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CAMPBELL BIOLOGY TENTH EDITION Reece • Urry • Cain • Wasserman • Minorsky • Jackson 34 The Origin and Evolution of Vertebrates Lecture Presentation by Nicole Tunbridge and Kathleen Fitzpatrick © 2014 Pearson Education, Inc. Half a Billion Years of Backbones Early in the Cambrian period, about 530 million years ago, an astonishing variety of invertebrate animals inhabited Earth’s oceans One type of animal gave rise to vertebrates, one of the most successful groups of animals The animals called vertebrates get their name from vertebrae, the series of bones that make up the backbone © 2014 Pearson Education, Inc. Figure 34.1 What is the relationship between this ancient organism and humans? © 2014 Pearson Education, Inc. One lineage of vertebrates colonized land 365 million years ago They gave rise to modern amphibians, reptiles (including birds), and mammals There are more than 57,000 species of vertebrates, including the largest organisms ever to live on Earth Vertebrates have great disparity, a wide range of differences within the group © 2014 Pearson Education, Inc. The Chordates 1. Chordates (phylum Chordata) are euterostomes, and have an internal skeleton, with muscles attached to the outer surface. A. Characteristics of Chordates 1. At some time during their life, all chordates have four basic characteristics. a. Notochord This supporting rod is located dorsally just below the nerve cord. It provides support and is replaced by the vertebral column in vertebrates. Dorsal tubular nerve cord This cord contains a fluid-filled canal. In vertebrates, this is the spinal cord and it is protected by vertebrae. Pharyngeal Pouches These openings function in feeding, gas exchange, or both. They are seen only during embryonic development in most vertebrates. In invertebrate chordates, fish, and amphibian larvae, they become functioning i. ii. b. i. ii. c. i. ii. iii. gills. iv. In terrestrial vertebrates, the pouches are modified for various purposes. v. In humans, the first pair of pouches becomes the auditory tubes, the second become tonsils, and the third and fourth pairs become the thymus and parathyroid glands. d. A postanal tail extends beyond the anus; in some this only appears in embryos. © 2014 Pearson Education, Inc. Concept 34.1: Chordates have a notochord and a dorsal, hollow nerve cord Chordates (phylum Chordata) are bilaterian animals that belong to the clade of animals known as Deuterostomia Chordates comprise all vertebrates and two groups of invertebrates, the urochordates and cephalochordates © 2014 Pearson Education, Inc. Phylogeny of living chordates Figure 34.2 Echinodermata ANCESTRAL DEUTEROSTOME Chordates Cephalochordata Urochordata Notochord Common ancestor of chordates Vertebrates Myxini Petromyzontida Jaws, mineralized skeleton Lobe-fins Actinistia Lungs or lung derivatives Dipnoi Lobed fins Amniotic egg Mammalia Milk © 2014 Pearson Education, Inc. Amniotes Reptilia Limbs with digits Tetrapods Amphibia Osteichthyans Actinopterygii Gnathostomes Chondrichthyes Vertebrae Figure 34.2a Phylogeny of living chordates (part 1) Echinodermata ANCESTRAL DEUTEROSTOME Cephalochordata Urochordata Notochord Common ancestor of chordates Vertebrae Jaws, mineralized skeleton © 2014 Pearson Education, Inc. Myxini Petromyzontida Chondrichthyes Figure 34.2b Phylogeny of living chordates (part 2) Chondrichthyes Actinopterygii Jaws, mineralized skeleton Actinistia Lungs or lung derivatives Dipnoi Lobed fins Amphibia Limbs with digits Reptilia Mammalia Amniotic egg Milk © 2014 Pearson Education, Inc. Derived Characters of Chordates All chordates share a set of derived characters Some species have some of these traits only during embryonic development Four key characters of chordates Notochord Dorsal, hollow nerve cord Pharyngeal slits or clefts Muscular, post-anal tail © 2014 Pearson Education, Inc. Figure 34.3 Chordate characteristics Notochord Dorsal, hollow nerve cord Muscle segments Mouth Anus Post-anal tail © 2014 Pearson Education, Inc. Pharyngeal slits or clefts Notochord The notochord is a longitudinal, flexible rod between the digestive tube and nerve cord It 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 © 2014 Pearson Education, Inc. 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 The nerve cord develops into the central nervous system: the brain and the spinal cord © 2014 Pearson Education, Inc. 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 Functions of pharyngeal slits Suspension-feeding structures in many invertebrate chordates Gas exchange in vertebrates (except vertebrates with limbs, the tetrapods) Develop into parts of the ear, head, and neck in tetrapods © 2014 Pearson Education, Inc. Muscular, Post-Anal Tail Chordates have a tail posterior to the anus In many species, the tail is greatly reduced during embryonic development The tail contains skeletal elements and muscles It provides propelling force in many aquatic species © 2014 Pearson Education, Inc. B. Nonvertebrate Chordates 1. Lancelets a. Lancelets are marine chordates, cephalochordates. b. Lancelets are named for their resemblance to a lancet—a two-edged surgical knife. c. They inhabit shallow coastal waters; they filter feed partly buried in sandy substrates. d. They feed on microscopic particles filtered from a constant stream of water that enters the mouth and exits through gill slits into an atrium that opens at the atriopore. e. Lancelets retain the four chordate characteristics as adults. f. The notochord extends from head to tail, accounting for their group name cephalochordates. g. They possess segmented muscles and the dorsal hollow nerve cord has periodic branches. 2. Sea Squirts a. Sea squirts, or urochordates, are also called tunicates. b. Adults have a body composed of an outer tunic; an excurrent siphon squirts out water when it is disturbed. c. The larvae are bilaterally symmetrical and have the four chordate characteristics. d. The larvae undergo metamorphosis to develop into sessile adults. e. Water passes into a pharynx and out numerous gill slits, the only chordate characteristic that remains in adults. f. It is hypothesized that sea squirts are directly related to vertebrates. A larva with the four chordate characteristics may have become sexually mature without developing the other sea squirt characteristics, and then evolved into a fishlike vertebrate. © 2014 Pearson Education, Inc. Lancelets Lancelets (Cephalochordata) are named for their bladelike shape They are marine suspension feeders that retain characteristics of the chordate body plan as adults © 2014 Pearson Education, Inc. Figure 34.UN01 Cephalochordata mini-tree Cephalochordata Urochordata Myxini Petromyzontida Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia Mammalia © 2014 Pearson Education, Inc. Figure 34.4 1 cm The lancelet Branchiostoma , a cephalochordate Cirri Notochord Mouth Dorsal, hollow nerve cord Pharyngeal slits Atrium Digestive tract Atriopore Segmental muscles Anus Tail © 2014 Pearson Education, Inc. Figure 34.4a 1 cm The lancelet Branchiostoma, a cephalochordate (part 1: photo) © 2014 Pearson Education, Inc. Tunicates Tunicates (Urochordata) are more closely related to other chordates than are lancelets Tunicates most resemble chordates during their larval stage, which may last only a few minutes As an adult, a tunicate draws in water through an incurrent siphon, filtering food particles When attacked, tunicates, or “sea squirts,” shoot water through their excurrent siphon © 2014 Pearson Education, Inc. Figure 34.UN02 Urochordata mini-tree Cephalochordata Urochordata Myxini Petromyzontida Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia Mammalia © 2014 Pearson Education, Inc. Figure 34.5 A tunicate, a urochordate Water flow Notochord Incurrent siphon to mouth Excurrent siphon Dorsal, hollow nerve cord Tail Excurrent siphon Incurrent siphon Muscle segments Intestine Stomach Atrium Pharynx with slits (a) A tunicate larva © 2014 Pearson Education, Inc. Excurrent siphon Anus Intestine Esophagus Atrium Pharynx with numerous slits Tunic Stomach (b) An adult tunicate (c) An adult tunicate Figure 34.5a A tunicate, a urochordate (part 1: art) Water flow Notochord Incurrent siphon to mouth Dorsal, hollow nerve cord Tail Excurrent siphon Incurrent siphon Muscle segments Intestine Stomach Atrium Pharynx with slits (a) A tunicate larva © 2014 Pearson Education, Inc. Excurrent siphon Anus Intestine Esophagus Stomach (b) An adult tunicate Atrium Pharynx with numerous slits Tunic Figure 34.5b A tunicate, a urochordate (part 2: photo Incurrent siphon to mouth Excurrent siphon Atrium Pharynx with numerous slits Tunic (c) An adult tunicate © 2014 Pearson Education, Inc. Tunicates are highly derived and have fewer Hox genes than other vertebrates © 2014 Pearson Education, Inc. Early Chordate Evolution Ancestral chordates may have resembled lancelets The same Hox genes that organize the vertebrate brain are expressed in the lancelet’s simple nerve cord tip Sequencing of the tunicate genome indicates that Genes associated with the heart and thyroid are common to all chordates Genes associated with transmission of nerve impulses are unique to vertebrates © 2014 Pearson Education, Inc. Figure 34.6 Myxini and Petromyzontida mini-tree BF1 Otx Hox3 Nerve cord of lancelet embryo BF1 Otx Hox3 Brain of vertebrate embryo (shown straightened) Forebrain Midbrain © 2014 Pearson Education, Inc. Hindbrain 29.2 The Vertebrates A. Characteristics of Vertebrates 1. As embryos, vertebrates have the four chordate characteristics. 2. Vertebrates also have these features: a. Vertebral column i. The embryonic notochord is replaced by a vertebral column. ii. Remnants of the notochord are seen in the intervertebral disks. iii. The vertebral column is part of a flexible, strong endoskeleton that is also evidence of segmentation. b. Skull i. A skull is an anterior component of the main axis of vertebrate endoskeleton; it encases the brain. ii. The high degree of cephalization in vertebrates is accompanied by complex sense organs. iii. The eyes developed as outgrowths of the brain. iv. The ears—equilibrium devices in water—function as sound-wave receivers in land vertebrates. c. Endoskeleton i. The endoskeleton and muscles together permit rapid and efficient movement. ii. The pectoral and pelvic fins of fish evolved into jointed appendages allowing vertebrates to move onto land. d. Internal organs i. Vertebrates possess a complete digestive system and a large coelom. ii. The circulatory system is closed and the blood is contained within blood vessels. iii. Gills or lungs provide efficient gas exchange. iv. The kidneys efficiently excrete nitrogenous waste and regulate water. v.Pearson Education, Reproduction is usually sexual with separate sexes. © 2014 Inc. Concept 34.2: Vertebrates are chordates that have a backbone A skeletal system and complex nervous system have allowed vertebrates efficiency at two essential tasks Capturing food Evading predators © 2014 Pearson Education, Inc. Derived Characters of Vertebrates Vertebrates have two or more sets of Hox genes; lancelets and tunicates have only one cluster Vertebrates have the following derived characters Vertebrae enclosing a spinal cord An elaborate skull Fin rays, in the aquatic forms © 2014 Pearson Education, Inc. Hagfishes and Lampreys Fossil evidence shows that the earliest vertebrates lacked jaws Only two lineages of jawless vertebrates remain today: the hagfishes and the lampreys Members of these groups lack a backbone The presence of rudimentary vertebrae and the results of phylogenetic analysis indicate that both hagfishes and lampreys are vertebrates © 2014 Pearson Education, Inc. Together, the hagfishes and lampreys form a clade of living jawless vertebrates, the cyclostomes Vertebrates with jaws make up a much larger clade, the gnathostomes © 2014 Pearson Education, Inc. Figure 34.UN03 Myxini and Petromyzontida mini-tree Cephalochordata Urochordata Myxini Petromyzontida Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia Mammalia © 2014 Pearson Education, Inc. Hagfishes Hagfishes (Myxini) are jawless vertebrates that have a cartilaginous skull, reduced vertebrae, and a flexible rod of cartilage derived from the notochord They have a small brain, eyes, ears, and tooth-like formations Hagfishes are marine; most are bottom-dwelling scavengers © 2014 Pearson Education, Inc. Figure 34.7 A hagfish Slime glands © 2014 Pearson Education, Inc. Lampreys Lampreys (Petromyzontida) are parasites that feed by clamping their mouth onto a live fish They inhabit various marine and freshwater habitats They have cartilaginous segments surrounding the notochord and arching partly over the nerve cord © 2014 Pearson Education, Inc. Figure 34.8 A sea lamprey © 2014 Pearson Education, Inc. Figure 34.8a A sea lamprey (part 1: lamprey) © 2014 Pearson Education, Inc. Figure 34.8b A sea lamprey (part 2: lamprey mouth) © 2014 Pearson Education, Inc. B. Vertebrate Evolution 1. Chordates (including vertebrates) appeared at the start of the Cambrian period. 2. The earliest vertebrates were fishes; most of which have jaws. 3. Jawed fish and other vertebrates are gnathostomes— animals with jaws. 4. Fish also had a bony skeleton, lungs, and fleshy fins, which were preadaptive for a land existence. 5. Amphibians are the first vertebrates to live on land and to have four limbs (tetrapods). 6. Amphibians are not fully adapted to living on land because they still have to reproduce in an aquatic environments. 7. Reptiles are fully adapted to living on land because they produce an amniotic egg. 8. Amniotes develop within an aquatic environment but of their own making. 9. In placental mammals, the fertilized egg develops inside the female, where it is surrounded by an amniotic membrane. 10. Another feature for living on land includes watertight skin, and can be seen in reptiles and mammals. © 2014 Pearson Education, Inc. Early Vertebrate Evolution Fossils from the Cambrian explosion document the transition to craniates The most primitive of the fossils are those of the 3cm-long Haikouella Haikouella had a well-formed brain, eyes, and muscular segments, but no skull or ear organs © 2014 Pearson Education, Inc. Fossil of an early chordate Figure 34.9 5 mm Segmented muscles Pharyngeal slits © 2014 Pearson Education, Inc. Figure 34.9a Fossil of an early chordate (part 1: photo) 5 mm © 2014 Pearson Education, Inc. The Fishes 1. Fishes are the largest group of vertebrates with nearly 28,000 recognized species. A. Jawless Fishes 1. Small, jawless, and finless ostracoderms are the earliest vertebrate fossils. 2. Today’s jawless fishes, or agnathans have a cartilaginous skeleton and persistent notochord. 3. They have smooth nonscaly skin. 4. They have cylindrical bodies and are up to a meter long. 5. Many lampreys are filter feeders similar to their ancestors. 6. Parasitic lampreys have a round muscular mouth equipped with teeth; they attach themselves to fish and suck nutrients from the host’s circulatory system. 7. Marine parasitic lampreys entered the Great Lakes and devastated the trout population in the 1950s. © 2014 Pearson Education, Inc. Conodonts were among the earliest vertebrates in the fossil record, dating from 500 to 200 million years ago They had mineralized skeletal elements in their mouth and pharynx Their fossilized dental elements are common in the fossil record © 2014 Pearson Education, Inc. Figure 34.10 A conodont Dental elements (within head) 0.5 cm © 2014 Pearson Education, Inc. Other groups of jawless vertebrates were armored with defensive plates of bone on their skin © 2014 Pearson Education, Inc. Figure 34.11 Jawless armored vertebrates Pteraspis Pharyngolepis © 2014 Pearson Education, Inc. Origins of Bone and Teeth Mineralization appears to have originated with vertebrate mouthparts The vertebrate endoskeleton became fully mineralized much later © 2014 Pearson Education, Inc. B. Fishes with Jaws 1. Fishes with jaws have: a. Ectothermy—they depend on the environment to regulate their body temperature. b. Gills are used for gas exchange. i. Jawed fish have a single-looped, closed circulatory pathway with a heart that pumps the blood first to the gills (for oxygen exchange) and then to the rest of the body. c. Cartilaginous or bony endoskeleton—the endoskeleton of jawed fishes includes the vertebral column, a skull with jaws, and paired pectoral and pelvic fins. i. Jaws evolved from the first pair of gill arches of agnathans; the second pair of arches became support structures for the jaws. d. Scales cover and protect the skin. 2. Placoderms are extinct jawed fishes of the Devonian Period. a. They were armored with heavy plates and had strong jaws. b. Like extant fishes, they had paired pectoral and pelvic fins. c. Paired fins allow a fish to balance and maneuver well in water; this helps predation. 3. Cartilaginous Fishes a. Sharks, rays, skates, and chimaeras are marine cartilaginous fishes (Chondrichthyes). b. They have a cartilaginous skeleton rather than bone. c. Five to seven gill slits are on both sides of the pharynx; they lack the gill covers found on bony fish. d. They have many openings to the gill chamber located behind the eyes called spiracles. e. Their body is covered with dermal denticles. f. The teeth of sharks are enlarged scales; there are many rows of replacement teeth growing behind the front teeth. g. They have three well developed senses to detect prey: i. The ability to sense electric currents in water; ii. A lateral line system senses pressure caused by fish swimming nearby and; iii. They have a keen sense of smell. h. The largest sharks are filter feeders, not predators; the basking and whale sharks eat tons of crustacea. i. Most sharks are fast, open-sea predators; a great white shark eats dolphins, sea lions and seals. © 2014 Pearson Education, Inc. Concept 34.3: Gnathostomes are vertebrates that have jaws Today, jawed vertebrates, or gnathostomes, outnumber jawless vertebrates Gnathostomes include sharks and their relatives, ray-finned fishes, lobe-finned fishes, amphibians, reptiles (including birds), and mammals © 2014 Pearson Education, Inc. Derived Characters of Gnathostomes Gnathostomes (“jaw mouth”) are named for their jaws, hinged structures that, especially with the help of teeth, are used to grip food items firmly and slice them The jaws are hypothesized to have evolved by modification of skeletal rods that supported the pharyngeal (gill) slits © 2014 Pearson Education, Inc. Figure 34.12 Possible step in the evolution of jawbones Gill slits Skeletal rods © 2014 Pearson Education, Inc. Cranium Modified skeletal rods Other characters common to gnathostomes Genome duplication, including duplication of Hox genes An enlarged forebrain associated with enhanced smell and vision In aquatic gnathostomes, the lateral line system, which is sensitive to vibrations © 2014 Pearson Education, Inc. Fossil Gnathostomes The earliest gnathostomes in the fossil record are an extinct lineage of armored vertebrates called placoderms They appeared about 440 million years ago © 2014 Pearson Education, Inc. Figure 34.13 Fossil of an early gnathostome…Dunkleosteus could exert a force of 8000 pounds per square inch. 0.5 m © 2014 Pearson Education, Inc. Another group of jawed vertebrates called acanthodians radiated during the Silurian and Devonian periods (444 to 359 million years ago) Three lineages of jawed vertebrates survive today: chondrichthyans, ray-finned fishes, and lobe-fins © 2014 Pearson Education, Inc. Chondrichthyans (Sharks, Rays, and Their Relatives) Chondrichthyans (Chondrichthyes) have a skeleton composed primarily of cartilage The largest and most diverse group of chondrichthyans includes the sharks, rays, and skates © 2014 Pearson Education, Inc. Figure 34.UN04 Chondrichthyes mini-tree, Cephalochordata Urochordata Myxini Petromyzontida Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia Mammalia © 2014 Pearson Education, Inc. Figure 34.14 Dorsal fins Pectoral fins Pelvic fins (a) Blacktip reef shark (Carcharhinus melanopterus) (c) Spotted ratfish (Hydrolagus colliei) © 2014 Pearson Education, Inc. (b) Southern stingray (Dasyatis americana) Figure 34.14a Dorsal fins Pectoral fins Pelvic fins (a) Blacktip reef shark (Carcharhinus melanopterus) © 2014 Pearson Education, Inc. j. Rays and skates live on the ocean floor; their pectoral fins are enlarged into winglike fins and they swim slowly. k. Stingrays have a venomous spine. l. Electric rays feed on fish that have been stunned with an electric shock that may reach over 300 volts. m. Sawfish rays have a large anterior “saw” that they use to slash through schools of fish. n. Chimaeras (ratfishes) live in cold marine waters and are known for their unusual shape and iridescent colors. © 2014 Pearson Education, Inc. Figure 34.14b (b) Southern stingray (Dasyatis americana) © 2014 Pearson Education, Inc. Figure 34.14c (c) Spotted ratfish (Hydrolagus colliei) © 2014 Pearson Education, Inc. Video: Manta Ray © 2014 Pearson Education, Inc. A second subclass is composed of a few dozen species of ratfishes, or chimaeras © 2014 Pearson Education, Inc. Sharks have a streamlined body and are swift swimmers The largest sharks are suspension feeders, but most are carnivores Sharks have a short digestive tract with a ridge called the spiral valve to increase the digestive surface area Sharks have acute senses including sight, smell, and the ability to detect electrical fields from nearby animals © 2014 Pearson Education, Inc. Shark eggs are fertilized internally but embryos can develop in different ways Oviparous: Eggs hatch outside the mother’s body Ovoviviparous: The embryo develops within the uterus and is nourished by the egg yolk Viviparous: The embryo develops within the uterus and is nourished through a yolk sac placenta from the mother’s blood © 2014 Pearson Education, Inc. The reproductive tract, excretory system, and digestive tract empty into a common cloaca Today, sharks are severely threatened by overfishing; Pacific populations have plummeted by up to 95% © 2014 Pearson Education, Inc. Ray-Finned Fishes and Lobe-Fins The vast majority of vertebrates belong to a clade of gnathostomes called Osteichthyes Nearly all living osteichthyans have a bony endoskeleton Osteichthyans include the bony fishes and tetrapods Aquatic osteichthyans are the vertebrates we informally call fishes © 2014 Pearson Education, Inc. Figure 34.UN05 Actinopterygii, Actinistia, and Dipnoi mini-tree Cephalochordata Urochordata Myxini Petromyzontida Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia Mammalia © 2014 Pearson Education, Inc. Most fishes breathe by drawing water over gills protected by an operculum Fishes control their buoyancy with an air sac known as a swim bladder Fishes have a lateral line system Most species are oviparous, but some have internal fertilization and birthing © 2014 Pearson Education, Inc. 4. Bony Fishes a. There are about 25,000 species of bony fishes (Osteichthyes). b. Bony fishes have a skeleton of bone; most are ray-finned with thin, bony rays supporting the fins. c. The gills of bony fishes do not open separately but instead are covered by an operculum. d. The swim bladder is a gas-filled sac whose pressure can be altered to regulate buoyancy and depth. e. Bony fish have a single-loop, closed cardiovascular system. f. They have a well-developed nervous system. g. Fish sperm and eggs are usually shed into water. h. For most fish, the fertilization and embryonic development occur outside the female’s body. i. The lobe-finned fishes include six species of lungfishes and two species of coelacanth. i. Their fleshy fins are supported by central bones. ii. Lungfishes have lungs and gills for gas exchange. iii. Lungfishes and lobe-finned fishes are grouped together as the Sarcopterygii. iv. Lungfishes live in stagnant freshwater or in ponds that dry up annually; they are found in Africa, South America, and Australia. v. Coelacanths live in deep oceans; once considered extinct, more than 200 have been captured since 1938. © 2014 Pearson Education, Inc. Figure 34.15 Anatomy of a trout, a ray-finned fish Cut edge of operculum Swim Spinal cord bladder Dorsal fin Brain Adipose fin (characteristic of trout) Caudal fin Nostril Anal fin Gills Kidney © 2014 Pearson Education, Inc. Liver Heart Stomach Anus Gonad Pelvic Intestine fin Lateral line Urinary bladder Ray-Finned Fishes Actinopterygii, the ray-finned fishes, include nearly all the familiar aquatic osteichthyans Ray-finned fishes originated during the Silurian period (444 to 416 million years ago) The fins, supported mainly by long, flexible rays, are modified for maneuvering, defense, and other functions © 2014 Pearson Education, Inc. ▼ Figure 34.16 Red lionfish (Pterois volitans) Yellowfin tuna (Thunnus albacares) ▼ Ray-finned fishes (Actinopterygii) ▼ ▼ © 2014 Pearson Education, Inc. Common sea horse, (Hippocampus ramulosus) Fine-spotted moray eel, (Gymnothorax dovii) Video: Clownfish and Anemone © 2014 Pearson Education, Inc. Video: Coral Reef © 2014 Pearson Education, Inc. Video: Seahorse Camouflage © 2014 Pearson Education, Inc. Industrial-scale fishing operations have driven many ray-finned fish populations to collapse Populations are also affected by dams that change water flow patterns, affecting prey capture, migration, and spawning © 2014 Pearson Education, Inc. Lobe-Fins The lobe-fins (Sarcopterygii) also originated in the Silurian period They have muscular pelvic and pectoral fins that they use to swim and “walk” underwater across the substrate © 2014 Pearson Education, Inc. Figure 34.17 A reconstruction of an ancient lobe-fin—discovered in 2009 5 cm Lower jaw © 2014 Pearson Education, Inc. Scaly covering Dorsal spine Figure 34.17a A reconstruction of an ancient lobe-fin (part 1: photo) 5 cm Lower jaw © 2014 Pearson Education, Inc. Scaly covering Dorsal spine Three lineages survive and include coelacanths, lungfishes, and tetrapods Coelacanths were thought to have become extinct 75 million years ago, but a living coelacanth was caught off the coast of South Africa in 1938 © 2014 Pearson Education, Inc. Figure 34.18 A coelacanth (Latimeria) © 2014 Pearson Education, Inc. The living lungfishes are all found in the Southern Hemisphere Though gills are the main organs for gas exchange, they can also surface to gulp air into their lungs The third surviving lineage of lobe-fins are tetrapods, a group that adapted to life on land © 2014 Pearson Education, Inc. 29.4 The Amphibians 1. Amphibians have the following characteristics: a. They are tetrapods; they have four limbs. i. The skeleton is well-developed for locomotion. b. Amphibians have smooth and nonscaly skin. i. The moist skin plays an active role in water balance, respiration, and temperature regulation. c. Amphibians usually have small lungs supplemented by gas exchange across porous skin. d. The single-loop circulatory path of fish is replaced by a closed double-loop circulatory system; however oxygen-rich blood mixes with some oxygen-poor blood. i. A three-chambered heart with a single ventricle pumps and two atria pumps blood to both the lungs and body. e. Amphibians have sense organs that are adapted to life on land. i. The brain is larger than that of fishes; their cerebral cortex is more developed. ii. A specialized tongue is used for catching prey. iii. The eyelids keep their eyes moist. iv. Amphibian ears are adapted for detecting sound waves; in turn, the larynx produces sounds. f. Amphibians are ectothermic, depending upon the environment to regulate body temperature. i. If winter temperature drops too low, temperate ectotherms become inactive and enter torpor. g. Amphibians return to the water to reproduce. i. They shed eggs into the water for external fertilization. ii. Generally, amphibian eggs are protected by a coat of jelly but not by a shell. iii. The young hatch into aquatic larvae with gills (tadpoles). iv. The aquatic larvae usually undergo metamorphosis to develop into a terrestrial adult. v. Some amphibians evolved mechanisms that allow them to bypass the aquatic larval stage. © 2014 Pearson Education, Inc. Concept 34.4: Tetrapods are gnathostomes that have limbs One of the most significant events in vertebrate history was when the fins of some lobe-fins evolved into the limbs and feet of tetrapods © 2014 Pearson Education, Inc. Derived Characters of Tetrapods Tetrapods have some specific adaptations Four limbs, and feet with digits A neck, which allows separate movement of the head Fusion of the pelvic girdle to the backbone The absence of gills (except some aquatic species) Ears for detecting airborne sounds © 2014 Pearson Education, Inc. A. Evolution of Amphibians 1. Amphibians evolved from the lobe-finned fishes with lungs by way of transitional forms. 2. Two hypotheses describe evolution of amphibians from lobe-finned fishes. a. Lobe-finned fishes that could move from pond-to-pond had an advantage over those that could not. b. The supply of food on land and the absence of predators promoted adaptation to land. 3. Paleontologists found a fossil, Tiktaalik roseae, from the late Devonian period in Arctic Canada that represents an intermediate between lobe-finned fishes and tetrapods with limbs. B. Diversity of Living Amphibians 1. Modern amphibians include three groups: salamanders and newts, frogs and toads, and caecilians. 2. Salamanders and newts have a long body and tail, and two pairs of legs 3. Their S-shaped locomotion is similar to fish movements. 4. Salamanders and newts are carnivorous, feeding on insects, snails, etc. 5. Salamanders practice internal fertilization; the males produce a spermatophore that females pick up with the cloaca (the common receptacle for the urinary, genital, and digestive canals). 6. Frogs and toads are tailless as adults; the hind limbs are specialized for jumping. 7. Glands in the skin secrete poisons; some tropical species often have brilliant warning coloration. 8. Frogs and toads have the head and trunk fused; frogs live near or in fresh water while toads live in damp places away from water. 9. Caecilians are legless; most burrow in soil and feed on worms, etc. © 2014 Pearson Education, Inc. The Origin of Tetrapods Tiktaalik, nicknamed a “fishapod,” shows both fish and tetrapod characteristics It had Fins, gills, lungs, and scales Ribs to breathe air and support its body A neck and shoulders Fins with the bone pattern of a tetrapod limb © 2014 Pearson Education, Inc. Figure 34.19 Fish Characters Tetrapod Characters Scales Fins Gills and lungs Neck Ribs Fin skeleton Flat skull Eyes on top of skull Discovery of a “fishapod”: Tiktaalik Shoulder bones Ribs Neck Scales Head Eyes on top of skull Humerus Flat skull Ulna “Wrist” Elbow Radius Fin © 2014 Pearson Education, Inc. Fin skeleton Figure 34.19a Shoulder bones Neck Head Eyes on top of skull Flat skull Fin © 2014 Pearson Education, Inc. Figure 34.19b Ribs © 2014 Pearson Education, Inc. Figure 34.19c Scales © 2014 Pearson Education, Inc. Figure 34.19d Humerus Ulna “Wrist” Elbow Radius Fin skeleton © 2014 Pearson Education, Inc. Tiktaalik could most likely prop itself on its fins, but not walk The first tetrapods appeared 365 million years ago © 2014 Pearson Education, Inc. Figure 34.20 Lungfishes Steps in the origin of limbs with digits Eusthenopteron Panderichthys Tiktaalik Acanthostega Limbs with digits Tulerpeton Amphibians Amniotes Silurian Devonian PALEOZOIC Carboniferous Permian 415 400 385 370 355 340 325 310 295 280 265 0 Time (millions of years ago) © 2014 Pearson Education, Inc. Key to limb bones Ulna Radius Humerus Figure 34.20a Lungfishes Eusthenopteron Panderichthys Tiktaalik Lobe-fins with limbs with digits Silurian Devonian PALEOZOIC Carboniferous Permian 415 400 385 370 355 340 325 310 295 280 265 0 Time (millions of years ago) © 2014 Pearson Education, Inc. Key to limb bones Ulna Radius Humerus Figure 34.20b Acanthostega Limbs with digits Tulerpeton Amphibians Amniotes Silurian Devonian PALEOZOIC Carboniferous Permian 415 400 385 370 355 340 325 310 295 280 265 0 Time (millions of years ago) © 2014 Pearson Education, Inc. Key to limb bones Ulna Radius Humerus Amphibians Amphibians (class Amphibia) are represented by about 6,150 species in three clades Urodela (salamanders) Anura (frogs) Apoda (caecilians) © 2014 Pearson Education, Inc. Figure 34.UN06 Amphibia mini-tree Cephalochordata Urochordata Myxini Petromyzontida Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia Mammalia © 2014 Pearson Education, Inc. Salamanders Salamanders (urodeles) are amphibians with tails Some are aquatic, but others live on land as adults or throughout life Paedomorphosis, the retention of juvenile features in sexually mature organisms, is common in aquatic species © 2014 Pearson Education, Inc. Figure 34.21 (b) Order Anura (a) Order Urodela Amphibians (c) Order Apoda © 2014 Pearson Education, Inc. Figure 34.21a (a) Order Urodela © 2014 Pearson Education, Inc. Frogs Frogs (anurans) lack tails and have powerful hind legs for locomotion on land Frogs with leathery skin are called “toads” © 2014 Pearson Education, Inc. Figure 34.21b Amphibians (part 2: frogs and toads) (b) Order Anura © 2014 Pearson Education, Inc. Caecilians Caecilians (apoda) are legless, nearly blind, and resemble earthworms The absence of legs is a secondary adaptation © 2014 Pearson Education, Inc. Figure 34.21c Amphibians (part 3: caecilians) (c) Order Apoda © 2014 Pearson Education, Inc. Lifestyle and Ecology of Amphibians Amphibian means “both ways of life,” referring to the metamorphosis of an aquatic larva into a terrestrial adult Tadpoles are herbivores that lack legs, but legs, lungs, external eardrums, and adaptations for carnivory may all arise during metamorphosis Most amphibians have moist skin that complements the lungs in gas exchange © 2014 Pearson Education, Inc. The “dual life” of a frog (Rana temporaria) Figure 34.22 (a) The tadpole © 2014 Pearson Education, Inc. (b) During metamorphosis (c) The adults return to water to mate. Figure 34.22a (a) The tadpole © 2014 Pearson Education, Inc. Figure 34.22b (b) During metamorphosis © 2014 Pearson Education, Inc. Figure 34.22c (c) The adults return to water to mate. © 2014 Pearson Education, Inc. Fertilization is external in most species, and the eggs require a moist environment In some species, males or females care for the eggs on their back, in their mouth, or in their stomach © 2014 Pearson Education, Inc. Figure 34.23 A mobile nursery © 2014 Pearson Education, Inc. Amphibian populations have been declining in recent decades The causes include a disease-causing chytrid fungus, habitat loss, climate change, and pollution © 2014 Pearson Education, Inc. The Reptiles 1. Reptiles (class Reptilia) are a successful group of terrestrial animals. 2. Reptiles have many characteristics showing that they are fully adapted to living on land. a. Reptiles have paired limbs adapted for climbing, running, paddling, or flying. b. Reptiles have a thick, scaly skin that is impermeable to water. i. Reptile’s protective skin prevents water loss but it also requires several molts a year. 3.Reptiles have efficient breathing - their lungs are more developed than in amphibians; air rhythmically moves in and out of the lungs due to an expandable rib cage. 4.Reptiles have efficient circulation. The heart prevents mixing of blood. Oxygen-rich blood is more fully separated from oxygen-poor blood. 5.Reptiles have efficient excretion. The kidneys are well-developed and excrete uric acid so less water is required. 6.Reptiles are ectothermic. i. They require a fraction of the food per body weight of birds and mammals. ii. They are behaviorally adapted to warm their body temperature by sunbathing. Reptiles have well-adapted reproduction. i. The sexes are separate and fertilization is internal. ii. The amniotic egg contains extraembryonic membranes. iii. Extraembryonic membranes are not a part of the embryo and are disposed of after development. iv. They protect the embryo, remove nitrogenous wastes, and provide oxygen, food, and water. v. The amnion is one extraembryonic membrane; it fills with fluid to provide a “pond” for the embryo to develop. © 2014 Pearson Education, Inc. Concept 34.5: Amniotes are tetrapods that have a terrestrially adapted egg Amniotes are a group of tetrapods whose living members are the reptiles, including birds, and mammals © 2014 Pearson Education, Inc. †Parareptiles Turtles Crocodilians Archosaurs Reptiles †Pterosaurs †Ornithischian ANCESTRAL AMNIOTE dinosaurs Saurischians Diapsids Dinosaurs A phylogeny of amniotes Figure 34.24 †Saurischian dinosaurs other than birds Birds †Plesiosaurs †Ichthyosaurs Lepidosaurs Synapsids © 2014 Pearson Education, Inc. Tuataras Squamates (lizards and snakes) Mammals Figure 34.24a †Parareptiles Turtles †Pterosaurs †Ornithischian Saurischians Diapsids Dinosaurs Archosaurs Reptiles © 2014 Pearson Education, Inc. Crocodilians dinosaurs † Saurischian dinosaurs other than birds Birds Figure 34.24b Diapsids †Plesiosaurs †Ichthyosaurs Lepidosaurs Synapsids © 2014 Pearson Education, Inc. Tuataras Squamates (lizards and snakes) Mammals A. Evolution of Amniotes 1. The amniotes consist of three lineages: a. The turtles, in which the skull is anapsid, having no openings behind the orbit—eye socket. b. All the other reptiles, in which the skull has two openings behind the orbit (diapsid). c. The mammals, in which the skull has one opening behind the orbit (synapsid). 2. The reptiles have no common ancestor; they are a paraphyletic group and not a monophyletic group. 3. Thecodonts are diapsids that gave rise to the ichthyosaurs, which returned to the aquatic environment, and the pterosaurs, which were terrestrial. a. The pterosaurs of the Jurassic period had a keel for attachment of flight muscles and air spaces in bones to reduce weight. b. The thecodonts gave rise to the crocodiles and dinosaurs. 4. Dinosaurs varied in size and behavior; some had a bipedal stance and gave rise to birds. 5. Dinosaurs dominated the earth for about 170 million years; then most died out at the end of the Cretaceous period (65 MYA). 6. One theory of mass extinction: a. A massive meteorite struck the Earth near the Yucatán Peninsula, resulting in cataclysmic events disrupting existing ecosystems, destroying many living things. b. An iridium layer, a mineral common in meteorites, occurs in rocks at the end of this period. © 2014 Pearson Education, Inc. Derived Characters of Amniotes Amniotes are named for the major derived character of the clade, the amniotic egg, which contains membranes that protect the embryo The extraembryonic membranes are the amnion, chorion, yolk sac, and allantois © 2014 Pearson Education, Inc. Figure 34.25 The amniotic egg Extraembryonic membranes Allantois Chorion Amniotic cavity with amniotic fluid Embryo Yolk (nutrients) Albumen Shell Amnion Yolk sac Extraembryonic membranes © 2014 Pearson Education, Inc. The amniotic egg was a key adaptation to life on land The amniotic eggs of most reptiles and some mammals have a shell Amniotes have other terrestrial adaptations, such as relatively impermeable skin and the ability to use the rib cage to ventilate the lungs © 2014 Pearson Education, Inc. Early Amniotes Living amphibians and amniotes split from a common ancestor about 350 million years ago Early amniotes were more tolerant of dry conditions than the first tetrapods The earliest amniotes were small predators with sharp teeth and long jaws © 2014 Pearson Education, Inc. Figure 34.26 Artist’s reconstruction of Hylonomus, an early amniote © 2014 Pearson Education, Inc. Reptiles The reptile clade includes the tuataras, lizards, snakes, turtles, crocodilians, birds, and some extinct groups Reptiles have scales that create a waterproof barrier Most reptiles lay shelled eggs on land © 2014 Pearson Education, Inc. Figure 34.UN07 Reptilia mini-tree Cephalochordata Urochordata Myxini Petromyzontida Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia Mammalia © 2014 Pearson Education, Inc. Figure 34.27 Hatching reptiles © 2014 Pearson Education, Inc. Most reptiles are ectothermic, absorbing external heat as the main source of body heat Ectotherms regulate their body temperature through behavioral adaptations Birds are endothermic, capable of maintaining body temperature through metabolism © 2014 Pearson Education, Inc. The Origin and Evolutionary Radiation of Reptiles Fossil evidence indicates that the earliest reptiles lived about 310 million years ago The first major group to emerge were parareptiles, which were mostly large, stocky quadrupedal herbivores © 2014 Pearson Education, Inc. As parareptiles were dwindling, the diapsids were diversifying The diapsids consisted of two main lineages: the lepidosaurs and the archosaurs The lepidosaurs include tuataras, lizards, snakes, and extinct mososaurs The archosaur lineage produced the crocodilians, pterosaurs, and dinosaurs © 2014 Pearson Education, Inc. Pterosaurs were the first tetrapods to exhibit flight The dinosaurs diversified into a vast range of shapes and sizes They included bipedal carnivores called theropods, the group from which birds are descended © 2014 Pearson Education, Inc. Fossil discoveries and research have led to the conclusion that many dinosaurs were agile and fast moving Paleontologists have also discovered signs of parental care among dinosaurs Some anatomical evidence supports the hypothesis that at least some dinosaurs were endotherms © 2014 Pearson Education, Inc. Dinosaurs, with the exception of birds, became extinct by the end of the Cretaceous Their extinction may have been partly caused by an asteroid © 2014 Pearson Education, Inc. C. Diversity of Living Reptiles 1. Most reptiles today live in the tropics or subtropics; lizards and snakes live on soil; turtles, crocodiles, and alligators live in water. 2. Turtles have a heavy shell fused to the ribs and to the thoracic vertebrae. a. Turtles lack teeth but use a sharp beak. b. Sea turtles must return to shore to lay eggs. 3. Lizards have four clawed legs and are carnivorous. a. Marine iguanas on the Galápagos Islands are adapted to spending time at sea. b. Chameleons live in trees, have a long sticky tongue to catch insects, and change color. c. Geckos are primarily nocturnal and have adhesive pads on their toes. d. Skinks have reduced limbs and shiny scales. 4. Snakes evolved from lizards and lost legs as an adaptation to burrowing. a. Their jaws can readily dislocate to engulf large food. b. A tongue collects airborne molecules to transfer them to Jacobson’s organ for tasting. c. Some snakes are poisonous and have special fangs to inject venom. d. Snakes have internal ears that can detect low-frequency sounds and vibrations. 5. Tuataras are lizardlike animals found in New Zealand. a. They possess a well-developed “third eye,” which is light sensitive and buried beneath the skin in the upper part of the head. b. They are the only member of an ancient group of reptiles that © 2014 Pearson Education, Inc. included the common ancestor of modern lizards and snakes. Turtles The phylogenetic position of turtles remains uncertain All turtles have a boxlike shell made of upper and lower shields that are fused to the vertebrae, clavicles, and ribs © 2014 Pearson Education, Inc. Figure 34.28 (a) Eastern box turtle (Terrapene carolina carolina) Extant reptiles (other than birds (d) Wagler’s pit viper (b) Tuatara (Sphenodon punctatus) (Tropidolaemus wagleri) (c) Australian thorny devil lizard (Moloch horridus) (e) American alligator (Alligator mississippiensis) © 2014 Pearson Education, Inc. Figure 34.28a (a) Eastern box turtle (Terrapene carolina carolina) © 2014 Pearson Education, Inc. Figure 34.28b (b) Tuatara (Sphenodon punctatus) © 2014 Pearson Education, Inc. Figure 34.28c (c) Australian thorny devil lizard (Moloch horridus) © 2014 Pearson Education, Inc. Figure 34.28d (d) Wagler’s pit viper (Tropidolaemus wagleri) © 2014 Pearson Education, Inc. Figure 34.28e (e) American alligator (Alligator mississippiensis) © 2014 Pearson Education, Inc. Video: Galápagos Marine Iguana © 2014 Pearson Education, Inc. Video: Snake Ritual Wrestling © 2014 Pearson Education, Inc. Video: Galápagos Tortoise © 2014 Pearson Education, Inc. Some turtles have adapted to deserts and others live entirely in ponds and rivers The largest turtles live in the sea Many species of sea turtles are endangered by accidental capture in fishing nets or development of beaches where they lay eggs © 2014 Pearson Education, Inc. Snakes are legless lepidosaurs that evolved from lizards Snakes are carnivorous, and have adaptations to aid in capture and consumption of prey including Chemical sensors Heat-detecting organs Venom Loosely articulated jawbones and elastic skin © 2014 Pearson Education, Inc. Crocodilians Crocodilians (alligators and crocodiles) belong to an archosaur lineage that dates back to the late Triassic Living crocodilians are restricted to warm regions © 2014 Pearson Education, Inc. D. Birds Birds share a common ancestor with crocodilians and have scales, a tail with vertebrae, and clawed feet. Feathers keep birds warm, and help birds fly and steer. Feathers are modified scales. Birds molt and replace their feathers annually. Birds have a modified skeleton. The collarbone is fused and the sternum has a keel. Other bones are fused to make the skeleton more rigid than the reptilian skeleton. The breast muscles are attached to the keel. Birds have modified respiration. Bird respiratory air sacs are extensive, even extending into some larger bones. i. Using a one-way flow of air, air sacs maximize gas exchange and oxygenation of blood. ii. Efficient supply of oxygen to muscles is vital for the level of muscle activity needed for flight. Birds are endothermic; they have the ability to maintain a constant, relatively high body temperature. Birds have well-developed sense organs and nervous system. Birds have very acute vision. Birds’ muscle reflexes are excellent. Bird migration allows use of widespread food sources; an enlarged portion of the brain is responsible for instinctive behaviors. E. Diversity of Living Birds Most birds can fly; some, however, some are flightless. Bird classification is based on beak and foot types, and to some extent on habitats and behaviors. Birds of prey have notched beaks and sharp talons. Shorebirds have long slender bills and long legs. Waterfowl have webbed toes and broad bills. © 2014 Pearson Education, Inc. Birds Birds are archosaurs, but almost every feature of their reptilian anatomy has undergone modification in their adaptation to flight © 2014 Pearson Education, Inc. Derived Characters of Birds Many characters of birds are adaptations that facilitate flight The major adaptation is wings with keratin feathers Other adaptations include lack of a urinary bladder, females with only one ovary, small gonads, and loss of teeth © 2014 Pearson Education, Inc. Figure 34.29 Form fits function: the avian wing and feather Finger 1 (b) Bone structure Palm Finger 2 (a) Wing Finger 3 Forearm Vane Shaft Wrist Shaft Barb Barbule Hook (c) Feather structure © 2014 Pearson Education, Inc. Figure 34.29a Finger 1 (b) Bone structure Palm (a) Wing Finger 2 Finger 3 Forearm Shaft Vane Wrist Shaft Barb Barbule Hook (c) Feather structure © 2014 Pearson Education, Inc. Figure 34.29c (b) Bone structure © 2014 Pearson Education, Inc. Video: Flapping Geese © 2014 Pearson Education, Inc. Video: Soaring Hawk © 2014 Pearson Education, Inc. Video: Swans Taking Flight © 2014 Pearson Education, Inc. Flight enhances hunting and scavenging, escape from terrestrial predators, and migration Flight requires a great expenditure of energy, acute vision, and fine muscle control © 2014 Pearson Education, Inc. The Origin of Birds Birds probably descended from small theropods, a group of carnivorous dinosaurs Early feathers might have evolved for insulation, camouflage, or courtship display © 2014 Pearson Education, Inc. By 160 million years ago, feathered theropods had evolved into birds Archaeopteryx remains the oldest bird known © 2014 Pearson Education, Inc. Figure 34.30 Was Archaeopteryx the first bird? Toothed beak Airfoil wing with contour feathers © 2014 Pearson Education, Inc. Long tail with many vertebrae Wing claw Living Birds Living birds belong to the clade Neornithes Several groups of birds are flightless The ratites, order Struthioniformes Penguins, order Sphenisciformes Certain species of rails, ducks, and pigeons © 2014 Pearson Education, Inc. Figure 34.31 An emu (Dromaius novaehollandiae), a flightless bird native to Australia © 2014 Pearson Education, Inc. Figure 34.32 A king penguin (Aptenodytes patagonicus) “flying” underwater © 2014 Pearson Education, Inc. The demands of flight have rendered the general body form of many flying birds similar to one another Bird species can be distinguished by characters including profile, color, flying style, behavior, beak shape, and foot structure © 2014 Pearson Education, Inc. Figure 34.33 Hummingbird feeding while hovering © 2014 Pearson Education, Inc. Figure 34.34 A specialized beak © 2014 Pearson Education, Inc. Figure 34.34a A specialized beak (part 1: greater flamingo) © 2014 Pearson Education, Inc. Figure 34.34b (part 2: greater flamingo beak) © 2014 Pearson Education, Inc. Figure 34.35 Feet adapted to perching © 2014 Pearson Education, Inc. The Mammals 1. The following characteristics distinguish mammals: a. Hair i. Hair provides insulation against heat loss. ii. Hair color can provide camouflage to blend into its surroundings. iii. Hair can serve sensory functions. b. Mammary glands i. Mammary glands enable females to feed young without deserting them to obtain food. ii. Nursing creates a bond between mother and offspring to ensure parental care while the young are helpless. c. Skeleton i. The mammal skull accommodates a larger brain relative to body size compared to reptiles’. ii. Their teeth are differentiated into molars and premolars. iii. The vertebral column provides more movement. d. Internal organs i. Gas exchange is efficiently accomplished by lungs. ii. Mammals possess a four-chambered heart and a double-loop circulatory system. iii. Kidneys are adapted to conserving water. iv. The nervous system and sensory organs are highly developed. e. Internal development i. In most mammals, the young are born alive after a period of development in the uterus. © 2014 Pearson Education, Inc. Concept 34.6: Mammals are amniotes that have hair and produce milk Mammals, class Mammalia, are represented by more than 5,300 species © 2014 Pearson Education, Inc. Figure 34.UN08 Mammalia mini-tree Cephalochordata Urochordata Myxini Petromyzontida Chondrichthyes Actinopterygii Actinistia Dipnoi Amphibia Reptilia Mammalia © 2014 Pearson Education, Inc. Derived Characters of Mammals Mammals have Mammary glands, which produce milk Hair A high metabolic rate, due to endothermy A larger brain than other vertebrates of equivalent size Differentiated teeth © 2014 Pearson Education, Inc. A. Evolution of Mammals 1. Mammals evolved during the Mesozoic Era from mammal-like synapsids. 2. True mammals appeared during the Triassic period, about the same time as the first dinosaurs. a. The first mammals were small, about the size of mice. b. Some of the earliest mammalian groups were monotremes and marsupials. c. Placental mammals evolved later to occupy habitats vacated by dinosaurs. 3. Monotremes a. Monotremes are mammals that have a cloaca and lay hardshelled amniote eggs. b. They are represented by the duckbill platypus and two species of the spiny anteaters. c. A female duckbill platypus lays her eggs in a burrow in the ground where she incubates them. d. After hatching, the young lick milk seeping from modified sweat glands on the abdomen. e. The spiny anteater has a pouch formed by swollen mammary glands and muscle; the egg moves from cloaca to pouch and hatches; the young remain for 53 days and live in the burrow where the mother feeds them. © 2014 Pearson Education, Inc. Early Evolution of Mammals Mammals are synapsids In the evolution of mammals from early synapsids, two bones that formerly made up the jaw joint were incorporated into the mammalian middle ear © 2014 Pearson Education, Inc. Figure 34.36 Biarmosuchus, an extinct synapsid The evolution of the mammalian ear bones Key Articular Quadrate Dentary Squamosal Temporal fenestra Jaw joint (a) Articular and quadrate bones in the jaw Middle ear Eardrum Stapes Inner ear Eardrum Middle ear Inner ear Stapes Sound Sound Incus (quadrate) Malleus (articular) Present-day reptile Present-day mammal (b) Articular and quadrate bones in the middle ear © 2014 Pearson Education, Inc. By the early Cretaceous, the three living lineages of mammals emerged: monotremes, marsupials, and eutherians Mammals did not undergo a significant adaptive radiation until after the Cretaceous © 2014 Pearson Education, Inc. Monotremes Monotremes are a small group of egg-laying mammals consisting of echidnas and the platypus © 2014 Pearson Education, Inc. Figure 34.37 Short-beaked echidna (Tachyglossus aculeatus), an Australian monotreme © 2014 Pearson Education, Inc. Figure 34.37a © 2014 Pearson Education, Inc. Figure 34.37b © 2014 Pearson Education, Inc. 4. Marsupials a. Marsupials begin development inside the mother’s body but are then born in a very immature state. b. The newborns crawl up into a pouch on their mother’s abdomen. c. Inside a pouch they attach to the nipples of the mother’s mammary glands and continue to develop. d. Today, most marsupials are found in Australia where they underwent adaptive radiation for several million years without competition from the placental mammals, only introduced recently. 5. Placental Mammals * Developing placental mammals are dependent on a placenta, an organ of exchange between maternal and fetal blood. * The placenta supplies nutrients to and removes wastes from the blood of developing offspring. * A placenta also allows a mother to move about while the offspring develop. * The placenta enables young to be born in a relatively advanced stage of development. * Placental mammals are very active animals; they possess acute senses and a relatively large brain. * The brains of placental animals have cerebral hemispheres proportionately larger than other animals. * The young go through a long period of dependency on their parents after birth. Most are terrestrial, but some are aquatic, and bats can fly. © 2014 Pearson Education, Inc. Marsupials Marsupials include opossums, kangaroos, and koalas The embryo develops within a placenta in the mother’s uterus A marsupial is born very early in its development It completes its embryonic development while nursing in a maternal pouch called a marsupium © 2014 Pearson Education, Inc. Australian marsupials Figure 34.38 (a) A young brushtail possum (b) A greater bilby © 2014 Pearson Education, Inc. Figure 34.38a (a) A young brushtail possum © 2014 Pearson Education, Inc. Figure 34.38b (b) A greater bilby © 2014 Pearson Education, Inc. In some species, such as the bandicoot, the marsupium opens to the rear of the mother’s body In Australia, convergent evolution has resulted in a diversity of marsupials that resemble the eutherians in other parts of the world © 2014 Pearson Education, Inc. Figure 34.39 Convergent evolution of marsupials and eutherians (placental mammals) Marsupial mammals Eutherian mammals © 2014 Pearson Education, Inc. Plantigale Deer mouse Marsupial mole Mole Sugar glider Wombat Tasmanian devil Kangaroo Flying squirrel Woodchuck Wolverine Patagonian cavy Figure 34.39a Marsupial mammals Eutherian mammals Plantigale Deer mouse Marsupial mole Mole Sugar glider © 2014 Pearson Education, Inc. Flying squirrel Figure 34.39b Marsupial mammals Wombat Tasmanian devil Kangaroo © 2014 Pearson Education, Inc. Eutherian mammals Woodchuck Wolverine Patagonian cavy Eutherians (Placental Mammals) Compared with marsupials, eutherians have a more complex placenta Young eutherians complete their embryonic development within a uterus, joined to the mother by the placenta Molecular and morphological data give conflicting dates on the diversification of eutherians © 2014 Pearson Education, Inc. Figure 34.40a Monotremes (5 species) Marsupials (324 species) ANCESTRAL MAMMAL Monotremata Marsupialia Eutherians (5,010 species) Exploring mammalian diversity (part 1: phylogenetic tree) © 2014 Pearson Education, Inc. Proboscidea Sirenia Tubulidentata Hyracoidea Afrosoricida Macroscelidea Xenarthra Rodentia Lagomorpha Primates Dermoptera Scandentia Carnivora Cetartiodactyla Perissodactyla Chiroptera Eulipotyphla Pholidota Figure 34.40aa Monotremes (5 species) Marsupials (324 species) ANCESTRAL MAMMAL Eutherians (5,010 species) © 2014 Pearson Education, Inc. Monotremata Marsupialia Proboscidea Sirenia Tubulidentata Hyracoidea Afrosoricida Macroscelidea Figure 34.40ab Exploring mammalian diversity (part 1b: phylogenetic tree Eutherians (5,010 species) Proboscidea Sirenia Tubulidentata Hyracoidea Afrosoricida Macroscelidea Xenarthra Rodentia Lagomorpha Primates Dermoptera Scandentia Carnivora Cetartiodactyla Perissodactyla Chiroptera Eulipotyphla Pholidota © 2014 Pearson Education, Inc. Figure 34.40b Orders and Examples Monotremata Main Characteristics Lay eggs; no nipples; young suck milk from fur of mother Platypuses, echidnas Orders and Examples Marsupialia Completes embryonic development in pouch on mother’s body Kangaroos, opossums, koalas Koala Echidna Proboscidea Long, muscular trunk; thick, loose skin; upper incisors elongated as tusks Elephants Tubulidentata Teeth consisting of many thin tubes cemented together; eats ants and termites Aardvarks African elephant Sirenia Aardvark Aquatic; finlike forelimbs and no hind limbs; herbivorous Manatees, dugongs Hyracoidea Short legs; stumpy tail; herbivorous; complex, multichambered stomach Hyraxes Manatee Xenarthra Rock hyrax Reduced teeth or no teeth; herbivorous (sloths) or carnivorous (anteaters, armadillos) Sloths, anteaters, armadillos Tamandua Lagomorpha Rabbits, hares, picas Rodentia Dogs, wolves, bears, cats, weasels, otters, seals, walruses Artiodactyls: sheep, pigs, cattle, deer, giraffes Primates Sharp, pointed canine teeth and molars for shearing; carnivorous Perissodactyla Opposable thumbs; forward-facing eyes; well-developed cerebral cortex; omnivorous Lemurs, monkeys, chimpanzees, gorillas, humans Golden lion tamarin Hooves with an odd number of toes on each foot; herbivorous Horses, zebras, tapirs, rhinoceroses Indian rhinoceros Hooves with an even number of toes on each foot; herbivorous Chiroptera Bats Frog-eating bat Bighorn sheep Cetaceans: whales, dolphins, porpoises Pacific whitesided porpoise © 2014 Pearson Education, Inc. Red squirrel Chisel-like incisors; hind legs longer than forelegs and adapted for running and jumping; herbivorous Coyote Cetartiodactyla Chisel-like, continuously growing incisors worn down by gnawing; herbivorous Squirrels, beavers, rats, porcupines, mice Jackrabbit Carnivora Main Characteristics Aquatic; streamlined body; paddle-like forelimbs and no hind limbs; thick layer of insulating blubber; carnivorous Eulipotyphla “Core insectivores”: some moles, some shrews Adapted for flight; broad skinfold that extends from elongated fingers to body and legs; carnivorous or herbivorous Eat mainly insects and other small invertebrates Star-nosed mole Figure 34.40ba Orders and Examples Main Characteristics Lay eggs; no nipples; young suck milk from fur of mother Monotremata Platypuses, echidnas Echidna Marsupialia Kangaroos, opossums, koalas Completes embryonic development in pouch on mother’s body Koala © 2014 Pearson Education, Inc. Figure 34.40bb Orders and Examples Main Characteristics Long, muscular trunk; thick, loose skin; upper incisors elongated as tusks Proboscidea Elephants African elephant Aquatic; finlike forelimbs and no hind limbs; herbivorous Sirenia Manatees, dugongs Manatee Teeth consisting of many thin tubes cemented together; eats ants and termites Tubulidentata Aardvarks Aardvark Hyracoidea Hyraxes Rock hyrax © 2014 Pearson Education, Inc. Short legs; stumpy tail; herbivorous; complex, multichambered stomach Figure 34.40bc Orders and Examples Xenarthra Sloths, anteaters, armadillos Tamandua Lagomorpha Rabbits, hares, picas Jackrabbit Main Characteristics Reduced teeth or no teeth; herbivorous (sloths) or carnivorous (anteaters, armadillos) Chisel-like incisors; hind legs longer than forelegs and adapted for running and jumping; herbivorous Rodentia Squirrels, beavers, rats, porcupines, mice Red squirrel Chisel-like, continuously growing incisors worn down by gnawing; herbivorous Primates Lemurs, monkeys, chimpanzees, gorillas, humans Golden lion tamarin Opposable thumbs; forward-facing eyes; well-developed cerebral cortex; omnivorous © 2014 Pearson Education, Inc. Figure 34.40bd Orders and Examples Carnivora Dogs, wolves, bears, cats, weasels, otters, seals, walruses Main Characteristics Sharp, pointed canine teeth and molars for shearing; carnivorous Coyote Cetartiodactyla Hooves with an even Artiodactyls: number of toes on each sheep, pigs, foot; herbivorous cattle, deer, Bighorn sheep giraffes Aquatic; streamlined Cetaceans: body; paddle-like whales, forelimbs and no hind dolphins, limbs; thick layer of porpoises Pacific whiteinsulating blubber; sided porpoise carnivorous © 2014 Pearson Education, Inc. Figure 34.40be Orders and Examples Main Characteristics Perissodactyla Hooves with an odd Horses, zebras, number of toes on tapirs, each foot; herbivorous rhinoceroses Indian rhinoceros Chiroptera Bats Adapted for flight; broad skinfold that extends from elongated fingers to body and Frog-eating bat legs; carnivorous or herbivorous Eulipotyphla “Core insectivores”: some moles, some shrews © 2014 Pearson Education, Inc. Eat mainly insects and other small invertebrates Star-nosed mole Video: Bat Licking Nectar © 2014 Pearson Education, Inc. Video: Bat Pollinating Agave Plant © 2014 Pearson Education, Inc. Video: Galápagos Sea Lion © 2014 Pearson Education, Inc. Video: Wolves Agonistic Behavior © 2014 Pearson Education, Inc. Video: Shark Eating a Seal © 2014 Pearson Education, Inc. Primates The mammalian order Primates includes lemurs, tarsiers, monkeys, and apes Humans are members of the ape group © 2014 Pearson Education, Inc. Derived Characters of Primates Most primates have hands and feet adapted for grasping, and flat nails © 2014 Pearson Education, Inc. Other derived characters of primates A large brain and short jaws Forward-looking eyes close together on the face, providing depth perception Complex social behavior and parental care A fully opposable thumb (in monkeys and apes) © 2014 Pearson Education, Inc. Living Primates There are three main groups of living primates Lemurs, lorises, and bush babies Tarsiers Anthropoids (monkeys and apes) © 2014 Pearson Education, Inc. Verreaux’s sifakas (Propithecus verreauxi), a type of lemur Figure 34.41 © 2014 Pearson Education, Inc. The oldest known anthropoid fossils, about 45 million years old, indicate that tarsiers are more closely related to anthropoids than to lemurs © 2014 Pearson Education, Inc. Figure 34.42 Lemurs, lorises, and bush babies ANCESTRAL PRIMATE New World monkeys Old World monkeys Gibbons Orangutans Gorillas Chimpanzees and bonobos Humans 70 60 50 40 30 20 10 Time (millions of years ago) © 2014 Pearson Education, Inc. 0 Anthropoids A phylogenetic tree of primates Tarsiers The first monkeys evolved in the Old World (Africa and Asia) In the New World (South America), monkeys first appeared roughly 25 million years ago New World and Old World monkeys underwent separate adaptive radiations during their many millions of years of separation © 2014 Pearson Education, Inc. Figure 34.43 (a) New World monkey: spider monkey (b) Old World monkey: macaque © 2014 Pearson Education, Inc. The other group of anthropoids consists of primates informally called apes This group includes gibbons, orangutans, gorillas, chimpanzees, bonobos, and humans Apes diverged from Old World monkeys about 25– 30 million years ago © 2014 Pearson Education, Inc. Figure 34.44 Nonhuman apes (a) Gibbon (b) Orangutan (c) Gorilla (d) Chimpanzees © 2014 Pearson Education, Inc. (e) Bonobos Video: Chimp Cracking Nut © 2014 Pearson Education, Inc. Concept 34.7: Humans are mammals that have a large brain and bipedal locomotion The species Homo sapiens is about 200,000 years old, which is very young, considering that life has existed on Earth for at least 3.5 billion years © 2014 Pearson Education, Inc. Derived Characters of Humans A number of characters distinguish humans from other apes Upright posture and bipedal locomotion Larger brains capable of language, symbolic thought, artistic expression, the manufacture and use of complex tools Reduced jawbones and jaw muscles Shorter digestive tract © 2014 Pearson Education, Inc. The human and chimpanzee genomes are 99% identical Changes in regulatory genes can have large effects © 2014 Pearson Education, Inc. The Earliest Hominins The study of human origins is known as paleoanthropology Hominins (formerly called hominids) are more closely related to humans than to chimpanzees Paleoanthropologists have discovered fossils of about 20 species of extinct hominins © 2014 Pearson Education, Inc. Figure 34.45 A timeline for selected hominin species Paranthropus robustus 0 Homo ergaster Paranthropus boisei 0.5 Homo Homo neanderthalensis sapiens ? 1.0 Australopithecus africanus 1.5 Millions of years ago 2.0 2.5 Kenyanthropus platyops Australopithecus garhi Australo3.0 pithecus anamensis 3.5 Homo habilis 4.0 4.5 Australopithecus afarensis 5.0 Ardipithecus ramidus 5.5 6.0 Homo erectus Orrorin tugenensis 6.5 7.0 © 2014 Pearson Education, Inc. Sahelanthropus tchadensis Homo rudolfensis Figure 34.45a Homo neander- Homo thalensis sapiens Paranthropus Homo robustus ergaster Millions of years ago 0 0.5 ? Paranthropus boisei 1.0 1.5 2.0 2.5 Australopithecus garhi 3.0 Australopithecus 3.5 africanus Homo erectus Homo habilis © 2014 Pearson Education, Inc. Homo rudolfensis Figure 34.45b Millions of years ago Kenyanthropus platyops 2.5 Australopithecus 3.0 anamensis 3.5 4.0 4.5 5.0 5.5 6.0 6.5 7.0 © 2014 Pearson Education, Inc. Australopithecus afarensis Ardipithecus ramidus Orrorin tugenensis Sahelanthropus tchadensis The oldest fossil evidence of hominins dates back to 6.5 million years ago Early hominins show evidence of small brains and increasing bipedalism © 2014 Pearson Education, Inc. Figure 34.46 © 2014 Pearson Education, Inc. Misconception: Early hominins were chimpanzees Correction: Hominins and chimpanzees shared a common ancestor Misconception: Human evolution is like a ladder leading directly to Homo sapiens Correction: Hominin evolution included many branches or coexisting species, though only humans survive today © 2014 Pearson Education, Inc. Figure 34.47 Evidence that hominins walked upright 3.5 million years ago (a) The Laetoli footprints © 2014 Pearson Education, Inc. (b) An artist’s reconstruction of A. afarensis “Robust” australopiths had sturdy skulls and powerful jaws “Gracile” australopiths were more slender and had lighter jaws © 2014 Pearson Education, Inc. Bipedalism Hominins began to walk long distances on two legs about 1.9 million years ago Bipedal walking was energy efficient in the arid environments inhabited by hominins at the time © 2014 Pearson Education, Inc. Tool Use The oldest evidence of tool use, cut marks on animal bones, is 2.5 million years old Fossil evidence indicates tool use may have originated prior to the evolution of large brains © 2014 Pearson Education, Inc. Early Homo 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 Stone tools have been found with H. habilis, giving this species its name, which means “handy man” © 2014 Pearson Education, Inc. Homo ergaster was the first fully bipedal, largebrained hominid The species existed between 1.9 and 1.5 million years ago Homo ergaster shows a significant decrease in sexual dimorphism (a size difference between sexes) compared with its ancestors © 2014 Pearson Education, Inc. Homo ergaster fossils were previously assigned to Homo erectus; most paleoanthropologists now recognize these as separate species © 2014 Pearson Education, Inc. Homo erectus originated in Africa by 1.8 million years ago It was the first hominin to leave Africa © 2014 Pearson Education, Inc. Neanderthals Neanderthals, Homo neanderthalensis, lived in Europe and the Near East from 350,000 to 28,000 years ago They were thick-boned with a larger brain, they buried their dead, and they made hunting tools Recent genetic analysis indicates that gene flow occurred between Neanderthals and Homo sapiens © 2014 Pearson Education, Inc. Homo Sapiens Homo sapiens appeared in Africa by 195,000 years ago All living humans are descended from these African ancestors © 2014 Pearson Education, Inc. Figure 34.UN10 Homo sapiens skull fossil, A 160,000-year-old fossil of Homo sapiens. © 2014 Pearson Education, Inc. The oldest fossils of Homo sapiens outside Africa date back about 115,000 years and are from the Middle East Humans first arrived in the New World sometime before 15,000 years ago In 2004, 18,000-year-old fossils were found in Indonesia, and a new small hominin was named: Homo floresiensis © 2014 Pearson Education, Inc. Homo sapiens were the first group to show evidence of symbolic and sophisticated thought In 2002, a 77,000-year-old artistic carving was found in South Africa © 2014 Pearson Education, Inc. B. Vertebrates and Human Medicine (Nature of Science reading) 1. There are many pharmaceutical products that come from vertebrates. a. The venom of the Thailand cobra is the source of Immunokine, which has been used for ten years in multiple sclerosis patients. b. ABT-594 from the poison dart frog is about 50 times more powerful than morphine without the addictive properties. 2. Animal pharming uses genetically altered vertebrates (mice, sheep, goats, cows, chickens, pigs) to produce medically useful pharmaceutical products. a. The human gene for some useful product is inserted into the embryo of the vertebrate. b. That embryo is implanted into a foster mother, which gives birth to the transgenic animal. 3. Xenotransplantation is the transplantation of nonhuman vertebrate tissues and organs into humans. a. The use of transgenic vertebrates for medical purposes raises many health and ethical concerns. © 2014 Pearson Education, Inc. Figure 34.50 © 2014 Pearson Education, Inc. Figure 34.UN11 Urochordata (tunicates) Marine suspension feeders; larvae display the derived traits of chordates Myxini (hagfishes) Jawless marine vertebrates with reduced vertebrae; have head that includes a skull and brain, eyes, and other sensory organs Jawless aquatic vertebrates with reduced vertebrae; typically feed by attaching to a live fish and ingesting its blood Aquatic gnathostomes; have cartilaginous skeleton, a derived trait formed by the reduction of an ancestral mineralized skeleton Aquatic gnathostomes; have bony skeleton and maneuverable fins supported by rays Chondrichthyes (sharks, rays, skates, ratfishes) Actinopterygii (ray-finned fishes) Amniotes: amniotic egg, rib cage ventilation Dipnoi (lungfishes) Tetrapods: four limbs, neck, fused pelvic girdle Lobe-fins: muscular fins or limbs Osteichthyans: bony skeleton Actinistia (coelacanths) © 2014 Pearson Education, Inc. Description Basal chordates; marine suspension feeders that exhibit four key derived characters of chordates Petromyzontida (lampreys) Gnathostomes: hinged jaws, four sets of Hox genes Vertebrates: Hox genes duplication, backbone of vertebrae Chordates: notochord; dorsal, hollow nerve cord; pharyngeal slits; post-anal tail Clade Cephalochordata (lancelets) Ancient lineage of aquatic lobe-fins still surviving in Indian Ocean Freshwater lobe-fins with both lungs and gills; sister group of tetrapods Amphibia (salamanders, frogs, caecilians) Reptilia (tuataras, lizards and snakes, turtles, crocodilians, birds) Have four limbs descended from modified fins; most have moist skin that functions in gas exchange; many live both in water (as larvae) and on land (as adults) One of two groups of living amniotes; have amniotic eggs and rib cage ventilation, key adaptations for life on land Mammalia (monotremes, marsupials, eutherians) Evolved from synapsid ancestors; include egg-laying monotremes (echidnas, platypus); pouched marsupials (such as kangaroos, opossums); and eutherians (placental mammals, such as rodents, primates) Figure 34.UN11a Clade Description Cephalochordata (lancelets) Basal chordates; marine suspension feeders that exhibit four key derived characters of chordates Urochordata (tunicates) Marine suspension feeders; larvae display the derived traits of chordates Myxini (hagfishes) Chondrichthyes (sharks, rays, skates, ratfishes) Actinopterygii (ray-finned fishes) Jawless marine vertebrates with reduced vertebrae; have head that includes a skull and brain, eyes, and other sensory organs Jawless aquatic vertebrates with reduced vertebrae; typically feed by attaching to a live fish and ingesting its blood Aquatic gnathostomes; have cartilaginous skeleton, a derived trait formed by the reduction of an ancestral mineralized skeleton Aquatic gnathostomes; have bony skeleton and maneuverable fins supported by rays Actinistia (coelacanths) Ancient lineage of aquatic lobe-fins still surviving in Indian Ocean Dipnoi (lungfishes) Freshwater lobe-fins with both lungs and gills; sister group of tetrapods Petromyzontida (lampreys) © 2014 Pearson Education, Inc.