Campbell Biology · Chapter 34
The Origin and Evolution of Vertebrates
pp. 697–735 · 8 sections
This chapter traces the animals with backbones, and their invertebrate chordate cousins, from small Cambrian swimmers to people. Each step adds a new feature: a notochord and nerve cord, a head, a backbone, jaws, limbs, a land-ready egg, hair and milk, and finally the upright walking and big brains of humans. Vertebrate groups aren't on the current AP exam, but the chapter is full of examples the course does test: reading family trees (Topic 7.9), fossils and homology as evidence (Topic 7.6) and endotherms versus ectotherms (Topic 8.2).
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34.1 The four chordate hallmarks and our invertebrate cousins
pp. 697–701
Chordate groups aren't in the current course, but the reasoning is: reading the chordate family tree (Topic 7.9), treating shared embryo features as evidence of common ancestry (Topic 7.6) and seeing how regulatory genes switched on in one region shape that region (Topic 6.6).
In the course: Topic 7.9 Phylogeny, Topic 7.6 Evidence of Evolution, Topic 6.6 Gene Expression and Cell Specialization (notes, videos and more questions)
Key points
- Vertebrates sit inside a bigger phylum, the chordates. Chordates are bilaterally symmetric deuterostomes, and their nearest relatives outside the phylum are echinoderms, such as sea urchins, and acorn worms.
- Every chordate shows four features at some stage, even if only as an embryo: a notochord, a hollow nerve cord running along the back, slits or clefts in the throat region (pharynx), and a muscular tail that continues past the anus.
- The notochord is a bendable supporting rod that lies between the gut and the nerve cord. In most vertebrates a jointed backbone takes over its job, and in you its leftovers help form the soft centers of the discs between your vertebrae.
- The chordate nerve cord starts as a flat strip of ectoderm on the embryo's back that curls up into a tube, which becomes the brain and spinal cord. Most other animals have a solid nerve cord on the belly side instead.
- Throat slits first served to strain food from water. In fishes they became gill slits, and in tetrapods the embryo's clefts never open but are reshaped into parts of the ear, head and neck.
- Two chordate groups have no backbone. Lancelets keep all four hallmarks as adults and branched off first. Tunicates show the hallmarks only as swimming larvae and lose most of them when they settle down as filter-feeding adults, yet DNA places tunicates closer to vertebrates than lancelets are.
- Lancelet embryos switch on the same region-labeling regulatory genes, in the same front-to-back order, as vertebrate embryos building a brain. That suggests the vertebrate brain is an expanded version of a small swelling at the front of an ancestral nerve cord.
Key terms (12)
- chordate
- A member of the phylum Chordata, the group that includes vertebrates, lancelets and tunicates. All of them have the four chordate hallmarks at some point in life.
- notochord
- A flexible rod of fluid-filled cells wrapped in tougher tissue that runs along a chordate's back. It's the body's first support, and muscles pull against it to bend the body.
- dorsal, hollow nerve cord
- A tube of nervous tissue along the back of a chordate, formed when a strip of ectoderm rolls up. In vertebrates it develops into the brain and spinal cord.
- pharyngeal slits or clefts
- Grooves along the sides of the throat region in every chordate embryo. They open as slits for feeding or breathing in aquatic chordates and are remodeled into other structures in tetrapods.
- post-anal tail
- A tail with muscle that extends behind the anus. Many chordates swim with it, while in humans it shrinks away before birth, leaving the tailbone.
- deuterostome
- A member of the animal group that includes chordates and echinoderms. In their embryos, the first opening that forms becomes the anus, not the mouth.
- lancelet
- A small, blade-shaped marine chordate (a cephalochordate) that keeps all four chordate hallmarks as an adult. It burrows in sand and strains food from water.
- tunicate
- A marine chordate (a urochordate), also called a sea squirt. Its tadpole-like larva shows the chordate hallmarks, but the adult is a sessile filter feeder that has lost most of them.
- suspension feeding
- Eating by straining tiny food particles out of the water, as lancelets and adult tunicates do with mucus across their throat slits.
- somite
- One of the paired blocks of mesoderm along both sides of the notochord in a chordate embryo. Somites give rise to the repeated muscle segments, and in vertebrates to vertebrae too.
- Hox genes
- A family of regulatory genes that tell embryo cells which part of the head-to-tail axis they belong to. They're arranged in clusters and have been kept across animal evolution.
- metamorphosis
- A major change in body form between the larva and the adult. A tunicate larva, for example, absorbs its tail and notochord when it settles and turns into a filter-feeding adult.
Check yourself: 34.1 The four chordate hallmarks and our invertebrate cousins
4 questions on 34.1 The four chordate hallmarks and our invertebrate cousins. Pick an answer to see if you got it, and why.
Divers collect a tiny, transparent marine larva. Under a microscope it has a stiff rod running the length of its body, a tubular nerve cord above that rod, several openings in the wall of its throat, and a muscular tail that extends well behind the anus. Which conclusion is best supported?
Which comparison correctly contrasts the main nerve cord of a chordate with that of a typical insect?
About four weeks after conception, a human embryo has a series of grooves and pouches along the sides of its neck region, like those that form gill slits in fish embryos. In humans, these become parts of the middle ear, the tonsils and some glands. How do these grooves count as evidence for evolution?
DNA comparisons place tunicates closer to vertebrates than lancelets are. Adult lancelets keep all four chordate hallmarks, but adult tunicates keep only their throat slits. Which inference about the common ancestor of all chordates is most reasonable?
0 of 4 answered
34.2 Getting a head: craniates and hagfishes
pp. 701–703
Craniates and hagfishes aren't in the current course. A few ideas carry over: gene duplication as a source of new genes (Topic 6.7), fossils and embryos as evidence (Topic 7.6), tree reading (Topic 7.9) and osmosis in hagfishes, whose body fluids are as salty as seawater (Topic 2.7). The book treats hagfishes as craniates without a backbone, but most biologists now count them as vertebrates.
In the course: Topic 6.7 Mutations, Topic 7.6 Evidence of Evolution, Topic 7.9 Phylogeny, Topic 2.7 Tonicity and Osmoregulation, Topic 7.2 Natural Selection (notes, videos and more questions)
Key points
- After the basic chordate body came a head: a brain at the front end of the nerve cord, eyes and other sense organs, and a skull around them. Chordates with a head are called craniates, and the head let them sense, steer and feed in more complicated ways.
- Craniates have extra copies of important developmental genes, including more than one Hox cluster, where lancelets have a single cluster. Spare copies can pick up mutations and take on new roles, which may have made more complex bodies possible.
- The neural crest is a band of embryo cells along the edges of the closing neural tube. These cells travel all over the body and build many vertebrate features, including parts of the skull, teeth, much of the peripheral nervous system and the deeper skin layer of the face.
- Craniates are more active than lancelets. They have bigger muscles, a gut that moves food with muscle, a heart of two or more chambers, hemoglobin-rich red blood cells, and kidneys. Muscles around their gill slits pump water across the gills for gas exchange.
- Fossils from China, about 530 million years old, catch the change in progress: one early chordate had a brain, eyes and gills but no skull, while a slightly more advanced one had cartilage capsules around its eyes and ears.
- Hagfishes are jawless, eel-shaped ocean scavengers with a skull of cartilage and a notochord kept for life. Attacked, they can release liters of slime that clogs a predator's gills. Unlike other vertebrates, their body fluids are about as salty as seawater.
- Update: the book places hagfishes outside the vertebrates because they seem to lack vertebrae. Since then, DNA studies have grouped hagfishes with lampreys (together called cyclostomes), and hagfish embryos turned out to form tiny vertebra-like pieces. So most biologists now treat hagfishes as vertebrates whose backbone was reduced.
Key terms (9)
- craniate
- A chordate with a head: a brain at the front of the nerve cord, sense organs and a skull. The word comes from cranium, the skull.
- cranium
- The skull, especially the part that encloses the brain. In hagfishes and sharks it's made of cartilage, and in most other vertebrates it's bone.
- neural tube
- The tube that forms when the strip of ectoderm along an embryo's back folds up. It becomes the brain and spinal cord.
- neural crest
- Bands of cells that form where the neural tube is closing in a vertebrate embryo. They migrate widely and build parts of the skull, teeth, many nerves and pigment cells.
- gene duplication
- A mutation that leaves an extra copy of a gene, or even a whole genome. The spare copy can change and take on a new job while the original keeps the old one.
- Hox cluster
- A row of Hox genes lying side by side on one chromosome. Lancelets have one cluster, while jawed vertebrates have four.
- hagfish
- A jawless, eel-shaped marine scavenger with a cartilage skull, a lifelong notochord and slime glands along its sides.
- cyclostome
- The group made of hagfishes and lampreys, the two living lines of jawless vertebrates. DNA evidence shows they share a common ancestor that jawed vertebrates don't.
- gill slits
- Throat openings that water flows through past gills, where gas exchange happens. In craniates, muscles and nerves around them let the animal pump water.
Check yourself: 34.2 Getting a head: craniates and hagfishes
4 questions on 34.2 Getting a head: craniates and hagfishes. Pick an answer to see if you got it, and why.
Researchers mark neural crest cells in a chick embryo with a dye that is passed on to all their descendant cells. Several days later, in which tissues would they most expect to find the dye?
After a gene is accidentally duplicated, one copy keeps doing the gene's original job. Why can the second copy evolve a new function without harming the organism?
Among chordates, craniates were the first to have red blood cells packed with hemoglobin. In a typical mammal, 100 mL of plasma alone dissolves about 0.3 mL of O₂. The same volume of whole blood holds about 15 g of hemoglobin, and each gram binds about 1.34 mL of O₂ (approximate real values). About how many times more O₂ can 100 mL of whole blood carry than 100 mL of plasma alone?
Hagfishes were long described as having no vertebrae. Researchers later found that hagfish embryos form small cartilage pieces along the notochord, much like the earliest vertebral elements of lamprey embryos. DNA data also group hagfishes with lampreys. What do these findings most strongly suggest?
0 of 4 answered
34.3 Backbones, lampreys and the first bone
pp. 703–704
Lampreys, conodonts and armored jawless fishes aren't in the current course. The sea lamprey's spread into the Great Lakes is a good example of an invasive species (Topic 8.7), and dating rocks with fossils ties to Topic 7.6.
In the course: Topic 8.7 Disruptions in Ecosystems, Topic 7.6 Evidence of Evolution, Topic 7.9 Phylogeny (notes, videos and more questions)
Key points
- Vertebrates split from the other craniates during the Cambrian. Another round of gene duplication, including a family of regulatory genes called Dlx, came along with a bigger skull and a backbone.
- Vertebrae can be as simple as small cartilage pieces lying along the notochord. In most vertebrates, though, they surround the spinal cord and take over the notochord's job of holding the body up. Fins stiffened by fin rays gave early swimmers more push and control.
- Lampreys are jawless vertebrates with a cartilage skeleton. Their notochord stays their main support for life, with small cartilage pieces along it that partly surround the nerve cord.
- Most adult lampreys are parasites. They attach to a living fish with a round sucking mouth, scrape through its skin with a toothed tongue and feed on its blood and tissue. Their larvae spend years buried in stream mud, filter-feeding.
- The sea lamprey spread into the upper Great Lakes through a shipping canal in the 1900s. Lake trout there had no history with the parasite, and their populations crashed until lampricide treatments of lamprey spawning streams brought the parasite under control.
- Conodonts were small early vertebrates with large eyes and hooked, mineralized feeding parts in the mouth. They were so abundant for about 300 million years that geologists use their tiny tooth-like fossils to date rock layers.
- Later jawless fishes had paired fins and bony armor. Under a microscope, the armor is built from tiny tooth-like units, which suggests that mineral-hardened tissue appeared first in the mouth, spread to the outer armor, and only later reached the inner skeleton, beginning with the skull.
Key terms (9)
- vertebrate
- A chordate with a backbone made of vertebrae, or at least vertebra-like pieces along the notochord. Fishes, amphibians, reptiles, birds and mammals are vertebrates.
- vertebra
- One of the small skeletal units that make up the backbone. In most vertebrates, each one wraps around and protects the spinal cord.
- cartilage
- A firm but bendable tissue made by cells surrounded by a protein-rich matrix. The skeletons of lampreys and sharks are mostly cartilage.
- lamprey
- A jawless vertebrate with a round sucking mouth. Most adults feed on the blood and tissue of other fish, while the larvae filter-feed in streams.
- conodont
- An extinct, slender early vertebrate with big eyes and hardened, hooked mouthparts. Its tooth-like fossils are common enough to help date rocks.
- mineralization
- Hardening of a tissue with minerals such as calcium phosphate. Bone, teeth and fish armor are mineralized tissues.
- fin ray
- A thin, stiff rod that supports a fish's fin and helps it push against water and steer.
- invasive species
- A species that spreads into a new area, often with human help, and harms the native species or ecosystem there.
- index fossil
- A fossil from a species that was widespread but lived for a limited time, so finding it helps date the rock layer it's in.
Check yourself: 34.3 Backbones, lampreys and the first bone
4 questions on 34.3 Backbones, lampreys and the first bone. Pick an answer to see if you got it, and why.
Fishery biologists tracked sea lampreys and lake trout in a large lake they had invaded. Starting in 1960, a chemical that kills lamprey larvae was applied to the streams where lampreys spawn (invented data): Year | Sea lamprey abundance (index) | Lamprey wounds per 100 lake trout 1950 | 90 | 35 1955 | 100 | 40 1960 | 98 | 39 1965 | 30 | 12 1970 | 18 | 7 1975 | 15 | 6 Which conclusion is best supported by the data?
Sea lamprey larvae spend several years buried in the mud of small streams, filter-feeding, before they change into parasitic adults that roam a large lake. To protect the lake's fish, managers treat certain stream stretches with a chemical that kills the larvae instead of treating the lake. Which reason best supports this plan?
In an adult lamprey, the main body support is a notochord with small cartilage pieces along it. In an adult trout, a column of bony vertebrae surrounds the spinal cord, and only small remnants of the notochord remain. What does this comparison suggest about how the backbone evolved?
One hypothesis says hardened (mineralized) tissue first evolved in the mouthparts of early vertebrates and only later spread to body armor and the inner skeleton. Which new discovery would most weaken this hypothesis?
0 of 4 answered
34.4 Jaws, sharks and the bony fishes
pp. 704–709
Fish groups aren't in the current course. What carries over: why a true 'bony fish' clade has to include you (Topic 7.9), lungs and swim bladders as homologous organs (Topic 7.6), and overfishing and dams as human disruptions (Topic 8.7).
In the course: Topic 7.9 Phylogeny, Topic 7.6 Evidence of Evolution, Topic 8.7 Disruptions in Ecosystems, Topic 2.2 Cell Size, Topic 8.1 Responses to the Environment (notes, videos and more questions)
Key points
- Gnathostomes are the vertebrates with jaws: sharks and rays, the two big groups of bony fishes, and every tetrapod. Since early in vertebrate history they have far outnumbered the two jawless groups that survive.
- Jaws probably evolved from skeletal bars that once held open the front gill slits. Jawed vertebrates also share four Hox clusters (the result of whole-genome duplication in early vertebrates), a larger forebrain for smell and vision, and, in water-dwelling forms, a lateral line that senses vibrations and water movement.
- Jawed fishes show up about 450 million years ago. Armored placoderms and spiny acanthodians are extinct, but by about 420 million years ago the three lineages alive today had split apart: the cartilage fishes, the lobe-fins and the ray-finned fishes.
- Sharks, rays and ratfishes (chondrichthyans) have skeletons mostly of cartilage. Traces of bone in their scales and teeth suggest their ancestors had more bone and the lineage later lost most of it. Sharks have rows of replaceable teeth, electric-field sensors, internal fertilization and a corkscrew fold in the intestine that adds surface area. Many species are now threatened by overfishing.
- Osteichthyans have bony skeletons. Most fishes ventilate their gills under a bony flap (the operculum), and many control buoyancy with a gas-filled swim bladder. Lungs came first in early bony fishes; in some later lines, they were modified into swim bladders.
- Ray-finned fishes are the largest vertebrate group, with more than 30,000 species, about half of all vertebrates. Lobe-fins have fleshy fins with a chain of bones inside. Their three surviving lines are coelacanths, lungfishes (which gulp air into lungs) and the tetrapods.
- Because tetrapods descend from lobe-fins, a 'bony fish' group that leaves out tetrapods isn't a true clade. Biologists include tetrapods, you included, in the osteichthyans.
Key terms (14)
- gnathostome
- A vertebrate with jaws. The name means 'jaw mouth,' and the group includes sharks, bony fishes and all tetrapods.
- lateral line system
- A row of sensors along each side of a fish's body that detects vibrations and changes in water movement around it.
- placoderm
- An extinct jawed fish covered in bony plates. Some were giant predators of Devonian seas.
- chondrichthyan
- A member of the group with sharks, rays, skates and ratfishes. Their skeletons are made mostly of cartilage.
- spiral valve
- A corkscrew-shaped fold inside a shark's intestine that adds surface area and slows food down, so more nutrients are absorbed.
- oviparous
- Describes animals that lay eggs, so the young develop and hatch in the outside world.
- ovoviviparous
- Describes animals that keep fertilized eggs inside the mother, where the embryos live on yolk and hatch before being born.
- viviparous
- Describes animals whose embryos develop inside the mother and get nourishment from her, then are born live.
- cloaca
- A shared chamber at the end of the gut where digestive, urinary and reproductive tracts empty, with one opening to the outside. Sharks, amphibians, reptiles and birds have one.
- osteichthyan
- A member of the clade of vertebrates with a bony skeleton. It includes ray-finned fishes, lobe-fins and, since they evolved from lobe-fins, the tetrapods.
- operculum
- The bony flap that covers and protects the gills of most bony fishes. Its movements help pump water over the gills.
- swim bladder
- A gas-filled sac in many fishes. Moving gas into or out of it lets a fish rise, sink or hover without swimming.
- ray-finned fish
- A bony fish whose fins are thin webs of skin held up by bony rays. Tuna, trout, eels and sea horses are examples.
- lobe-fin
- A bony fish with fleshy, muscular fins built around a central chain of bones. Coelacanths and lungfishes are the living fish members.
Check yourself: 34.4 Jaws, sharks and the bony fishes
4 questions on 34.4 Jaws, sharks and the bony fishes. Pick an answer to see if you got it, and why.
Which observation best supports the hypothesis that vertebrate jaws evolved from the skeletal supports of the front gill slits?
An old museum label defines 'Pisces' (fishes) as all jawed vertebrates except tetrapods. A biologist objects that this group is not a clade. Which is the best reason for the objection?
In the bony-fish family tree, the earliest-branching living ray-finned fishes (such as bichirs) have lunglike organs they use to breathe air, and lungfishes, which are lobe-fins, have lungs too. Only later-branching ray-finned fishes have a swim bladder used for buoyancy. Which conclusion is most parsimonious?
Some open-ocean sharks die if they are held motionless, while bottom-resting nurse sharks can lie still on the seafloor for hours. Which explanation best accounts for the difference?
0 of 4 answered
34.5 Limbs, the move onto land and amphibians
pp. 709–713
Tetrapod and amphibian groups aren't in the current course, but Tiktaalik is a favorite example of a transitional fossil (Topic 7.6), the fish-to-tetrapod tree is good practice for Topic 7.9, and amphibian declines are an example of ecosystem disruption (Topic 8.7).
In the course: Topic 7.6 Evidence of Evolution, Topic 7.9 Phylogeny, Topic 8.7 Disruptions in Ecosystems, Topic 2.2 Cell Size (notes, videos and more questions)
Key points
- In the Devonian, the paired fins of one lobe-fin lineage became limbs with digits. These were the first tetrapods ('four feet'), the group that now includes amphibians, reptiles (birds too) and mammals. The oldest full tetrapod skeletons are about 365 to 370 million years old, and fossil footprints suggest limbed animals existed even earlier.
- Life on land reshaped more than the limbs. Tetrapods gained a neck so the head can move on its own, hip bones joined firmly to the backbone so the hind legs can push the body along, and, in most adults, no gills: the embryo's throat clefts become parts of the ear and certain glands instead.
- Limbs didn't appear out of nowhere. Lobe-fins living in shallow, oxygen-poor Devonian waters already had lungs and sturdy fins with bones inside, so the tetrapod body was a remodeled fish body.
- Tiktaalik, a roughly 375-million-year-old fossil from the Canadian Arctic, mixes fish and tetrapod traits: scales, fins and gills, but also lungs, strong ribs, a neck, and fin bones arranged like an upper arm, forearm and wrist. Update: footprints with toes about 395 million years old, found in Poland, suggest limbed tetrapods were already walking before Tiktaalik lived, so it is best seen as a late-surviving close relative of tetrapod ancestors, not a direct ancestor.
- Living amphibians come in three groups: salamanders, which keep their tails; frogs, which lose the tail as adults and hop with strong hind legs; and caecilians, legless burrowers whose ancestors had legs.
- Many frogs live a double life: a tadpole with gills, a tail and a plant-based diet transforms into an air-breathing, insect-eating adult. Most amphibians stay tied to moisture, because their eggs have no shell and dry out quickly, and many rely on thin, moist skin for much of their gas exchange.
- Amphibians are declining worldwide. Habitat loss, climate change, pollution and a deadly chytrid fungus have caused steep drops in hundreds of species and dozens of extinctions.
Key terms (12)
- tetrapod
- A vertebrate with four limbs that have digits, or one descended from such animals. Amphibians, reptiles, birds and mammals are tetrapods, even snakes and whales.
- digit
- A finger or toe. Limbs ending in digits are the key trait of tetrapods.
- pelvic girdle
- The hip bones that attach the hind limbs to the body. In tetrapods they're fused to the backbone, so pushes from the legs move the whole body.
- transitional fossil
- A fossil that shows a mix of traits from an older group and a newer one, documenting a step in an evolutionary change.
- amphibian
- A tetrapod in the group that includes frogs, salamanders and caecilians. Most need water or damp places to reproduce.
- salamander
- An amphibian that keeps its tail as an adult. Many walk by bending their body from side to side.
- frog
- A tailless amphibian with long hind legs for jumping. Toads are frogs with drier, bumpier skin.
- caecilian
- A legless, burrowing amphibian of tropical regions. It evolved from ancestors that had legs.
- tadpole
- The aquatic larva of a frog, with gills and a finned tail. It usually feeds on algae and plants.
- paedomorphosis
- Keeping juvenile features into adulthood, like an axolotl salamander that breeds while still having gills.
- cutaneous respiration
- Gas exchange through the skin. Many amphibians get much of their oxygen this way, and some salamanders have no lungs at all.
- chytrid fungus
- A fungus that infects amphibian skin and disrupts its water and salt balance. It has driven many frog species into decline or extinction.
Check yourself: 34.5 Limbs, the move onto land and amphibians
4 questions on 34.5 Limbs, the move onto land and amphibians. Pick an answer to see if you got it, and why.
The hypothesis that tetrapods evolved from lobe-finned fishes makes predictions about the fossil record. Which discovery would most strongly contradict it?
The fin of a Devonian lobe-fin contains one bone nearest the body, then two side-by-side bones, then a cluster of small bones. A bird's wing has the same one-two-many layout. What best explains the match?
In a fish, the pelvic fins aren't connected to the backbone. In a tetrapod, the hip bones are fused to it. Why would this difference matter for an animal walking on land?
Lungless salamanders take in all their oxygen through their skin, and nearly all of them are small. Suppose one salamander were twice as long, twice as wide and twice as tall as another of the same shape. Its skin area would be about 4 times larger and its volume about 8 times larger. How would its skin area per unit of body volume compare with the smaller salamander's?
0 of 4 answered
34.6 Amniotes: eggs for dry land, reptiles and birds
pp. 713–720
Reptile and bird groups aren't in the current course, but endotherms versus ectotherms is tested in Topic 8.2, and the idea that birds belong inside the reptile clade is tree reading from Topic 7.9.
In the course: Topic 8.2 Energy Flow Through Ecosystems, Topic 7.9 Phylogeny, Topic 7.6 Evidence of Evolution, Topic 7.2 Natural Selection (notes, videos and more questions)
Key points
- Amniotes are the reptiles (birds included) and the mammals. They're named for the amniotic egg, in which the embryo grows four membranes around itself: the amnion surrounds it with cushioning fluid, the yolk sac holds stored food, the allantois collects wastes, and the chorion, together with the allantois, exchanges gases.
- Carrying its own water, the embryo can develop on dry land. Most reptiles add a leathery or hard shell that slows water loss. Most mammals dropped the shell and keep the embryo, still wrapped in an amnion, inside the mother.
- Amniotes also fill their lungs by expanding the rib cage, which moves more air than the throat pumping amphibians use, and they have tougher, less leaky skin, so they lose less water.
- Reptiles have scales made of keratin. Because most lay shelled eggs, sperm must reach the egg inside the female before the shell forms. Lizards, snakes, turtles and crocodilians are ectotherms that get most of their body heat from their surroundings, often by basking, so they need only a small fraction of the food that a same-sized mammal eats. Birds are endotherms.
- Living reptiles include tuataras, squamates (lizards and snakes, which descend from legged lizards), turtles, crocodilians and birds. DNA studies now put turtles closest to the crocodilian-and-bird group (archosaurs), not off on an early side branch.
- Birds are living dinosaurs that descend from two-legged, meat-eating theropods. Many nonbird dinosaurs had feathers, so feathers came before flight and probably first served for warmth or display. Archaeopteryx, about 150 million years old, had feathered wings but kept teeth, clawed fingers and a long bony tail. All other dinosaurs died out 66 million years ago.
- Flight is costly. Birds save weight with air-filled bones, a toothless beak and, in most females, a single ovary. They power it with big breast muscles anchored to a keeled breastbone, a four-chambered heart, air sacs that keep air moving through the lungs, and a high, steady body temperature.
Key terms (14)
- amniote
- A member of the clade of tetrapods with an amniotic egg: reptiles (including birds) and mammals.
- amniotic egg
- An egg with four membranes around the embryo that let it develop away from water. It's the key trait of amniotes.
- amnion
- The membrane that encloses the embryo in a pool of fluid, protecting it from drying out and from bumps.
- chorion
- The outermost membrane around the embryo. With the allantois, it handles the exchange of oxygen and carbon dioxide.
- yolk sac
- The membrane around the yolk, the embryo's food store. Its blood vessels carry nutrients into the embryo.
- allantois
- A membrane sac that stores the embryo's nitrogen wastes and also helps with gas exchange.
- ectotherm
- An animal that gets most of its body heat from its surroundings, like a lizard basking on a rock. Ectotherms need much less food than endotherms of the same size.
- endotherm
- An animal that keeps its body warm with heat from its own metabolism, like birds and mammals. It costs a lot of food energy.
- squamate
- A member of the reptile group made of lizards and snakes.
- archosaur
- The reptile clade that includes crocodilians, extinct dinosaurs and pterosaurs, and birds.
- theropod
- A two-legged, mostly meat-eating dinosaur. Tyrannosaurus was one, and birds evolved from this group.
- feather
- A branched structure made of keratin that grows from bird skin. Feathers insulate, shape wings for flight and are used in display.
- keel
- A ridge down the middle of a bird's breastbone. The large flight muscles attach to it.
- air sacs
- Thin-walled sacs connected to a bird's lungs that keep air flowing through the lungs in one direction, so birds get oxygen very efficiently.
Check yourself: 34.6 Amniotes: eggs for dry land, reptiles and birds
4 questions on 34.6 Amniotes: eggs for dry land, reptiles and birds. Pick an answer to see if you got it, and why.
Eggs of a frog and a lizard, each starting at 2.00 g, were kept in dry air at 30 °C for 48 hours. The frog eggs, with only a jelly coat, fell to 0.60 g. The lizard eggs, which have a leathery shell, fell to 1.84 g. Which statement is best supported?
In a developing turtle egg, researchers block blood flow through the chorion and allantois but leave the other membranes working. Which process would most directly fail?
The table shows the body temperatures of two animals held at different air temperatures (invented data). Air temperature (°C) | Animal X body temp (°C) | Animal Y body temp (°C) 5 | 37 | 7 15 | 37 | 16 25 | 37 | 26 35 | 38 | 34 Which statement is best supported?
A 50 g lizard resting at 25 °C uses about 1.8 kJ of energy per day. A 50 g mouse at the same temperature uses about 60 kJ per day (invented but realistic values). The lizard's daily energy use is about what percent of the mouse's?
0 of 4 answered
34.7 Mammals, their origins and the primates
pp. 720–728
Mammal groups and primate traits aren't tested, but this section is full of examples you'll use: jaw bones that became ear bones (Topic 7.6), mammals' radiation after the dinosaurs and marsupials' radiation in Australia (Topic 7.10), reading primate trees (Topic 7.9) and mammals as endotherms (Topic 8.2).
In the course: Topic 7.6 Evidence of Evolution, Topic 7.9 Phylogeny, Topic 7.10 Speciation, Topic 8.2 Energy Flow Through Ecosystems, Topic 7.2 Natural Selection (notes, videos and more questions)
Key points
- Mammals are amniotes that feed their young milk from mammary glands and have hair. They're endotherms with a four-chambered heart and a diaphragm that helps fill the lungs. Most have big brains for their size, care for their young for a long time, and have teeth of different shapes for cutting, piercing and grinding.
- Mammals evolved from synapsids, amniotes with one opening in the skull behind each eye where jaw muscles attach. Over more than 100 million years, synapsid jaws were reshaped: a new jaw joint formed, and two bones from the old joint became the hammer (malleus) and anvil (incus) of the middle ear.
- True mammals appeared by the Jurassic and stayed mostly small while dinosaurs ruled. After the nonbird dinosaurs died out 66 million years ago, mammals underwent an adaptive radiation into large plant-eaters, predators, fliers and swimmers.
- Living mammals form three groups. Monotremes (the platypus and echidnas) lay eggs and have no nipples. Marsupials are born tiny and finish developing while nursing, usually in a pouch. Eutherians, or placental mammals, have a long pregnancy supported by a complex placenta.
- Marsupials came to dominate Australia, which has been isolated for tens of millions of years. Many resemble placental mammals that fill similar roles elsewhere, a result of convergent evolution. When a land bridge formed between North and South America about 3 million years ago, animals crossed in both directions.
- Primates (lemurs and lorises, tarsiers, monkeys and apes) evolved in trees. They have grasping hands and feet with flat nails, big brains and short faces, forward-facing eyes for judging depth, and long care of their young. Monkeys and apes can press the thumb fully against each fingertip.
- Tarsiers are closer to monkeys and apes (the anthropoids) than to lemurs. New World monkeys, some with grasping tails, and Old World monkeys don't make up one clade. Apes split from Old World monkeys roughly 25 million years ago and have no tail.
Key terms (13)
- mammary gland
- A gland that makes milk to feed young. Every mammal has them, and they give the group its name.
- synapsid
- A member of the amniote line that led to mammals, marked by one opening in the skull behind each eye.
- diaphragm
- A sheet of muscle under the lungs. When it contracts, the chest expands and air rushes in.
- monotreme
- An egg-laying mammal. The platypus and echidnas are the only living ones, and their young lap milk from the mother's skin.
- marsupial
- A mammal born at a very early stage that completes its development while nursing, often in a pouch. Kangaroos and opossums are marsupials.
- eutherian
- A placental mammal, whose young develop for a long time inside the mother, fed through a complex placenta.
- placenta
- An organ formed from the mother's uterine lining and the embryo's membranes, where nutrients and oxygen pass to the embryo and wastes pass back.
- convergent evolution
- When unrelated lineages evolve similar features because they face similar challenges, like a marsupial and a placental mammal that both dig tunnels.
- adaptive radiation
- The rapid branching of one lineage into many species that fill different roles, often after a mass extinction or after reaching a new place.
- primate
- A mammal in the order with lemurs, tarsiers, monkeys and apes. Most have grasping hands, forward-facing eyes and large brains.
- opposable thumb
- A thumb that can touch the tips of the other fingers on the same hand, giving a strong, precise grip.
- anthropoid
- A member of the primate group that includes monkeys and apes, humans among them.
- prehensile tail
- A tail that can grip branches like an extra hand, found in some New World monkeys.
Check yourself: 34.7 Mammals, their origins and the primates
4 questions on 34.7 Mammals, their origins and the primates. Pick an answer to see if you got it, and why.
Biologists claim that the malleus and incus, two tiny bones in the mammalian middle ear, are homologous to bones that form the jaw joint in reptiles. Which observation would best support this claim?
Echidnas lay eggs, as reptiles and birds do. Marsupials and placental mammals give birth to live young. Phylogenies place monotremes on the earliest branch of the mammal tree. Which interpretation fits best?
Early marsupials reached Australia while it was still joined to Antarctica, and Australia then drifted off on its own. Today Australian marsupials include grazers, burrowers, tree-dwellers, gliders and predators. Which process best describes this pattern?
Echidnas, egg-laying mammals of Australia and New Guinea, and hedgehogs, placental mammals of Europe, Asia and Africa, both have backs covered in sharp spines made of modified hair, eat insects and other small invertebrates, and can curl into a defensive ball. DNA and many other traits show that echidnas are far more closely related to the platypus than to hedgehogs. How should a biologist building a mammal phylogeny treat the spiny coat?
0 of 4 answered
34.8 Human evolution: upright walking, big brains and a bushy tree
pp. 728–733
Hominin species aren't in the current course, but human evolution often shows up as tree reading (Topic 7.9): know why humans didn't evolve from chimpanzees. Fossils and DNA as evidence fit Topic 7.6, Neanderthal interbreeding tests the species concept (Topic 7.10), and founder effects fit Topic 7.4. Several of the book's dates are out of date; this page uses current ones.
In the course: Topic 7.9 Phylogeny, Topic 7.6 Evidence of Evolution, Topic 7.10 Speciation, Topic 7.4 Population Genetics (notes, videos and more questions)
Key points
- Compared with other apes, humans walk upright on two legs, have much larger brains, use language and symbols, make complex tools, and have smaller jaws and teeth. Human and chimpanzee DNA match at about 99% of comparable sites, but small differences, especially in genes that control other genes, can have big effects.
- Hominins are humans plus all the extinct species on our side of the split from chimpanzees. The oldest candidates, from Africa, are 6 to 7 million years old. Chimpanzees aren't our ancestors: they're our closest living relatives and have been evolving on their own branch just as long.
- Human traits didn't all evolve together. Upright walking came early, shown by a skull opening for the spinal cord placed underneath and by footprints left about 3.6 million years ago, while brains stayed ape-sized for millions of years. Biologists call this mosaic evolution.
- From about 4 to 2 million years ago, Africa held many australopiths, such as the small, upright Australopithecus afarensis and the heavy-jawed 'robust' species. Often several hominin species lived at once, so human evolution is a branching bush, not a ladder.
- Update: stone tools from Kenya are about 3.3 million years old, older than the earliest Homo fossils (about 2.8 million years old). Homo erectus, which includes the African fossils some call Homo ergaster, had a tall, long-legged body, and it spread beyond Africa about 2 million years ago, earlier than any other hominin we know of.
- Update: the oldest Homo sapiens fossils, from Morocco, are about 300,000 years old, and DNA from living people points to an African origin. Modern humans spread across the world, reaching the Americas at least 15,000 and probably more than 20,000 years ago.
- Update: sequenced Neanderthal genomes show they interbred with modern humans; most people with ancestry from outside Africa carry roughly 1 to 2% Neanderthal DNA. Neanderthals died out about 40,000 years ago. Other recent relatives include the Denisovans and the small Homo floresiensis of Indonesia, whose bones are now dated to about 60,000 to 100,000 years ago, far older than the book's estimate.
Key terms (11)
- hominin
- A human or any extinct species that shares a more recent ancestor with us than with chimpanzees.
- bipedalism
- Walking upright on two legs. In hominins it evolved millions of years before large brains did.
- foramen magnum
- The hole in the underside of the skull that the spinal cord passes through. Placed underneath the skull, it signals an upright posture.
- paleoanthropology
- The study of human origins through fossils, tools and other remains.
- australopith
- One of a varied group of small-brained, upright-walking hominins that lived in Africa about 4 to 2 million years ago.
- mosaic evolution
- When different traits of a lineage evolve at different times and rates, like upright walking long before big brains in hominins.
- Homo erectus
- A long-legged, larger-brained hominin species that appeared about 2 million years ago and was the first to spread from Africa into Asia.
- Neanderthal
- Homo neanderthalensis, a stocky, large-brained hominin of Europe and western Asia that made tools, buried its dead and interbred with modern humans.
- Denisovan
- An extinct group of hominins known mostly from DNA in a few bones and teeth from Asia. Some living people, especially in Oceania, carry Denisovan DNA.
- Homo sapiens
- Our own species, which arose in Africa about 300,000 years ago and has since spread worldwide.
- founder effect
- Loss of genetic variation when a small group splits off to start a new population, carrying only part of the original population's alleles.
Check yourself: 34.8 Human evolution: upright walking, big brains and a bushy tree
4 questions on 34.8 Human evolution: upright walking, big brains and a bushy tree. Pick an answer to see if you got it, and why.
A classmate says, 'If humans evolved from apes, why are there still chimpanzees?' Which response is most accurate?
In fossil hominins, a foramen magnum (the opening where the spinal cord enters the skull) placed underneath the skull is used as a sign of upright walking. A later study found that mammals that hop on two legs, such as jerboas and kangaroo rats, also have this opening set farther forward and under the skull than their four-legged relatives. Why does this finding matter for reading hominin fossils?
Genome studies estimate how much DNA in present-day people comes from Neanderthals. Rounded averages (published estimates vary from study to study): Population | Neanderthal-derived DNA (%) East Asian | 2.2 European | 1.9 South Asian | 1.9 West African | 0.5 Neanderthals lived only in Europe and western Asia. Which explanation fits the pattern best?
DNA evidence shows that Neanderthals and modern humans produced offspring whose descendants are alive today. Under the biological species concept, what does this evidence suggest?
0 of 4 answered