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Chapter 34 Vertebrates
Overview: Half a Billion Years of Backbones <ul><li>Early in the Cambrian period, about 530 million years ago, an astonish...
Fig. 34-1
<ul><li>The animals called  vertebrates  get their name from vertebrae, the series of bones that make up the backbone </li...
Concept 34.1: Chordates have a notochord and a dorsal, hollow nerve cord <ul><li>Vertebrates are a subphylum within the ph...
Fig. 34-2 Lobed fins Legs Amniotic egg Milk Jaws, mineralized skeleton Lungs or lung derivatives Vertebral column Head Not...
Fig. 34-2a Lobed fins Legs Amniotic egg Milk Jaws, mineralized skeleton Lungs or lung derivatives Vertebral column Head No...
Fig. 34-2b Lobed fins Legs Amniotic egg Milk Jaws, mineralized skeleton Lungs or lung derivatives Vertebral column Head Ch...
Fig. 34-2c Lobed fins Legs Amniotic egg Milk Jaws, mineralized skeleton Lungs or lung derivatives Vertebral column Chondri...
Fig. 34-2d Lobed fins Legs Amniotic egg Milk Jaws, mineralized skeleton Lungs or lung derivatives Chondrichthyes (sharks, ...
Fig. 34-2e Lobed fins Legs Amniotic egg Milk Lungs or lung derivatives Mammalia (mammals) Actinopterygii (ray-finned fishe...
Fig. 34-2f Lobed fins Legs Amniotic egg Milk Mammalia (mammals) Actinistia (coelacanths) Amphibia (frogs, salamanders) Dip...
Fig. 34-2g Legs Amniotic egg Milk Mammalia (mammals) Amphibia (frogs, salamanders) Reptilia (turtles, snakes, crocodiles, ...
Fig. 34-2h Amniotic egg Milk Mammalia (mammals) Reptilia (turtles, snakes, crocodiles, birds) Amniotes
Derived Characters of Chordates <ul><li>All chordates share a set of derived characters </li></ul><ul><li>Some species hav...
Fig. 34-3 Dorsal, hollow nerve cord Anus Muscular, post-anal tail Pharyngeal slits or clefts Notochord Mouth Muscle segments
Notochord <ul><li>The  notochord  is a longitudinal, flexible rod between the digestive tube and nerve cord </li></ul><ul>...
Dorsal, Hollow Nerve Cord <ul><li>The nerve cord of a chordate embryo develops from a plate of ectoderm that rolls into a ...
Pharyngeal Slits or Clefts <ul><li>In most chordates, grooves in the pharynx called  pharyngeal clefts  develop into slits...
Muscular, Post-Anal Tail <ul><li>Chordates have a tail posterior to the anus </li></ul><ul><li>In many species, the tail i...
Lancelets <ul><li>Lancelets  (Cephalochordata) are named for their bladelike shape </li></ul><ul><li>They are marine suspe...
Fig. 34-UN1 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amp...
Fig. 34-4 Dorsal, hollow nerve cord Notochord Tail Cirri Mouth Pharyngeal slits Digestive tract Atrium Atriopore Segmental...
Tunicates <ul><li>Tunicates  (Urochordata) are more closely related to other chordates than are lancelets </li></ul><ul><l...
Fig. 34-UN2 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amp...
Fig. 34-5 Tunic Water flow Excurrent siphon Atrium An adult tunicate Pharynx with slits Anus Atrium Excurrent siphon Incur...
<ul><li>Tunicates most resemble chordates during their larval stage, which may last only a few minutes </li></ul>
Early Chordate Evolution <ul><li>Ancestral chordates may have resembled lancelets </li></ul><ul><li>Genome sequencing of t...
Fig. 34-6 BF1 Brain of vertebrate embryo (shown straightened) Hindbrain Forebrain Midbrain Nerve cord of lancelet embryo B...
Concept 34.2: Craniates are chordates that have a head <ul><li>The origin of a head opened up a completely new way of feed...
Derived Characters of Craniates <ul><li>Craniates have two clusters of  Hox  genes; lancelets and tunicates have only one ...
Fig. 34-7 Migrating neural crest cells Notochord Dorsal edges of neural plate Neural crest Neural tube
<ul><li>In aquatic craniates the pharyngeal clefts evolved into gill slits </li></ul><ul><li>Craniates have a higher metab...
The Origin of Craniates <ul><li>Fossils from the Cambrian explosion 530 million years ago document the transition to crani...
Fig. 34-8 Segmented muscles Pharyngeal slits 5 mm
Fig. 34-8a 5 mm
Fig. 34-8b Segmented muscles Pharyngeal slits
<ul><li>In other Cambrian rocks, paleontologists have found fossils of even more advanced chordates, such as  Myllokunming...
Hagfishes <ul><li>The least derived surviving craniate lineage is Myxini, the hagfishes </li></ul><ul><li>Hagfishes have a...
Fig. 34-UN3 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amp...
Fig. 34-9 Slime glands
Concept 34.3: Vertebrates are craniates that have a backbone <ul><li>During the Cambrian period, a lineage of craniates ev...
Derived Characters of Vertebrates <ul><li>Vertebrates underwent a second gene duplication involving the  Dlx  family of tr...
Lampreys <ul><li>Lampreys (Petromyzontida) represent the oldest living lineage of vertebrates </li></ul><ul><li>They are j...
Fig. 34-UN4 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amp...
Fig. 34-10
Fig. 34-10a
Fig. 34-10b
Fossils of Early Vertebrates <ul><li>Conodonts  were the first vertebrates with mineralized skeletal elements in their mou...
Fig. 34-11 Dental elements
<ul><li>Other armored, jawless vertebrates had defensive plates of bone on their skin </li></ul>
Fig. 34-12 Pteraspis Pharyngolepis
Origins of Bone and Teeth <ul><li>Mineralization appears to have originated with vertebrate mouthparts </li></ul><ul><li>T...
Concept 34.4: Gnathostomes are vertebrates that have jaws <ul><li>Today, jawed vertebrates, or  gnathostomes , outnumber j...
Derived Characters of Gnathostomes <ul><li>Gnathostomes have jaws that might have evolved from skeletal supports of the ph...
Fig. 34-13-1 Skeletal rods Cranium Gill slits Mouth
Fig. 34-13-2 Skeletal rods Cranium Gill slits Mouth
Fig. 34-13-3 Skeletal rods Cranium Gill slits Mouth
<ul><li>Other characters common to gnathostomes: </li></ul><ul><ul><li>An additional duplication of  Hox  genes </li></ul>...
Fossil Gnathostomes <ul><li>The earliest gnathostomes in the fossil record are an extinct lineage of armored vertebrates c...
Fig. 34-14 0.5 m
<ul><li>Another group of jawed vertebrates called acanthodians radiated during the Devonian period </li></ul>
Chondrichthyans (Sharks, Rays, and Their Relatives) <ul><li>Chondrichthyans  (Chondrichthyes) have a skeleton composed pri...
Fig. 34-UN5 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amp...
Fig. 34-15 Pelvic fins Pectoral fins (c) Spotted ratfish ( Hydrolagus colliei ) (a) Blacktip reef shark ( Carcharhinus    ...
Fig. 34-15a Pelvic fins Pectoral fins (a) Blacktip reef shark ( Carcharhinus melanopterus )
Fig. 34-15b (b) Southern stingray ( Dasyatis americana )
<ul><li>A second subclass is composed of a few dozen species of ratfishes </li></ul>
Fig. 34-15c (c) Spotted ratfish ( Hydrolagus colliei )
<ul><li>Most sharks   </li></ul><ul><ul><li>Have a streamlined body and are swift swimmers </li></ul></ul><ul><ul><li>Are ...
<ul><li>Shark eggs are fertilized internally but embryos can develop in different ways: </li></ul><ul><ul><li>Oviparous : ...
<ul><li>The reproductive tract, excretory system, and digestive tract empty into a common  cloaca </li></ul>
Ray-Finned Fishes and Lobe-Fins <ul><li>The vast majority of vertebrates belong to a clade of gnathostomes called Osteicht...
<ul><li>Nearly all living  osteichthyans  have a bony endoskeleton </li></ul><ul><li>Aquatic osteichthyans are the vertebr...
Fig. 34-UN6 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amp...
Fig. 34-16 Intestine Adipose fin (characteristic of trout) Cut edge of operculum Swim bladder Caudal fin Lateral line Urin...
Ray-Finned Fishes <ul><li>Class Actinopterygii, the  ray-finned fishes , includes nearly all the familiar aquatic osteicht...
Fig. 34-17 (a) Yellowfin tuna ( Thunnus albacares ) (b) Clownfish ( Amphiprion ocellaris ) (c) Sea horse ( Hippocampus    ...
Fig. 34-17a (a) Yellowfin tuna ( Thunnus albacares )
Fig. 34-17b (b) Clownfish ( Amphiprion ocellaris )
Fig. 34-17c (c) Sea horse ( Hippocampus ramulosus )
Fig. 34-17d (d) Fine-spotted moray eel ( Gymnothorax dovii )
Lobe-Fins <ul><li>The  lobe-fins  (Sarcopterygii) have muscular pelvic and pectoral fins </li></ul><ul><li>Three lineages ...
Fig. 34-18
Concept 34.5: Tetrapods are gnathostomes that have limbs <ul><li>One of the most significant events in vertebrate history ...
Derived Characters of Tetrapods <ul><li>Tetrapods  have some specific adaptations: </li></ul><ul><ul><li>Four limbs, and f...
The Origin of Tetrapods <ul><li>In one lineage of lobe-fins, the fins became progressively more limb-like while the rest o...
Fig. 34-19 Tetrapod limb skeleton Bones supporting gills
<ul><li>Extraordinary fossil discoveries over the past 20 years have allowed paleontologists to reconstruct the origin of ...
Fig. 34-20 Ray-finned fishes Coelacanths Lungfishes Eusthenopteron Panderichthys Tiktaalik Elginerpeton Metaxygnathus Acan...
Amphibians <ul><li>Amphibians  (class Amphibia) are represented by about 6,150 species of organisms in three orders </li><...
Fig. 34-UN7 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amp...
Fig. 34-21 (a) Order Urodela (b) Order Anura (c) Order Apoda
Fig. 34-21a (a) Order Urodela
<ul><li>Order Anura includes frogs and toads, which lack tails </li></ul>
Fig. 34-21b (b) Order Anura
<ul><li>Order Apoda includes caecilians, which are legless and resemble worms </li></ul>
Fig. 34-21c (c) Order Apoda
<ul><li>Amphibian  means “both ways of life,” referring to the metamorphosis of an aquatic larva into a terrestrial adult ...
Fig. 34-22 (c) Mating adults (a) Tadpole (b) During metamorphosis
Fig. 34-22a (a) Tadpole
Fig. 34-22b (b) During metamorphosis
Fig. 34-22c (c) Mating adults
Fig. 34-23
Concept 34.6: Amniotes are tetrapods that have a terrestrially adapted egg <ul><li>Amniotes  are a group of tetrapods whos...
Fig. 34-24 Reptiles ANCESTRAL AMNIOTE Diapsids Synapsids Lepidosaurs Archosaurs Dinosaurs Saurischians Mammals Squamates T...
Derived Characters of Amniotes <ul><li>Amniotes are named for the major derived character of the clade, the  amniotic egg ...
Fig. 34-25 Yolk sac Amniotic cavity with amniotic fluid Chorion Amnion Albumen Yolk (nutrients) Allantois Embryo Shell
<ul><li>Amniotes have other terrestrial adaptations, such as relatively impermeable skin and the ability to use the rib ca...
Early Amniotes <ul><li>Living amphibians and amniotes split from a common ancestor about 370 million years ago </li></ul><...
Reptiles <ul><li>The  reptile  clade includes the tuataras, lizards, snakes, turtles, crocodilians, birds, and the extinct...
Fig. 34-UN8 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amp...
Fig. 34-26
<ul><li>Most reptiles are  ectothermic , absorbing external heat as the main source of body heat </li></ul><ul><li>Birds a...
The Origin and Evolutionary Radiation of Reptiles <ul><li>The oldest reptilian fossils date to about 310 million years ago...
<ul><li>As parareptiles were dwindling, the  diapsids  were diversifying </li></ul><ul><li>The diapsids consisted of two m...
<ul><li>The dinosaurs diversified into a vast range of shapes and sizes </li></ul><ul><li>They included bipedal carnivores...
<ul><li>Dinosaurs, with the exception of birds, became extinct by the end of the Cretaceous </li></ul><ul><li>Their extinc...
Lepidosaurs <ul><li>One surviving lineage of lepidosaurs is represented by two species of lizard-like reptiles called tuat...
Fig. 34-27 (a) Tuatara ( Sphenodon punctatus ) (c) Wagler’s pit viper ( Tropidolaemus wagleri ) (b) Australian thorny devi...
Fig. 34-27a (a) Tuatara ( Sphenodon punctatus )
<ul><li>The other major living lineage of lepidosaurs consists of the squamates, the lizards and snakes </li></ul><ul><li>...
Fig. 34-27b (b) Australian thorny devil lizard ( Moloch horridus )
<ul><li>Snakes are legless lepidosaurs that evolved from lizards </li></ul>Video: Snake Ritual Wrestling
Fig. 34-27c (c) Wagler’s pit viper ( Tropidolaemus wagleri )
Turtles <ul><li>Turtles are the most distinctive group of reptiles alive today </li></ul><ul><li>All turtles have a boxlik...
Fig. 34-27d (d) Eastern box turtle ( Terrapene carolina carolina )
Alligators and Crocodiles <ul><li>Crocodilians (alligators and crocodiles) belong to an archosaur lineage that dates back ...
Fig. 34-27e (e) American alligator ( Alligator mississippiensis )
Birds <ul><li>Birds are archosaurs, but almost every feature of their reptilian anatomy has undergone modification in thei...
<ul><li>Many characters of birds are adaptations that facilitate flight </li></ul><ul><li>The major adaptation is wings wi...
Fig. 34-28 (a) Wing (b) Bone structure (c) Feather structure Finger 1 Finger 2 Finger 3 Palm Hook Vane Barbule Barb Shaft ...
<ul><li>Flight enhances hunting and scavenging, escape from terrestrial predators, and migration </li></ul><ul><li>Flight ...
<ul><li>Birds probably descended from small theropods, a group of carnivorous dinosaurs </li></ul><ul><li>By 150 million y...
Fig. 34-29 Airfoil wing with contour feathers Toothed beak Wing claw Long tail with many vertebrae
<ul><li>Living birds belong to the clade Neornithes </li></ul><ul><li>Several groups of birds are flightless </li></ul><ul...
<ul><li>The demands of flight have rendered the general body form of many flying birds similar to one another </li></ul><u...
Fig. 34-30 (a) Emu (b) Mallards (c) Laysan albatrosses (d) Barn swallows
Fig. 34-30a (a) Emu
Fig. 34-30b (b) Mallards
Fig. 34-30c (c) Laysan albatrosses
Fig. 34-30d (d) Barn swallows
Concept 34.7: Mammals are amniotes that have hair and produce milk <ul><li>Mammals , class Mammalia, are represented by mo...
Fig. 34-UN9 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amp...
Derived Characters of Mammals <ul><li>Mammals have </li></ul><ul><ul><li>Mammary glands, which produce milk </li></ul></ul...
Early Evolution of Mammals <ul><li>Mammals evolved from  synapsids  in the late Triassic period </li></ul><ul><li>Two bone...
Fig. 34-31 (b) In mammals, the articular and quadrate bones are incorporated into the middle ear. (a) In  Biarmosuchus,  a...
<ul><li>By the early Cretaceous, the three living lineages of mammals emerged: monotremes, marsupials, and eutherians </li...
Monotremes <ul><li>Monotremes  are a small group of egg-laying mammals consisting of echidnas and the platypus </li></ul>
Fig. 34-32
Marsupials <ul><li>Marsupials  include opossums, kangaroos, and koalas </li></ul><ul><li>The embryo develops within a  pla...
Fig. 34-33 (a) A young brushtail possum (b) Long-nosed bandicoot
Fig. 34-33a (a) A young brushtail possum
<ul><li>In some species, such as the bandicoot, the marsupium opens to the rear of the mother’s body </li></ul>
Fig. 34-33b (b) Long-nosed bandicoot
<ul><li>In Australia, convergent evolution has resulted in a diversity of marsupials that resemble the eutherians in other...
Fig. 34-34 Plantigale Marsupial mammals Eutherian mammals Marsupial mammals Eutherian mammals Marsupial mole Flying squirr...
Eutherians (Placental Mammals) <ul><li>Compared with marsupials,  eutherians  have a longer period of pregnancy </li></ul>...
Fig. 34-35a ANCESTRAL MAMMAL Monotremata Marsupialia Monotremes (5 species) Marsupials (324 species) Eutherians (5,010 spe...
Fig. 34-35b
Fig. 34-35c
Fig. 34-35d
Fig. 34-35e
Fig. 34-35f
Fig. 34-35g
Fig. 34-35h
Video: Bat Licking Nectar Video: Bat Pollinating Agave Plant Video: Galápagos Sea Lion Video: Wolf Agonistic Behavior
Primates <ul><li>The mammalian order Primates includes lemurs, tarsiers, monkeys, and apes </li></ul><ul><li>Humans are me...
<ul><li>Most primates have hands and feet adapted for grasping </li></ul>Derived Characters of Primates
<ul><li>Other derived characters of primates:   </li></ul><ul><ul><li>A large brain and short jaws </li></ul></ul><ul><ul>...
<ul><li>There are three main groups of living primates: </li></ul><ul><ul><li>Lemurs, lorises, and pottos  </li></ul></ul>...
Fig. 34-36
<ul><li>The oldest known anthropoid fossils, about 45 million years old, indicate that tarsiers are more closely related t...
Fig. 34-37 Lemurs, lorises, and pottos Tarsiers New World monkeys Old World monkeys Gibbons Orangutans Gorillas Chimpanzee...
<ul><li>The first monkeys evolved in the Old World (Africa and Asia) </li></ul><ul><li>In the New World (South America), m...
Fig. 34-38 (a) New World monkey (b) Old World monkey
Fig. 34-38a (a) New World monkey
Fig. 34-38b (b) Old World monkey
<ul><li>The other group of anthropoids consists of primates informally called apes </li></ul><ul><li>This group includes g...
Fig. 34-39 (e) Bonobos (a) Gibbon (d) Chimpanzees (b) Orangutan (c) Gorilla
Fig. 34-39a (a) Gibbon
Fig. 34-39b (b) Orangutan
Fig. 34-39c (c) Gorilla
Fig. 34-39d (d) Chimpanzees
Fig. 34-39e (e) Bonobos
Concept 34.8: Humans are mammals that have a large brain and bipedal locomotion <ul><li>The species  Homo sapiens  is abou...
Derived Characters of Humans <ul><li>A number of characters distinguish humans from other apes: </li></ul><ul><ul><li>Upri...
The Earliest Hominins <ul><li>The study of human origins is known as  paleoanthropology </li></ul><ul><li>Hominins  (forme...
Fig. 34-40 Homo erectus Homo habilis Homo sapiens Homo neanderthalensis ? Homo ergaster Paranthropus robustus Paranthropus...
<ul><li>Hominins originated in Africa about 6–7 million years ago </li></ul><ul><li>Early hominins had a small brain but p...
<ul><li>Two common misconceptions about early hominins: </li></ul><ul><ul><li>Thinking of them as chimpanzees </li></ul></...
Australopiths <ul><li>Australopiths are a paraphyletic assemblage of hominins living between 4 and 2 million years ago </l...
Fig. 34-41 (c) An artist’s reconstruction of what  A. afarensis  may have looked like (a)  Australopithecus afarensis  ske...
Fig. 34-41a (a)  Australopithecus afarensis  skeleton
Fig. 34-41b (b)   The Laetoli   footprints
Fig. 34-41c (c) An artist’s reconstruction of what  A. afarensis may have looked like
Bipedalism <ul><li>Hominins began to walk long distances on two legs about 1.9 million years ago </li></ul>
Tool Use <ul><li>The oldest evidence of tool use, cut marks on animal bones, is 2.5 million years old </li></ul>
Early  Homo <ul><li>The earliest fossils placed in our genus  Homo  are those of  Homo habilis , ranging in age from about...
<ul><li>Homo ergaster  was the first fully bipedal, large-brained hominid </li></ul><ul><li>The species existed between 1....
<ul><li>Homo ergaster  fossils were previously assigned to  Homo erectus ; most paleoanthropologists now recognize these a...
Fig. 34-42
<ul><li>Homo erectus  originated in Africa by 1.8 million years ago </li></ul><ul><li>It was the first hominin to leave Af...
Neanderthals <ul><li>Neanderthals,  Homo neanderthalensis,  lived in Europe and the Near East from 200,000 to 28,000 years...
Fig. 34-43 Chimpanzees Chimpanzees European and other living humans Neanderthals Living Europeans Other living humans Nean...
Fig. 34-43a Chimpanzees Neanderthals Living Europeans Other living humans Hypothesis: Neanderthals gave rise to European h...
Fig. 34-43b Chimpanzees European and other living humans Neanderthal 1 Neanderthal 2 RESULTS
Homo Sapiens <ul><li>Homo sapiens  appeared in Africa by 195,000 years ago </li></ul><ul><li>All living humans are descend...
Fig. 34-44
<ul><li>The oldest fossils of  Homo sapiens  outside Africa date back about 115,000 years and are from the Middle East </l...
<ul><li>Rapid expansion of our species may have been preceded by changes to the brain that made cognitive innovations poss...
Fig. 34-45
Fig. 34-UN10
Fig. 34-UN10a
Fig. 34-UN10b
Fig. 34-UN10c
Fig. 34-UN10d
Fig. 34-UN10e
Fig. 34-UN10f
Fig. 34-UN10g
Fig. 34-T1
Fig. 34-UN11
Fig. 34-UN12
You should now be able to: <ul><li>List the derived traits for: chordates, craniates, vertebrates, gnathostomes, tetrapods...
<ul><li>Define and distinguish among gnathostomes, tetrapods, and amniotes </li></ul><ul><li>Describe an amniotic egg and ...
<ul><li>Define the term hominin </li></ul><ul><li>Describe the evolution of Homo sapiens from australopith ancestors, and ...
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  • Figure 34.1 Are humans among the descendants of this ancient organism?
  • Figure 34.2 Phylogeny of living chordates
  • Figure 34.2 Phylogeny of living chordates
  • Figure 34.2 Phylogeny of living chordates
  • Figure 34.2 Phylogeny of living chordates
  • Figure 34.2 Phylogeny of living chordates
  • Figure 34.2 Phylogeny of living chordates
  • Figure 34.2 Phylogeny of living chordates
  • Figure 34.2 Phylogeny of living chordates
  • Figure 34.2 Phylogeny of living chordates
  • Figure 34.3 Chordate characteristics
  • Figure 34.4 The lancelet Branchiostoma , a cephalochordate
  • Figure 34.5 A tunicate, a urochordate
  • Figure 34.6 Expression of developmental genes in lancelets and vertebrates
  • Figure 34.7 The neural crest, embryonic source of many unique craniate characters
  • Figure 34.8 Fossil of an early chordate
  • Figure 34.8 Fossil of an early chordate
  • Figure 34.8 Fossil of an early chordate
  • Figure 34.9 A hagfish
  • Figure 34.10 A sea lamprey
  • Figure 34.10 A sea lamprey
  • Figure 34.10 A sea lamprey
  • Figure 34.11 A conodont
  • Figure 34.12 Jawless armored vertebrates
  • Figure 34.13 Hypothesis for the evolution of vertebrate jaws
  • Figure 34.13 Hypothesis for the evolution of vertebrate jaws
  • Figure 34.13 Hypothesis for the evolution of vertebrate jaws
  • Figure 34.14 Fossil of an early gnathostome
  • Figure 34.15 Chondrichthyans
  • Figure 34.15 Chondrichthyans
  • Figure 34.15 Chondrichthyans
  • Figure 34.15 Chondrichthyans
  • Figure 34.16 Anatomy of a trout, a ray-finned fish
  • Figure 34.17 Ray-finned fishes (class Actinopterygii)
  • Figure 34.17 Ray-finned fishes (class Actinopterygii)
  • Figure 34.17 Ray-finned fishes (class Actinopterygii)
  • Figure 34.17 Ray-finned fishes (class Actinopterygii)
  • Figure 34.17 Ray-finned fishes (class Actinopterygii)
  • Figure 34.18 A coelacanth ( Latimeria )
  • Figure 34.19 Acanthostega , a Devonian relative of tetrapods
  • Figure 34.20 The origin of tetrapods
  • Figure 34.21 Amphibians
  • Figure 34.21 Amphibians
  • Figure 34.21 Amphibians
  • Figure 34.21 Amphibians
  • Figure 34.22 The “dual life” of a frog ( Rana temporaria )
  • Figure 34.22 The “dual life” of a frog ( Rana temporaria )
  • Figure 34.22 The “dual life” of a frog ( Rana temporaria )
  • Figure 34.22 The “dual life” of a frog ( Rana temporaria )
  • Figure 34.23 A mobile nursery
  • Figure 34.24 A phylogeny of amniotes
  • Figure 34.25 The amniotic egg
  • Figure 34.26 Hatching reptiles
  • Figure 34.27 Extant reptiles (other than birds)
  • Figure 34.27 Extant reptiles (other than birds)
  • Figure 34.27 Extant reptiles (other than birds)
  • Figure 34.27 Extant reptiles (other than birds)
  • Figure 34.27 Extant reptiles (other than birds)
  • Figure 34.27 Extant reptiles (other than birds)
  • Figure 34.28 Form fits function: the avian wing and feather
  • Figure 34.29 Artist’s reconstruction of Archaeopteryx , the earliest known bird
  • Figure 34.30 A small sample of living birds
  • Figure 34.30 A small sample of living birds
  • Figure 34.30 A small sample of living birds
  • Figure 34.30 A small sample of living birds
  • Figure 34.30 A small sample of living birds
  • Figure 34.31 The evolution of the mammalian ear bones
  • Figure 34.32 Short-beaked echidna ( Tachyglossus aculeatus ), an Australian monotreme
  • Figure 34.33 Australian marsupials
  • Figure 34.33 Australian marsupials
  • Figure 34.33 Australian marsupials
  • Figure 34.34 Evolutionary convergence of marsupials and eutherians (placental mammals)
  • Figure 34.35 Mammalian diversity
  • Figure 34.35 Mammalian diversity
  • Figure 34.35 Mammalian diversity
  • Figure 34.35 Mammalian diversity
  • Figure 34.35 Mammalian diversity
  • Figure 34.35 Mammalian diversity
  • Figure 34.35 Mammalian diversity
  • Figure 34.35 Mammalian diversity
  • Figure 34.36 Coquerel’s sifakas ( Propithecus verreauxi coquereli ), a type of lemur
  • Figure 34.37 A phylogenetic tree of primates
  • Figure 34.38 New World monkeys and Old World monkeys
  • Figure 34.38 New World monkeys and Old World monkeys
  • Figure 34.38 New World monkeys and Old World monkeys
  • Figure 34.39 Nonhuman apes
  • Figure 34.39 Nonhuman apes
  • Figure 34.39 Nonhuman apes
  • Figure 34.39 Nonhuman apes
  • Figure 34.39 Nonhuman apes
  • Figure 34.39 Nonhuman apes
  • Figure 34.40 A timeline for some selected hominin species
  • Figure 34.41 Upright posture predates an enlarged brain in human evolution
  • Figure 34.41 Upright posture predates an enlarged brain in human evolution
  • Figure 34.41 Upright posture predates an enlarged brain in human evolution
  • Figure 34.41 Upright posture predates an enlarged brain in human evolution
  • Figure 34.42 Fossil and artist’s reconstruction of Homo ergaster
  • Figure 34.43 Did Neanderthals give rise to European humans?
  • Figure 34.43 Did Neanderthals give rise to European humans?
  • Figure 34.43 Did Neanderthals give rise to European humans?
  • Figure 34.44 A 160,000-year-old fossil of Homo sapiens
  • Figure 34.45 Art, a human hallmark
  • Transcript of "Chapter 34 Presentation"

    1. 1. Chapter 34 Vertebrates
    2. 2. Overview: Half a Billion Years of Backbones <ul><li>Early in the Cambrian period, about 530 million years ago, an astonishing variety of animals inhabited Earth’s oceans </li></ul><ul><li>One type of animal gave rise to vertebrates, one of the most successful groups of animals </li></ul>
    3. 3. Fig. 34-1
    4. 4. <ul><li>The animals called vertebrates get their name from vertebrae, the series of bones that make up the backbone </li></ul><ul><li>There are about 52,000 species of vertebrates, including the largest organisms ever to live on the Earth </li></ul><ul><li>Vertebrates have great disparity , a wide range of differences within the group </li></ul>
    5. 5. Concept 34.1: Chordates have a notochord and a dorsal, hollow nerve cord <ul><li>Vertebrates are a subphylum within the phylum Chordata </li></ul><ul><li>Chordates are bilaterian animals that belong to the clade of animals known as Deuterostomia </li></ul><ul><li>Two groups of invertebrate deuterostomes, the urochordates and cephalochordates, are more closely related to vertebrates than to other invertebrates </li></ul>
    6. 6. Fig. 34-2 Lobed fins Legs Amniotic egg Milk Jaws, mineralized skeleton Lungs or lung derivatives Vertebral column Head Notochord Common ancestor of chordates ANCESTRAL DEUTERO- STOME Echinodermata (sister group to chordates) Chondrichthyes (sharks, rays, chimaeras) Cephalochordata (lancelets) Urochordata (tunicates) Myxini (hagfishes) Petromyzontida (lampreys) Mammalia (mammals) Actinopterygii (ray-finned fishes) Actinistia (coelacanths) Amphibia (frogs, salamanders) Dipnoi (lungfishes) Reptilia (turtles, snakes, crocodiles, birds) Chordates Craniates Vertebrates Gnathostomes Lobe-fins Osteichthyans Tetrapods Amniotes
    7. 7. Fig. 34-2a Lobed fins Legs Amniotic egg Milk Jaws, mineralized skeleton Lungs or lung derivatives Vertebral column Head Notochord Common ancestor of chordates Chondrichthyes (sharks, rays, chimaeras) Cephalochordata (lancelets) Urochordata (tunicates) Myxini (hagfishes) Petromyzontida (lampreys) Mammalia (mammals) Actinopterygii (ray-finned fishes) Actinistia (coelacanths) Amphibia (frogs, salamanders) Dipnoi (lungfishes) Reptilia (turtles, snakes, crocodiles, birds) Chordates Craniates Vertebrates Gnathostomes Lobe-fins Osteichthyans Tetrapods Amniotes
    8. 8. Fig. 34-2b Lobed fins Legs Amniotic egg Milk Jaws, mineralized skeleton Lungs or lung derivatives Vertebral column Head Chondrichthyes (sharks, rays, chimaeras) Myxini (hagfishes) Petromyzontida (lampreys) Mammalia (mammals) Actinopterygii (ray-finned fishes) Actinistia (coelacanths) Amphibia (frogs, salamanders) Dipnoi (lungfishes) Reptilia (turtles, snakes, crocodiles, birds) Craniates Vertebrates Gnathostomes Lobe-fins Osteichthyans Tetrapods Amniotes
    9. 9. Fig. 34-2c Lobed fins Legs Amniotic egg Milk Jaws, mineralized skeleton Lungs or lung derivatives Vertebral column Chondrichthyes (sharks, rays, chimaeras) Petromyzontida (lampreys) Mammalia (mammals) Actinopterygii (ray-finned fishes) Actinistia (coelacanths) Amphibia (frogs, salamanders) Dipnoi (lungfishes) Reptilia (turtles, snakes, crocodiles, birds) Vertebrates Gnathostomes Lobe-fins Osteichthyans Tetrapods Amniotes
    10. 10. Fig. 34-2d Lobed fins Legs Amniotic egg Milk Jaws, mineralized skeleton Lungs or lung derivatives Chondrichthyes (sharks, rays, chimaeras) Mammalia (mammals) Actinopterygii (ray-finned fishes) Actinistia (coelacanths) Amphibia (frogs, salamanders) Dipnoi (lungfishes) Reptilia (turtles, snakes, crocodiles, birds) Gnathostomes Lobe-fins Osteichthyans Tetrapods Amniotes
    11. 11. Fig. 34-2e Lobed fins Legs Amniotic egg Milk Lungs or lung derivatives Mammalia (mammals) Actinopterygii (ray-finned fishes) Actinistia (coelacanths) Amphibia (frogs, salamanders) Dipnoi (lungfishes) Reptilia (turtles, snakes, crocodiles, birds) Lobe-fins Osteichthyans Tetrapods Amniotes
    12. 12. Fig. 34-2f Lobed fins Legs Amniotic egg Milk Mammalia (mammals) Actinistia (coelacanths) Amphibia (frogs, salamanders) Dipnoi (lungfishes) Reptilia (turtles, snakes, crocodiles, birds) Lobe-fins Tetrapods Amniotes
    13. 13. Fig. 34-2g Legs Amniotic egg Milk Mammalia (mammals) Amphibia (frogs, salamanders) Reptilia (turtles, snakes, crocodiles, birds) Tetrapods Amniotes
    14. 14. Fig. 34-2h Amniotic egg Milk Mammalia (mammals) Reptilia (turtles, snakes, crocodiles, birds) Amniotes
    15. 15. Derived Characters of Chordates <ul><li>All chordates share a set of derived characters </li></ul><ul><li>Some species have some of these traits only during embryonic development </li></ul><ul><li>Four key characters of chordates: </li></ul><ul><ul><li>Notochord </li></ul></ul><ul><ul><li>Dorsal, hollow nerve cord </li></ul></ul><ul><ul><li>Pharyngeal slits or clefts </li></ul></ul><ul><ul><li>Muscular, post-anal tail </li></ul></ul>
    16. 16. Fig. 34-3 Dorsal, hollow nerve cord Anus Muscular, post-anal tail Pharyngeal slits or clefts Notochord Mouth Muscle segments
    17. 17. Notochord <ul><li>The notochord is a longitudinal, flexible rod between the digestive tube and nerve cord </li></ul><ul><li>It provides skeletal support throughout most of the length of a chordate </li></ul><ul><li>In most vertebrates, a more complex, jointed skeleton develops, and the adult retains only remnants of the embryonic notochord </li></ul>
    18. 18. Dorsal, Hollow Nerve Cord <ul><li>The nerve cord of a chordate embryo develops from a plate of ectoderm that rolls into a tube dorsal to the notochord </li></ul><ul><li>The nerve cord develops into the central nervous system: the brain and the spinal cord </li></ul>
    19. 19. Pharyngeal Slits or Clefts <ul><li>In most chordates, grooves in the pharynx called pharyngeal clefts develop into slits that open to the outside of the body </li></ul><ul><li>Functions of pharyngeal slits : </li></ul><ul><ul><li>Suspension-feeding structures in many invertebrate chordates </li></ul></ul><ul><ul><li>Gas exchange in vertebrates (except vertebrates with limbs, the tetrapods) </li></ul></ul><ul><ul><li>Develop into parts of the ear, head, and neck in tetrapods </li></ul></ul>
    20. 20. Muscular, Post-Anal Tail <ul><li>Chordates have a tail posterior to the anus </li></ul><ul><li>In many species, the tail is greatly reduced during embryonic development </li></ul><ul><li>The tail contains skeletal elements and muscles </li></ul><ul><li>It provides propelling force in many aquatic species </li></ul>
    21. 21. Lancelets <ul><li>Lancelets (Cephalochordata) are named for their bladelike shape </li></ul><ul><li>They are marine suspension feeders that retain characteristics of the chordate body plan as adults </li></ul>
    22. 22. Fig. 34-UN1 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia
    23. 23. Fig. 34-4 Dorsal, hollow nerve cord Notochord Tail Cirri Mouth Pharyngeal slits Digestive tract Atrium Atriopore Segmental muscles Anus 2 cm
    24. 24. Tunicates <ul><li>Tunicates (Urochordata) are more closely related to other chordates than are lancelets </li></ul><ul><li>They are marine suspension feeders commonly called sea squirts </li></ul><ul><li>As an adult, a tunicate draws in water through an incurrent siphon, filtering food particles </li></ul>
    25. 25. Fig. 34-UN2 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia
    26. 26. Fig. 34-5 Tunic Water flow Excurrent siphon Atrium An adult tunicate Pharynx with slits Anus Atrium Excurrent siphon Incurrent siphon to mouth Dorsal, hollow nerve cord Incurrent siphon Excurrent siphon Muscle segments Notochord Tail Stomach Intestine Intestine Esophagus Stomach Pharynx with slits A tunicate larva
    27. 27. <ul><li>Tunicates most resemble chordates during their larval stage, which may last only a few minutes </li></ul>
    28. 28. Early Chordate Evolution <ul><li>Ancestral chordates may have resembled lancelets </li></ul><ul><li>Genome sequencing of tunicates has identified genes shared by tunicates and vertebrates </li></ul><ul><li>Gene expression in lancelets holds clues to the evolution of the vertebrate form </li></ul>
    29. 29. Fig. 34-6 BF1 Brain of vertebrate embryo (shown straightened) Hindbrain Forebrain Midbrain Nerve cord of lancelet embryo BF1 Hox3 Otx Otx Hox3
    30. 30. Concept 34.2: Craniates are chordates that have a head <ul><li>The origin of a head opened up a completely new way of feeding for chordates: active predation </li></ul><ul><li>Craniates share some characteristics: a skull, brain, eyes, and other sensory organs </li></ul>
    31. 31. Derived Characters of Craniates <ul><li>Craniates have two clusters of Hox genes; lancelets and tunicates have only one cluster </li></ul><ul><li>One feature unique to craniates is the neural crest , a collection of cells near the dorsal margins of the closing neural tube in an embryo </li></ul><ul><li>Neural crest cells give rise to a variety of structures, including some of the bones and cartilage of the skull </li></ul>
    32. 32. Fig. 34-7 Migrating neural crest cells Notochord Dorsal edges of neural plate Neural crest Neural tube
    33. 33. <ul><li>In aquatic craniates the pharyngeal clefts evolved into gill slits </li></ul><ul><li>Craniates have a higher metabolism and are more muscular than tunicates and lancelets </li></ul><ul><li>Craniates have a heart with at least two chambers, red blood cells with hemoglobin, and kidneys </li></ul>
    34. 34. The Origin of Craniates <ul><li>Fossils from the Cambrian explosion 530 million years ago document the transition to craniates </li></ul><ul><li>The most primitive of the fossils are those of the 3-cm-long Haikouella </li></ul><ul><li>Haikouella had a well-formed brain, eyes, and muscular segments, but not a skull </li></ul>
    35. 35. Fig. 34-8 Segmented muscles Pharyngeal slits 5 mm
    36. 36. Fig. 34-8a 5 mm
    37. 37. Fig. 34-8b Segmented muscles Pharyngeal slits
    38. 38. <ul><li>In other Cambrian rocks, paleontologists have found fossils of even more advanced chordates, such as Myllokunmingia </li></ul><ul><li>Myllokunmingia had a skull and was a true craniate </li></ul>
    39. 39. Hagfishes <ul><li>The least derived surviving craniate lineage is Myxini, the hagfishes </li></ul><ul><li>Hagfishes have a cartilaginous skull and axial rod of cartilage derived from the notochord, but lack jaws and vertebrae </li></ul>
    40. 40. Fig. 34-UN3 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia
    41. 41. Fig. 34-9 Slime glands
    42. 42. Concept 34.3: Vertebrates are craniates that have a backbone <ul><li>During the Cambrian period, a lineage of craniates evolved into vertebrates </li></ul><ul><li>Vertebrates became more efficient at capturing food and avoiding being eaten </li></ul>
    43. 43. Derived Characters of Vertebrates <ul><li>Vertebrates underwent a second gene duplication involving the Dlx family of transcription factors </li></ul><ul><li>Vertebrates have the following derived characters: </li></ul><ul><ul><li>Vertebrae enclosing a spinal cord </li></ul></ul><ul><ul><li>An elaborate skull </li></ul></ul><ul><ul><li>Fin rays, in the aquatic forms </li></ul></ul>
    44. 44. Lampreys <ul><li>Lampreys (Petromyzontida) represent the oldest living lineage of vertebrates </li></ul><ul><li>They are jawless vertebrates inhabiting various marine and freshwater habitats </li></ul><ul><li>They have cartilaginous segments surrounding the notochord and arching partly over the nerve cord </li></ul>
    45. 45. Fig. 34-UN4 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia
    46. 46. Fig. 34-10
    47. 47. Fig. 34-10a
    48. 48. Fig. 34-10b
    49. 49. Fossils of Early Vertebrates <ul><li>Conodonts were the first vertebrates with mineralized skeletal elements in their mouth and pharynx </li></ul>
    50. 50. Fig. 34-11 Dental elements
    51. 51. <ul><li>Other armored, jawless vertebrates had defensive plates of bone on their skin </li></ul>
    52. 52. Fig. 34-12 Pteraspis Pharyngolepis
    53. 53. Origins of Bone and Teeth <ul><li>Mineralization appears to have originated with vertebrate mouthparts </li></ul><ul><li>The vertebrate endoskeleton became fully mineralized much later </li></ul>
    54. 54. Concept 34.4: Gnathostomes are vertebrates that have jaws <ul><li>Today, jawed vertebrates, or gnathostomes , outnumber jawless vertebrates </li></ul>
    55. 55. Derived Characters of Gnathostomes <ul><li>Gnathostomes have jaws that might have evolved from skeletal supports of the pharyngeal slits </li></ul>
    56. 56. Fig. 34-13-1 Skeletal rods Cranium Gill slits Mouth
    57. 57. Fig. 34-13-2 Skeletal rods Cranium Gill slits Mouth
    58. 58. Fig. 34-13-3 Skeletal rods Cranium Gill slits Mouth
    59. 59. <ul><li>Other characters common to gnathostomes: </li></ul><ul><ul><li>An additional duplication of Hox genes </li></ul></ul><ul><ul><li>An enlarged forebrain associated with enhanced smell and vision </li></ul></ul><ul><ul><li>In aquatic gnathostomes, the lateral line system , which is sensitive to vibrations </li></ul></ul>
    60. 60. Fossil Gnathostomes <ul><li>The earliest gnathostomes in the fossil record are an extinct lineage of armored vertebrates called placoderms </li></ul>
    61. 61. Fig. 34-14 0.5 m
    62. 62. <ul><li>Another group of jawed vertebrates called acanthodians radiated during the Devonian period </li></ul>
    63. 63. Chondrichthyans (Sharks, Rays, and Their Relatives) <ul><li>Chondrichthyans (Chondrichthyes) have a skeleton composed primarily of cartilage </li></ul><ul><li>The cartilaginous skeleton evolved secondarily from an ancestral mineralized skeleton </li></ul><ul><li>The largest and most diverse group of chondrichthyans includes the sharks, rays, and skates </li></ul>Video: Shark Eating Seal Video: Manta Ray
    64. 64. Fig. 34-UN5 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia
    65. 65. Fig. 34-15 Pelvic fins Pectoral fins (c) Spotted ratfish ( Hydrolagus colliei ) (a) Blacktip reef shark ( Carcharhinus       melanopterus ) (b) Southern stingray ( Dasyatis americana )
    66. 66. Fig. 34-15a Pelvic fins Pectoral fins (a) Blacktip reef shark ( Carcharhinus melanopterus )
    67. 67. Fig. 34-15b (b) Southern stingray ( Dasyatis americana )
    68. 68. <ul><li>A second subclass is composed of a few dozen species of ratfishes </li></ul>
    69. 69. Fig. 34-15c (c) Spotted ratfish ( Hydrolagus colliei )
    70. 70. <ul><li>Most sharks </li></ul><ul><ul><li>Have a streamlined body and are swift swimmers </li></ul></ul><ul><ul><li>Are carnivores </li></ul></ul><ul><ul><li>Have a short digestive tract; a ridge called the spiral valve increases the digestive surface area </li></ul></ul><ul><ul><li>Have acute senses </li></ul></ul>
    71. 71. <ul><li>Shark eggs are fertilized internally but embryos can develop in different ways: </li></ul><ul><ul><li>Oviparous : eggs hatch outside the mother’s body </li></ul></ul><ul><ul><li>Ovoviviparous : the embryo develops within the uterus and is nourished by the egg yolk </li></ul></ul><ul><ul><li>Viviparous : the embryo develops within the uterus and is nourished through a yolk sac placenta from the mother’s blood </li></ul></ul>
    72. 72. <ul><li>The reproductive tract, excretory system, and digestive tract empty into a common cloaca </li></ul>
    73. 73. Ray-Finned Fishes and Lobe-Fins <ul><li>The vast majority of vertebrates belong to a clade of gnathostomes called Osteichthyes </li></ul><ul><li>Osteichthyes includes the bony fish and tetrapods </li></ul>
    74. 74. <ul><li>Nearly all living osteichthyans have a bony endoskeleton </li></ul><ul><li>Aquatic osteichthyans are the vertebrates we informally call fishes </li></ul><ul><li>Most fishes breathe by drawing water over gills protected by an operculum </li></ul><ul><li>Fishes control their buoyancy with an air sac known as a swim bladder </li></ul>
    75. 75. Fig. 34-UN6 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia
    76. 76. Fig. 34-16 Intestine Adipose fin (characteristic of trout) Cut edge of operculum Swim bladder Caudal fin Lateral line Urinary bladder Pelvic fin Anus Dorsal fin Spinal cord Brain Nostril Gills Kidney Heart Liver Gonad Anal fin Stomach
    77. 77. Ray-Finned Fishes <ul><li>Class Actinopterygii, the ray-finned fishes , includes nearly all the familiar aquatic osteichthyans </li></ul><ul><li>The fins, supported mainly by long, flexible rays, are modified for maneuvering, defense, and other functions </li></ul>Video: Seahorse Camouflage Video: Clownfish and Anemone Video: Coral Reef
    78. 78. Fig. 34-17 (a) Yellowfin tuna ( Thunnus albacares ) (b) Clownfish ( Amphiprion ocellaris ) (c) Sea horse ( Hippocampus       ramulosus ) (d) Fine-spotted moray eel ( Gymnothorax dovii )
    79. 79. Fig. 34-17a (a) Yellowfin tuna ( Thunnus albacares )
    80. 80. Fig. 34-17b (b) Clownfish ( Amphiprion ocellaris )
    81. 81. Fig. 34-17c (c) Sea horse ( Hippocampus ramulosus )
    82. 82. Fig. 34-17d (d) Fine-spotted moray eel ( Gymnothorax dovii )
    83. 83. Lobe-Fins <ul><li>The lobe-fins (Sarcopterygii) have muscular pelvic and pectoral fins </li></ul><ul><li>Three lineages survive and include coelacanths, lungfishes, and tetrapods </li></ul>
    84. 84. Fig. 34-18
    85. 85. Concept 34.5: Tetrapods are gnathostomes that have limbs <ul><li>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 </li></ul>
    86. 86. Derived Characters of Tetrapods <ul><li>Tetrapods have some specific adaptations: </li></ul><ul><ul><li>Four limbs, and feet with digits </li></ul></ul><ul><ul><li>Ears for detecting airborne sounds </li></ul></ul>
    87. 87. The Origin of Tetrapods <ul><li>In one lineage of lobe-fins, the fins became progressively more limb-like while the rest of the body retained adaptations for aquatic life </li></ul><ul><li>For example, Acanthostega lived in Greenland 365 million years ago </li></ul>
    88. 88. Fig. 34-19 Tetrapod limb skeleton Bones supporting gills
    89. 89. <ul><li>Extraordinary fossil discoveries over the past 20 years have allowed paleontologists to reconstruct the origin of tetrapods </li></ul>
    90. 90. Fig. 34-20 Ray-finned fishes Coelacanths Lungfishes Eusthenopteron Panderichthys Tiktaalik Elginerpeton Metaxygnathus Acanthostega Ichthyostega Hynerpeton Amphibians Greerpeton Amniotes PALEOZOIC Carboniferous Silurian Devonian Permian 0 265 280 295 310 325 340 355 370 385 400 415 430 Time (millions of years ago)
    91. 91. Amphibians <ul><li>Amphibians (class Amphibia) are represented by about 6,150 species of organisms in three orders </li></ul><ul><li>Order Urodela includes salamanders, which have tails </li></ul>
    92. 92. Fig. 34-UN7 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia
    93. 93. Fig. 34-21 (a) Order Urodela (b) Order Anura (c) Order Apoda
    94. 94. Fig. 34-21a (a) Order Urodela
    95. 95. <ul><li>Order Anura includes frogs and toads, which lack tails </li></ul>
    96. 96. Fig. 34-21b (b) Order Anura
    97. 97. <ul><li>Order Apoda includes caecilians, which are legless and resemble worms </li></ul>
    98. 98. Fig. 34-21c (c) Order Apoda
    99. 99. <ul><li>Amphibian means “both ways of life,” referring to the metamorphosis of an aquatic larva into a terrestrial adult </li></ul><ul><li>Most amphibians have moist skin that complements the lungs in gas exchange </li></ul><ul><li>Fertilization is external in most species, and the eggs require a moist environment </li></ul>
    100. 100. Fig. 34-22 (c) Mating adults (a) Tadpole (b) During metamorphosis
    101. 101. Fig. 34-22a (a) Tadpole
    102. 102. Fig. 34-22b (b) During metamorphosis
    103. 103. Fig. 34-22c (c) Mating adults
    104. 104. Fig. 34-23
    105. 105. Concept 34.6: Amniotes are tetrapods that have a terrestrially adapted egg <ul><li>Amniotes are a group of tetrapods whose living members are the reptiles, including birds, and mammals </li></ul>
    106. 106. Fig. 34-24 Reptiles ANCESTRAL AMNIOTE Diapsids Synapsids Lepidosaurs Archosaurs Dinosaurs Saurischians Mammals Squamates Tuataras Plesiosaurs Ichthyosaurs Birds Saurischian dinosaurs other than birds Ornithischian dinosaurs Pterosaurs Crocodilians Turtles Parareptiles
    107. 107. Derived Characters of Amniotes <ul><li>Amniotes are named for the major derived character of the clade, the amniotic egg , which contains membranes that protect the embryo </li></ul><ul><li>The extraembryonic membranes are the amnion, chorion, yolk sac, and allantois </li></ul>
    108. 108. Fig. 34-25 Yolk sac Amniotic cavity with amniotic fluid Chorion Amnion Albumen Yolk (nutrients) Allantois Embryo Shell
    109. 109. <ul><li>Amniotes have other terrestrial adaptations, such as relatively impermeable skin and the ability to use the rib cage to ventilate the lungs </li></ul>
    110. 110. Early Amniotes <ul><li>Living amphibians and amniotes split from a common ancestor about 370 million years ago </li></ul><ul><li>Early amniotes were more tolerant of dry conditions than early tetrapods </li></ul>
    111. 111. Reptiles <ul><li>The reptile clade includes the tuataras, lizards, snakes, turtles, crocodilians, birds, and the extinct dinosaurs </li></ul><ul><li>Reptiles have scales that create a waterproof barrier </li></ul><ul><li>They lay shelled eggs on land </li></ul>
    112. 112. Fig. 34-UN8 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia
    113. 113. Fig. 34-26
    114. 114. <ul><li>Most reptiles are ectothermic , absorbing external heat as the main source of body heat </li></ul><ul><li>Birds are endothermic , capable of keeping the body warm through metabolism </li></ul>
    115. 115. The Origin and Evolutionary Radiation of Reptiles <ul><li>The oldest reptilian fossils date to about 310 million years ago </li></ul><ul><li>The first major group to emerge were parareptiles , which were mostly large, stocky herbivores </li></ul>
    116. 116. <ul><li>As parareptiles were dwindling, the diapsids were diversifying </li></ul><ul><li>The diapsids consisted of two main lineages: the lepidosaurs and the archosaurs </li></ul><ul><li>The lepidosaurs include tuataras, lizards, and snakes </li></ul><ul><li>The archosaur lineage produced the crocodilians, pterosaurs , and dinosaurs </li></ul>
    117. 117. <ul><li>The dinosaurs diversified into a vast range of shapes and sizes </li></ul><ul><li>They included bipedal carnivores called theropods </li></ul><ul><li>Fossil discoveries and research have led to the conclusion that many dinosaurs were agile and fast moving </li></ul><ul><li>Paleontologists have also discovered signs of parental care among dinosaurs </li></ul>
    118. 118. <ul><li>Dinosaurs, with the exception of birds, became extinct by the end of the Cretaceous </li></ul><ul><li>Their extinction may have been partly caused by an asteroid </li></ul>
    119. 119. Lepidosaurs <ul><li>One surviving lineage of lepidosaurs is represented by two species of lizard-like reptiles called tuataras </li></ul>
    120. 120. Fig. 34-27 (a) Tuatara ( Sphenodon punctatus ) (c) Wagler’s pit viper ( Tropidolaemus wagleri ) (b) Australian thorny devil lizard ( Moloch horridus ) (e) American alligator ( Alligator mississippiensis ) (d) Eastern box turtle ( Terrapene carolina carolina )
    121. 121. Fig. 34-27a (a) Tuatara ( Sphenodon punctatus )
    122. 122. <ul><li>The other major living lineage of lepidosaurs consists of the squamates, the lizards and snakes </li></ul><ul><li>Lizards are the most numerous and diverse reptiles, apart from birds </li></ul>Video: Galápagos Marine Iguana
    123. 123. Fig. 34-27b (b) Australian thorny devil lizard ( Moloch horridus )
    124. 124. <ul><li>Snakes are legless lepidosaurs that evolved from lizards </li></ul>Video: Snake Ritual Wrestling
    125. 125. Fig. 34-27c (c) Wagler’s pit viper ( Tropidolaemus wagleri )
    126. 126. Turtles <ul><li>Turtles are the most distinctive group of reptiles alive today </li></ul><ul><li>All turtles have a boxlike shell made of upper and lower shields that are fused to the vertebrae, clavicles, and ribs </li></ul><ul><li>Some turtles have adapted to deserts and others live entirely in ponds and rivers </li></ul>Video: Galápagos Tortoise
    127. 127. Fig. 34-27d (d) Eastern box turtle ( Terrapene carolina carolina )
    128. 128. Alligators and Crocodiles <ul><li>Crocodilians (alligators and crocodiles) belong to an archosaur lineage that dates back to the late Triassic </li></ul>
    129. 129. Fig. 34-27e (e) American alligator ( Alligator mississippiensis )
    130. 130. Birds <ul><li>Birds are archosaurs, but almost every feature of their reptilian anatomy has undergone modification in their adaptation to flight </li></ul>
    131. 131. <ul><li>Many characters of birds are adaptations that facilitate flight </li></ul><ul><li>The major adaptation is wings with keratin feathers </li></ul><ul><li>Other adaptations include lack of a urinary bladder, females with only one ovary, small gonads, and loss of teeth </li></ul>Derived Characters of Birds
    132. 132. Fig. 34-28 (a) Wing (b) Bone structure (c) Feather structure Finger 1 Finger 2 Finger 3 Palm Hook Vane Barbule Barb Shaft Wrist Forearm Shaft
    133. 133. <ul><li>Flight enhances hunting and scavenging, escape from terrestrial predators, and migration </li></ul><ul><li>Flight requires a great expenditure of energy, acute vision, and fine muscle control </li></ul>
    134. 134. <ul><li>Birds probably descended from small theropods, a group of carnivorous dinosaurs </li></ul><ul><li>By 150 million years ago, feathered theropods had evolved into birds </li></ul><ul><li>Archaeopteryx remains the oldest bird known </li></ul>The Origin of Birds
    135. 135. Fig. 34-29 Airfoil wing with contour feathers Toothed beak Wing claw Long tail with many vertebrae
    136. 136. <ul><li>Living birds belong to the clade Neornithes </li></ul><ul><li>Several groups of birds are flightless </li></ul><ul><ul><li>The ratites , order Struthioniformes </li></ul></ul><ul><ul><li>Penguins, order Sphenisciformes </li></ul></ul><ul><ul><li>Certain species of rails, ducks, and pigeons </li></ul></ul>Living Birds
    137. 137. <ul><li>The demands of flight have rendered the general body form of many flying birds similar to one another </li></ul><ul><li>Foot structure in birds shows considerable variation </li></ul>Video: Flapping Geese Video: Swans Taking Flight Video: Soaring Hawk
    138. 138. Fig. 34-30 (a) Emu (b) Mallards (c) Laysan albatrosses (d) Barn swallows
    139. 139. Fig. 34-30a (a) Emu
    140. 140. Fig. 34-30b (b) Mallards
    141. 141. Fig. 34-30c (c) Laysan albatrosses
    142. 142. Fig. 34-30d (d) Barn swallows
    143. 143. Concept 34.7: Mammals are amniotes that have hair and produce milk <ul><li>Mammals , class Mammalia, are represented by more than 5,300 species </li></ul>
    144. 144. Fig. 34-UN9 Cephalochordata Urochordata Myxini Petromyzontida Mammalia Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia
    145. 145. Derived Characters of Mammals <ul><li>Mammals have </li></ul><ul><ul><li>Mammary glands, which produce milk </li></ul></ul><ul><ul><li>Hair </li></ul></ul><ul><ul><li>A larger brain than other vertebrates of equivalent size </li></ul></ul><ul><ul><li>Differentiated teeth </li></ul></ul>
    146. 146. Early Evolution of Mammals <ul><li>Mammals evolved from synapsids in the late Triassic period </li></ul><ul><li>Two bones that formerly made up the jaw joint were incorporated into the mammalian middle ear </li></ul>
    147. 147. Fig. 34-31 (b) In mammals, the articular and quadrate bones are incorporated into the middle ear. (a) In Biarmosuchus, an early synapsid, the articular and quadrate bones formed the jaw joint. Middle ear Temporal fenestra Jaw joint Eardrum Present-day reptile Present-day mammal Malleus (articular) Incus (quadrate) Sound Stapes Inner ear Eardrum Middle ear Sound Inner ear Stapes Key Quadrate Articular Squamosal Dentary
    148. 148. <ul><li>By the early Cretaceous, the three living lineages of mammals emerged: monotremes, marsupials, and eutherians </li></ul><ul><li>Mammals did not undergo a significant adaptive radiation until after the Cretaceous </li></ul>
    149. 149. Monotremes <ul><li>Monotremes are a small group of egg-laying mammals consisting of echidnas and the platypus </li></ul>
    150. 150. Fig. 34-32
    151. 151. Marsupials <ul><li>Marsupials include opossums, kangaroos, and koalas </li></ul><ul><li>The embryo develops within a placenta in the mother’s uterus </li></ul><ul><li>A marsupial is born very early in its development </li></ul><ul><li>It completes its embryonic development while nursing in a maternal pouch called a marsupium </li></ul>
    152. 152. Fig. 34-33 (a) A young brushtail possum (b) Long-nosed bandicoot
    153. 153. Fig. 34-33a (a) A young brushtail possum
    154. 154. <ul><li>In some species, such as the bandicoot, the marsupium opens to the rear of the mother’s body </li></ul>
    155. 155. Fig. 34-33b (b) Long-nosed bandicoot
    156. 156. <ul><li>In Australia, convergent evolution has resulted in a diversity of marsupials that resemble the eutherians in other parts of the world </li></ul>
    157. 157. Fig. 34-34 Plantigale Marsupial mammals Eutherian mammals Marsupial mammals Eutherian mammals Marsupial mole Flying squirrel Sugar glider Deer mouse Mole Tasmanian devil Wombat Kangaroo Woodchuck Patagonian cavy Wolverine
    158. 158. Eutherians (Placental Mammals) <ul><li>Compared with marsupials, eutherians have a longer period of pregnancy </li></ul><ul><li>Young eutherians complete their embryonic development within a uterus, joined to the mother by the placenta </li></ul><ul><li>Molecular and morphological data give conflicting dates on the diversification of eutherians </li></ul>
    159. 159. Fig. 34-35a ANCESTRAL MAMMAL Monotremata Marsupialia Monotremes (5 species) Marsupials (324 species) Eutherians (5,010 species) Xenarthra Rodentia Lagomorpha Primates Dermoptera (flying lemurs) Scandentia (tree shrews) Carnivora Cetartiodactyla Perissodactyla Chiroptera Eulipotyphla Pholidota (pangolins) Proboscidea Sirenia Tubulidentata Hyracoidea Afrosoricida (golden moles and tenrecs) Macroscelidea (elephant shrews)
    160. 160. Fig. 34-35b
    161. 161. Fig. 34-35c
    162. 162. Fig. 34-35d
    163. 163. Fig. 34-35e
    164. 164. Fig. 34-35f
    165. 165. Fig. 34-35g
    166. 166. Fig. 34-35h
    167. 167. Video: Bat Licking Nectar Video: Bat Pollinating Agave Plant Video: Galápagos Sea Lion Video: Wolf Agonistic Behavior
    168. 168. Primates <ul><li>The mammalian order Primates includes lemurs, tarsiers, monkeys, and apes </li></ul><ul><li>Humans are members of the ape group </li></ul>
    169. 169. <ul><li>Most primates have hands and feet adapted for grasping </li></ul>Derived Characters of Primates
    170. 170. <ul><li>Other derived characters of primates: </li></ul><ul><ul><li>A large brain and short jaws </li></ul></ul><ul><ul><li>Forward-looking eyes close together on the face, providing depth perception </li></ul></ul><ul><ul><li>Complex social behavior and parental care </li></ul></ul><ul><ul><li>A fully opposable thumb (in monkeys and apes) </li></ul></ul>
    171. 171. <ul><li>There are three main groups of living primates: </li></ul><ul><ul><li>Lemurs, lorises, and pottos </li></ul></ul><ul><ul><li>Tarsiers </li></ul></ul><ul><ul><li>Anthropoids (monkeys and apes) </li></ul></ul>Living Primates
    172. 172. Fig. 34-36
    173. 173. <ul><li>The oldest known anthropoid fossils, about 45 million years old, indicate that tarsiers are more closely related to anthropoids than to lemurs </li></ul>
    174. 174. Fig. 34-37 Lemurs, lorises, and pottos Tarsiers New World monkeys Old World monkeys Gibbons Orangutans Gorillas Chimpanzees and bonobos Humans 0 10 20 30 40 50 60 Time (millions of years ago) ANCESTRAL PRIMATE Anthropoids
    175. 175. <ul><li>The first monkeys evolved in the Old World (Africa and Asia) </li></ul><ul><li>In the New World (South America), monkeys first appeared roughly 25 million years ago </li></ul><ul><li>New World and Old World monkeys underwent separate adaptive radiations during their many millions of years of separation </li></ul>
    176. 176. Fig. 34-38 (a) New World monkey (b) Old World monkey
    177. 177. Fig. 34-38a (a) New World monkey
    178. 178. Fig. 34-38b (b) Old World monkey
    179. 179. <ul><li>The other group of anthropoids consists of primates informally called apes </li></ul><ul><li>This group includes gibbons, orangutans, gorillas, chimpanzees, bonobos, and humans </li></ul><ul><li>Apes diverged from Old World monkeys about 20–25 million years ago </li></ul>Video: Chimp Agonistic Behavior Video: Chimp Cracking Nut Video: Gibbons Brachiating
    180. 180. Fig. 34-39 (e) Bonobos (a) Gibbon (d) Chimpanzees (b) Orangutan (c) Gorilla
    181. 181. Fig. 34-39a (a) Gibbon
    182. 182. Fig. 34-39b (b) Orangutan
    183. 183. Fig. 34-39c (c) Gorilla
    184. 184. Fig. 34-39d (d) Chimpanzees
    185. 185. Fig. 34-39e (e) Bonobos
    186. 186. Concept 34.8: Humans are mammals that have a large brain and bipedal locomotion <ul><li>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 </li></ul>
    187. 187. Derived Characters of Humans <ul><li>A number of characters distinguish humans from other apes: </li></ul><ul><ul><li>Upright posture and bipedal locomotion </li></ul></ul><ul><ul><li>Larger brains </li></ul></ul><ul><ul><li>Language capabilities and symbolic thought </li></ul></ul><ul><ul><li>The manufacture and use of complex tools </li></ul></ul><ul><ul><li>Shortened jaw </li></ul></ul><ul><ul><li>Shorter digestive tract </li></ul></ul>
    188. 188. The Earliest Hominins <ul><li>The study of human origins is known as paleoanthropology </li></ul><ul><li>Hominins (formerly called hominids) are more closely related to humans than to chimpanzees </li></ul><ul><li>Paleoanthropologists have discovered fossils of about 20 species of extinct hominins </li></ul>
    189. 189. Fig. 34-40 Homo erectus Homo habilis Homo sapiens Homo neanderthalensis ? Homo ergaster Paranthropus robustus Paranthropus boisei Australopithecus africanus Australopithecus garhi Australopithecus afarensis Sahelanthropus tchadensis Orrorin tugenensis Ardipithecus ramidus Australo- pithecus anamensis Kenyanthropus platyops Homo rudolfensis 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
    190. 190. <ul><li>Hominins originated in Africa about 6–7 million years ago </li></ul><ul><li>Early hominins had a small brain but probably walked upright </li></ul>
    191. 191. <ul><li>Two common misconceptions about early hominins: </li></ul><ul><ul><li>Thinking of them as chimpanzees </li></ul></ul><ul><ul><li>Imagining human evolution as a ladder leading directly to Homo sapiens </li></ul></ul>
    192. 192. Australopiths <ul><li>Australopiths are a paraphyletic assemblage of hominins living between 4 and 2 million years ago </li></ul><ul><li>Some species walked fully erect </li></ul><ul><li>“ Robust” australopiths had sturdy skulls and powerful jaws </li></ul><ul><li>“ Gracile” australopiths were more slender and had lighter jaws </li></ul>
    193. 193. Fig. 34-41 (c) An artist’s reconstruction of what A. afarensis may have looked like (a) Australopithecus afarensis skeleton (b) The Laetoli footprints
    194. 194. Fig. 34-41a (a) Australopithecus afarensis skeleton
    195. 195. Fig. 34-41b (b) The Laetoli footprints
    196. 196. Fig. 34-41c (c) An artist’s reconstruction of what A. afarensis may have looked like
    197. 197. Bipedalism <ul><li>Hominins began to walk long distances on two legs about 1.9 million years ago </li></ul>
    198. 198. Tool Use <ul><li>The oldest evidence of tool use, cut marks on animal bones, is 2.5 million years old </li></ul>
    199. 199. Early Homo <ul><li>The earliest fossils placed in our genus Homo are those of Homo habilis , ranging in age from about 2.4 to 1.6 million years </li></ul><ul><li>Stone tools have been found with H. habilis , giving this species its name, which means “handy man” </li></ul>
    200. 200. <ul><li>Homo ergaster was the first fully bipedal, large-brained hominid </li></ul><ul><li>The species existed between 1.9 and 1.5 million years ago </li></ul><ul><li>Homo ergaster shows a significant decrease in sexual dimorphism (a size difference between sexes) compared with its ancestors </li></ul>
    201. 201. <ul><li>Homo ergaster fossils were previously assigned to Homo erectus ; most paleoanthropologists now recognize these as separate species </li></ul>
    202. 202. Fig. 34-42
    203. 203. <ul><li>Homo erectus originated in Africa by 1.8 million years ago </li></ul><ul><li>It was the first hominin to leave Africa </li></ul>
    204. 204. Neanderthals <ul><li>Neanderthals, Homo neanderthalensis, lived in Europe and the Near East from 200,000 to 28,000 years ago </li></ul><ul><li>They were thick-boned with a larger brain, they buried their dead, and they made hunting tools </li></ul>
    205. 205. Fig. 34-43 Chimpanzees Chimpanzees European and other living humans Neanderthals Living Europeans Other living humans Neanderthal 1 Neanderthal 2 Hypothesis: Neanderthals gave rise to European humans. Expected phylogeny: RESULTS EXPERIMENT
    206. 206. Fig. 34-43a Chimpanzees Neanderthals Living Europeans Other living humans Hypothesis: Neanderthals gave rise to European humans. Expected phylogeny: EXPERIMENT
    207. 207. Fig. 34-43b Chimpanzees European and other living humans Neanderthal 1 Neanderthal 2 RESULTS
    208. 208. Homo Sapiens <ul><li>Homo sapiens appeared in Africa by 195,000 years ago </li></ul><ul><li>All living humans are descended from these African ancestors </li></ul>
    209. 209. Fig. 34-44
    210. 210. <ul><li>The oldest fossils of Homo sapiens outside Africa date back about 115,000 years and are from the Middle East </li></ul><ul><li>Humans first arrived in the New World sometime before 15,000 years ago </li></ul><ul><li>In 2004, 18,000 year old fossils were found in Indonesia, and a new small hominin was named: Homo floresiensis </li></ul>
    211. 211. <ul><li>Rapid expansion of our species may have been preceded by changes to the brain that made cognitive innovations possible </li></ul><ul><ul><li>For example, the FOXP2 gene is essential for human language, and underwent intense natural selection during the last 200,000 years </li></ul></ul><ul><li>Homo sapiens were the first group to show evidence of symbolic and sophisticated thought </li></ul>
    212. 212. Fig. 34-45
    213. 213. Fig. 34-UN10
    214. 214. Fig. 34-UN10a
    215. 215. Fig. 34-UN10b
    216. 216. Fig. 34-UN10c
    217. 217. Fig. 34-UN10d
    218. 218. Fig. 34-UN10e
    219. 219. Fig. 34-UN10f
    220. 220. Fig. 34-UN10g
    221. 221. Fig. 34-T1
    222. 222. Fig. 34-UN11
    223. 223. Fig. 34-UN12
    224. 224. You should now be able to: <ul><li>List the derived traits for: chordates, craniates, vertebrates, gnathostomes, tetrapods, amniotes, birds, mammals, primates, humans </li></ul><ul><li>Explain what Haikouella and Myllokunmingia tell us about craniate evolution </li></ul><ul><li>Describe the trends in mineralized structures in early vertebrates </li></ul><ul><li>Describe and distinguish between Chondrichthyes and Osteichthyes, noting the main traits of each group </li></ul>
    225. 225. <ul><li>Define and distinguish among gnathostomes, tetrapods, and amniotes </li></ul><ul><li>Describe an amniotic egg and explain its significance in the evolution of reptiles and mammals </li></ul><ul><li>Explain why the reptile clade includes birds </li></ul><ul><li>Explain the significance of Archaeopteryx </li></ul><ul><li>Distinguish among monotreme, marsupial, and eutherian mammals </li></ul>
    226. 226. <ul><li>Define the term hominin </li></ul><ul><li>Describe the evolution of Homo sapiens from australopith ancestors, and clarify the order in which distinctive human traits arose </li></ul><ul><li>Explain the significance of the FOXP2 gene </li></ul>
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