LECTURE NOTES

CHAPTER 30 LECTURE NOTES

I. Vertebrates belong to the phylum Chordata

The phylum Chordata includes three subphyla: Urochordata, Cephalochordata and Vertebrata.

A. Chordate Characteristics

Chordates are deuterostomes with four unique characteristics which appear at some time

during the animal's life. These characteristics are: the notochord; a dorsal, hollow nerve

cord; pharyngeal slits; and a muscular postanal tail.

1. Notochord

Notochord = A longitudinal, flexible rod located between the gut and nerve cord.

Present in all chordate embryos-
Composed of large, fluid-filled cells encased in a stiff, fibrous tissue.
Extends through most of the length of the animal as a simple skeleton.

In some invertebrate chordates and primitive vertebrates it persists to support the adult.

In most vertebrates, a more complex, jointed skeleton develops and the notochord is retained in adults as the gelatinous material of the discs between the vertebrae.

2. Dorsal, Hollow Nerve Cord

Develops in the embryo from a plate of dorsal ectoderm that rolls into a tube located dorsal to the notochord.

Unique to chordates.

=> Other animal phyla have solid, usually ventral nerve cords.

The brain and spinal cord (central nervous system) develops from this nerve cord.

3. Pharyngeal Slits

Chordates have a complete digestive system (mouth and anus). The pharynx is the mregion just posterior to the mouth and it opens to the outside through several pairs of slits.

The presence of these pharyngeal slits permits water entering the mouth to exit without passing through the entire digestive system.
These pharyngeal gill slits function for suspension-feeding in invertebrate chordates.
They have become modified for gas exchange and other functions during the evolution of vertebrates.

4. Muscular Postanal Tail

A tail extending beyond the anus, it is found in most chordates and contains skeletal elements and muscles.

Provides much of the propulsive force in many aquatic species.

The digestive tract in most nonchordates extends nearly the whole length of the body.

II. Invertebrate chordates provide clues to the origin of vertebrates

A. Subphylum Urochordata

Species in the subphylum Urochordata are called tunicates.

Entire animal is cloaked in a tunic made of a celluloselike carbohydrate.
Most are sessile marine animals which adhere to rocks, docks and boats.
Some species are planktonic, while others are colonial.

The tunicates are filter feeders. (See Campbell, Figure 30.2b)

Seawater enters through an incurrent siphon, passes through the slits of the pharynx into a chambertcalled the atrium, and exits via an excurrent siphon, the atriopore.
Food filtered from the water by a mucous net of the pharynx is moved by cilia into the intestine.
The anus empties into the excurrent siphon.
When disturbed, tunicates eject a jet of water through the excurrent siphon, so they are commonly called sea squirts.

Adult tunicates bear little resemblance to other chordates.

They lack a notochord', a nerve cord and tail-
They possess only pharyngeal slits.

Larval tunicates are free swimmers and possess all four chordate characteristics-

Larva attach by the head on a surface and undergo metamorphosis to adult form.

B. Subphylum Cephalochordata

Animals in the subphylum Cephalochordata are known as lancelets due to their bladelike shape. (See Campbell, Figure 30.3) Chordate characteristics are prominent and persist in the adult. These include:

Notochord
Dorsal nerve cord
Numerous gill slits
Postanal tail

Cephalochordates are marine filter feeders.

They burrow tail first into the sand with only the anterior exposed.
Water is drawn into mouth by ciliary action and food is trapped on a mucous net secreted across the pharyngeal slits.
Water exits through the slits and trapped food passes down the digestive tube-

Cephalochordates are feeble swimmers with fishlike motions.

Frequently move to new locations
Muscle segments are serially arranged in chevronlike rows, and coordinated contraction flexes the notochord from side to side in a sinusoidal pattern.
Muscle segments develop from blocks of mesoderm called somites that are arranged along each side of the notochord in the embryo.
The serial musculature is evidence of chordate segmentation which developed independently of that in annelids and arthropods.

C. Relationship of Invertebrate Chordates to the Vertebrates

Vertebrates first appear in the fossil records in Cambrian rocks.

Fossilized invertebrates {about 550 million years old) resembling cephalochordates are found in Burgess Shale of British Columbia.
This is about 50 million years older than the oldest known vertebrates.

Most zoologists feel the vertebrate ancestors possessed all four chordate characteristics and were suspension-feeders.

They may have resembled lance lets but were less specialized.
Information provided by molecular systematics supports the idea that cephalochordates are the closest relatives of vertebrates.

Cephalochordates and vertebrates may have evolved from a sessile ancestor by

paedogenesis.

Paedogenesis = Precocious attainment of sexual maturity in a larva.

Cephalochordates more closely resemble urochordate larvae than adult urochordates.
Changes in the developmental control genes can alter the timing of developmental events (i.e. gonad maturation).
Zoologists postulate that some early urochordatelike larval forms became sexually mature and reproduced before undergoing metamorphosis.
If reproducing larvae were successful, natural selection may have reinforced the absence of metamorphosis and a vertebrae life cycle may have evolved.

The divergence of cephalochordates from vertebrates occurred about 500 million years ago.

The differences between the two groups can be viewed as vertebrate adaptations to larger size and a more mobile life style.

III. The evolution of vertebrate characteristics is associated with increased size and activity

Vertebrates have retained the chordate features while adding other specializations. These

synapomorphies distinguish the vertebrates from urochordates and cephalochordates.

The unique vertebrate structures probably evolved in association with increased size and more

active foraging for food. The unique vertebrate adaptations include:

Vertebrates show a much greater degree of cephalization than cephalochordates.

=> The brain and sense organs are located at the anterior end which is the portion of the body which is in contact with the most environmental stimuli.

A skeleton including a cranium and vertebral column is the main axis of the body, replacing the notochord as the basic skeleton.

=> The cranium protects the brain.

=> The vertebral column provides support and a strong, jointed anchor that provides leverage to the segmental swimming muscles.

=> The axial skeleton made larger size and stronger, faster movement possible.

=> Most vertebrates also have ribs (anchor muscles and protect internal organs) and an appendicular skeleton supporting two pairs of appendages.

The vertebrate skeleton may be composed of bone, cartilage, or a combination of the two.

=> A majority of the skeleton is a non-living matrix which is secreted and maintained by living skeletal cells.

=> The living vertebrate endoskeleton can grow with the animal unlike the exoskeleton of arthropods.

The neural crest, a group of embryonic cells found only in vertebrates, contributes to the formation of certain skeletal components and many other structures distinguishing vertebrates from other chordates.

=> The dorsal, hollow nerve cord develops from an infolding of the edges of an ectodermal plate on the surface of the embryo.

=> The neural crest forms near the dorsal margins of the tube resulting from this

infolding.

=> Cells from the neural crest then migrate to various specific areas of the embryo and help form a variety of structures including some of the bones and cartilage of the head.

Some anatomical adaptations also support the greater metabolic demands of increased activity .

The generation of A TP by cellular respiration, to replace the energy used by vertebrates in obtaining food or escaping predators, consumes oxygen.

=> The respiratory and circulatory systems of vertebrates show adaptations which support the mitochondria of muscles and other active tissues.

Vertebrates have a closed circulatory system composed of a ventral chambered (2 to 4) heart, arteries, capillaries and veins.

=> The heart pumps the blood through the system.

=>The blood becomes oxygenated as it passes through the capillaries of the gills or lungs.

The more active the life style, the larger the amounts of organic molecules necessary to produce energy.

=> Vertebrates have several adaptations for feeding, digestion and nutrient absorption.

=> For example, muscles in the walls of the digestive tract move food from organ to organ along the tract.

IV. Vertebrate diversity and phylogeny: an overview

Seven extant classes comprise the subphylum Vertebrata: Agnatha, Chondrichthyes,

Osteichthyes, Amphibia, Reptilia, Aves, and Mammalia. The first three of these classes are fishes, while the remaining four are the tetrapod vertebrates.

Tetrapod = An animal possessing two pairs of limbs that support it on land.

In addition to being tetrapods, the reptiles, birds and mammals have other adaptations for a terrestrial lifestyle which are not found in the amphibians.

The amniotic egg (a shelled, water resistant egg) allows completion of the life cycle on land.
Most mammals do not lay eggs but retain other features of the amniotic condition, consequently, they are also considered amniotes along with the birds and reptiles.

It is recommended that figure 30.6 be reviewed to establish the relationships among the classes of the subphylum Vertebrata.

V. Agnathans are jawless vertebrates

The class Agnatha contains the jawless fishes and the extinct, heavily armored ostracoderms.

The oldest fossilized vertebrates are classified as agnathans. Some are found in Cambrian strata but most date to the Ordovician and Silurian (400 to 500 million years ago).

Early agnathans were small, less than 50 cm in length-
They were jawless with oval or slitlike mouths, most lacked paired fins and were bottom- dwellers.
Some were active and had paired fins-
Were probably bottom- or suspension-feeders that trapped organic debris in their gill slits.
Ostracoderms and most other agnathans declined and disappeared during the Devonian.

Extant forms include about 60 species of lampreys and hagfishes which lack paired appendages and external armor.

Lampreys are eel-shaped and feed by clamping their round mouths onto live fish.

Once attached, they use a rasping tongue to penetrate the skin and feed on the prey's blood.
Sea lampreys spend their larval development in freshwater streams and migrate to the sea or lakes as they mature.
Larva are suspension-feeders that resemble lancelets.
Some lamprey species feed only as larvae. Once they mature and reproduce, they die within a few days.

Hagfishes superficially resemble lampreys.

They are scavengers without rasping mouthparts.
Some species will feed on sick or dead fish while others feed on marine worms.
Lack a larval stage and are entirely marine.

VI. Placoderms were armored fishes with jaws and paired fins

The class Placodermi is a group of armored fishes which replaced agnathans during the late Silurian and early Devonian.

Most were less than 1 m in length, but some were up to 10 m long.
Differed from agnathans in that they possessed paired fins and hinged jaws.
Paired fins enhanced swimming ability and hinged jaws allowed more varied feeding

habits including predation.

Hinged jaws evolved as modifications of the skeletal rods which previously supported the anterior pharyngeal (gill) slits.

Remaining gill slits retained function as major gas exchange sites.
Hinged jaws of vertebrates work in an up and down direction those in arthropods work from side to side.

Placoderms and another group of jawed fishes, the acanthodians, radiated during the Devonian period (the Age of Fishes) and many new forms evolved in fresh and salt waters.

Placodenns and acanthodians disappeared by the start of the Carboniferous period (350 million years ago).
Ancestors of the placoderms and acanthodians also gave rise to early sharks ( class Chondrichthyes) and bony fishes ( class Osteichthyes ).

VII. Sharks and their relatives have adaptations for powerful swimming

The class Chondrichthyes contains about 750 extant species of cartilaginous fishes (e.g. sharks, skates, rays).

Species in the class Chondrichthyes have flexible skeletons composed of cartilage, well- developed jaws and paired fins.

The ancestors of members of this class had bony skeletons.
The characteristic cartilaginous skeleton is thus a derived characteristic, having evolved secondarily.
The developmental sequence in cartilaginous fishes differs from other vertebrates in that the initial (first) cartilaginous skeleton does not become ossified.
The cartilaginous skeleton is more elastic and lighter than a bone skeleton.

Sharks have streamlined bodies and are swift swimmers.

The tail provides propulsion.
The dorsal fins serve as stabilizers.
Pectoral and pelvic fins produce lift.
Some buoyancy is provided by large amounts of oil stored in liver, but most swim continuously to remain in the water column.

=> Continual swimming also produces water flow through mouth and over gills for gas exchange.

=> Some sharks are known to rest on the sea floor and in caves; water is continuously pumped over the gills while resting.

Most sharks are carnivorous, although the)argest (whale shark) is a suspension feeder.

Prey may be swallowed whole or pieces may be tom from large prey.
Teeth evolved as modified scales.
The digestive tract is proportionately shorter than in other vertebrates-

=> A spiral valve, which increases surface area and slows food movement, is present in the intestine.

Sharks possess sharp vision (can not distinguish color), and olfactory senses and other adaptations to their life style.

Electric sensory regions that detect muscle contractions of prey are located on the head.
A lateral line system is present along the flanks.

=> It is composed of rows of microscopic organs sensitive to water pressure changes and detects vibrations.

A pair of auditory organs also detect sound waves passing through the water.

Sharks reproduce sexually with internal fertilization.

A pair of claspers on the pelvic fins of males transfers sperm into the female reproductive tract.
Some species are oviparous, some are ovoviviparous, and a few are viviparous.
A cloaca (common chamber for reproductive, digestive and excretory systems) is present.

Rays are adapted to a bottom-dwelling life style.

They have dorsoventrally flattened bodies.
Their jaws are used to crush mollusks and crustaceans.
Enlarged pectoral fins provide propulsion for swimming.
The tail in many species is whiplike and, in some, bears venomous barbs.

VIII. Bony fishes are the most abundant and diverse vertebrates

The class Osteichthyes contains the bony fishes which are represented by more than 30,000 extant species.

Abundant in marine and fresh waters.
Range from 1 cm to 6 m in length.
Skeleton is bony, reinforced with a matrix of calcium phosphate.
Skin is covered with flattened bony scales.
Skin glands produce mucus that reduces drag when swimming.
A lateral line system is present as a row of tiny pits in the skin on both sides of the body.

Gas exchange occurs by drawing water over the four or five pairs of gills located in chambers covered by an operculum.

Water is drawn into the mouth, through the pharynx and out between the gills by movement of the operculum and contraction of muscles within the gill chambers.

=> Allows bony fishes to breath while stationary .

A swim bladder located dorsal to the digestive system provides buoyancy.

Transfer of gases between blood and swim bladder varies bladder inflation and adjusts the density of the fish.

Bony fishes are very maneuverable swimmers. The flexible fins provide better steering and propulsion than the stiff fins of sharks-

The fastest bony fish can swim to 80 km per hour in short bursts.
A fusiform body shape is common to all fast fishes and aquatic mammals.

=> This body shape reduces drag produced by the density ofwater (convergent evolution).

Most bony fish are oviparous and utilize external fertilization.

Some are ovoviviparous or viviparous and utilize internal fertilization.
Some display complex mating behavior.

The cartilaginous and bony fishes diversified during the Devonian and Carboniferous periods.

Sharks arose in the sea, bony fishes probably originated in fresh water.
The swim bladder was modified from lungs of ancestral fishes which supplemented the gills for gas exchange in stagnant waters.

The two extant subclasses of bony fishes had diverged by the end of the Devonian: the Actinopterygii (ray-finned fishes) and the Sarcopterygii (fleshy-finned fishes).

The subclass Actinopterygii includes fish with fins supported mainly by flexible rays.

These are the most familiar fishes-
They spread from fresh water to the seas and many returned to fresh water during evolution of the taxon.
Some bony fish (e.g. salmon and sea-run trout) reproduce in fresh water and mature in the sea.

The subclass Sarcopterygii includes three orders of fleshy-finned fishes that evolved in fresh water: lungfishes, coelocanths and rhipidistans.

The ancestors of these fishes continued to use their lungs to aid the gills in gas exchange.

Coelocanths and rhipidistians are referred to as lobe-finned fishes.

Their fins were fleshy, muscular and supported by extensions of the bony skeleton.
Many were large, bottom-dwelling forms that used their paired fins to walk on the substratum.
The only extant species, the coelocanth, is marine and lungless. It belongs to a lineage that became marine at some point in its evolution.
All rhipidistians are extinct.

Three genera of lungfishes exist in the Southern Hemisphere.

They live in stagnant ponds and swamps where they surface to gulp air into lungs connected to the pharynx.
When ponds dry, lungfishes burrow in the mud and aestivate.

Lobe-finned fishes of Devonian were numerous and important in vertebrate genealogy because they gave rise to amphibians. (See Campbell, Figure 30.13)

IX. Amphibians are the oldest class of tetrapods

The first vertebrates to move onto land were members of the class Amphibia.

Today there are about 4,000 extant species of frogs, salamanders and caecilians.

A. Early Amphibians

Early amphibians evolved from lobe- finned fishes that adapted to environmental variations (drought and flooding) of the Devonian.

The skeletal structure of the lobed fins suggest they could have assisted in movement on land.
Fossillobe-fins, such as the rhipidistian Eusthenopteron, exhibited many anatomical similarities to early amphibians. (See Campbell, Figure, 30.14a)

Oldest amphibian fossils are from the late Devonian (350 million years ago).

Early amphibians were predators that ate insects and other invertebrates that had moved previously onto land.
Amphibians were the only vertebrates on land in the late Devonian and early Carboniferous.

Radiation of forms occurred during the early Carboniferous period.

Some forms reached 4 m in length and some resembled reptiles.
Amphibians began to decline during the late Carboniferous. At the beginning of the Triassic period (230 million years ago), most of the survivors resembled modern species.

B. Modern Amphibians

There are three extant orders of amphibians: Urodela (salamanders), Anura (frogs and toads) and Apoda (caecilians).

The Order Urodela (salamanders) contains about 400 species.

Some are aquatic and some are terrestrial.
Terrestrial forms walk with a side-to-side bending of the body. Aquatic forms swim sinusoidally or walk along the bottom of streams or ponds.

The Order Anura (frogs and toads) contains about 3,500 species which are better adapted to the terrestrial habitat than urodeles.

Enlarged hindlegs provide better movement (hopping) than in urodeles.
They capture prey by flicking the sticky tongue which is attached anteriorly.
Predator avoidance aided by camouflage color patterns and distasteful or poisonous mucus secreted by skin glands.

=> Bright coloration is common in poisonous species.

The Order Apoda ( caecilians ) contains about 150 species.

They are legless and almost blind.
Most species burrow in moist tropical soils; a few species inhabit freshwater ponds and streams.

The Frog Life Cycle:

The frog exhibits a metamorphosis from the larval to adult stage. (See Campbell,

Figure 30.16)

The tadpole (larval stage) is usually aquatic, herbivorous and possesses internal gills, a lateral line system and a long, finned tail.
The tadpole lacks legs and swims by undulating the tail.
During metamorphosis, 1egs develop and the gills and lateral line system disappear.
A young frog is tetrapod, has air-breathing lungs, a pair of external eardrums and a digestive system that can digest animal protein.
The adult is usually terrestrial or semiaquatic and a predator.

Many amphibians, including some frogs, do not have a tadpole stage-

Some species in each order are strictly aquatic while others are strictly terrestrial.
Urodeles and apodans have larva that more closely resemble adults and both larva and adults are carnivorous.
Paedogenesis is common in some groups of urodeles.

Most amphibians maintain close ties with water and are most abundant in damp habitats-

Terrestrial forms in arid habitats spend much of their time in burrows where humidity is high.
Gas exchange is primarily cutaneous and terrestrial forms must keep the skin moist.

=> Lungs can aid in gas exchange although most are small and inefficient. Some forms lack lungs.

=> Many species also exchange gases across moist surfaces of the mouth.

Amphibians are dioecious.

The reproduce sexually usually with external fertilization in water ( e.g. ponds, streams, temporary pools).
In frogs, the male grasps the female and sperm are released as the female sheds her eggs. (See Campbell, Figure 30.16a)
Eggs are unshelled and produced in large numbers by most species.

Some species exhibit parental behavior and produce small numbers of eggs.

Males or females (species dependent) incubate eggs on their back, in the mouth or in the stomach.
Some tropical species lay eggs in a moist foamy nest that prevents drying-
Some species are ovoviviparous and a few are viviparous with the eggs developing in the female's reproductive tract.

Amphibians exhibit complex and diverse social behavior especially during breeding season (e.g. vocalization by male anurans, migrations, navigation or chemical signaling).

X. Evolution of the amniotic egg expanded the success of vertebrates on land

Many specialized adaptations for living in a terrestrial habitat were necessary for reptiles to evolve from their amphibian ancestor .

The amniotic egg was important as its development broke the last ties with the aquatic environment by allowing life cycles to be completed on land.

=> Amphibians must deposit their shell-less eggs in water.

=> The shell of the amniotic egg helps prevent desiccation.

The extraembryonic membranes within the egg develop from tissue layers that grow out from the embryo.

=> These specialized membranes function in gas exchange, transfer of stored nutrients to the embryo, and waste storage.

=> One of these membranes, the amnion, encloses a compartment filled with amniotic fluid that bathes the embryo and provides a cushion against shocks.

Birds and mammals evolved from reptiles.

Birds produce shelled amniotic eggs.
Placental mammals do not produce a shelled egg, the developing embryo is nourished by the mother after it becomes implanted in the uterine wall.

=> The extraembryonic membranes are retained and serve important functions in embryo development.

XI. A reptilian heritage is evident in all amniotes

The class Reptilia is a diverse group with about 7,000 extant species and a wide array of extinct forms.

Reptiles possess several adaptations to live on land not found in amphibians.

A. Reptilian Characteristics

Adaptations for a terrestrial existence include:

Scales contain the protein keratin which helps prevent dehydration.
Gas.exchange via lungs although many turtles also use moist cloacal surfaces-
Dioecious with sexual reproduction and internal fertilization.

=> Most are oviparous and produce an amniotic egg.

=> Some species of snakes and lizards are viviparous with the young obtaining nutrients from the mother across a "placenta" which forms from the extraembryonic membranes.

Reptiles are ectotherms and use behavioral adaptations to regulate their body temperature.

Ectotherm = An animal that uses behavioral adaptations to absorb solar energy and regulate its body temperature-

Due to ectothermy, reptiles can survive on a much lower caloric intake than mammals of comparable size.

Reptiles were abundant and diverse in the Mesozoic era.

B. The Age of Reptiles

I. Origin and Early Evolutionary Radiation of Reptiles

The oldest reptilian fossils are found in upper Carboniferous rock (300 million years old).

Ancestors were probably Devonian amphibians.
Two waves of adaptive radiation resulted in reptiles being the dominant terrestrial vertebrates for 200 million years.

The first reptilian radiation was in the early Permian period and gave rise to two main evolutionary branches: the synapsids and sauropsids.

Synapsids were terrestrial predators and gave rise to the therapsid lineage which were mammal-Iike reptiles.

=> Therapsids were large, dog-sized predators from which mammals are believed to have evolved.

Sauropsids gave rise to the modern amniote groups other than mammals. They split into two lineages early in their history: the anapsids and the diapsids.

=> The anapsids are presently represented only by the turtles.

=> The extant diapsids are represented by the lizards, snakes, and crocodilians; dinosaurs and some other extinct groups were also diapsids.

2. Dinosaurs and Their Relatives

The second reptilian radiation began in the late Triassic (about 220 million years ago ) and several lineages evolved during this event.

Two groups are most important: the dinosaurs and pterosaurs (flying reptiles).
Dinosaut:s varied in body shape, size and habitat.

=> Some fossilized forms measure 45 meters in length.

Pterosaurs had wings formed from skin stretched from the body wall, along the forelimb to the tip of an elongate finger and supported by stiff fibers.

Evidence indicates that dinosaurs were agile, fast moving and social, some may have even exhibited parental care of the young.

There is also some anatomical evidence supporting the hypothesis that dinosaurs were endothermic.

Endothermy = The ability to keep the body warm through an animal's own metabolism.

Skeptics of this hypothesis feel that the Mesozoic climate was warm and consistent, and that basking may have been sufficient for maintaining body temperature.
Low surface-to-volume ratios of large forms reduced fluctuations of body temperature vs air temperature; thus, dinosaurs may not have been endothermic.

3. The Cretaceous Crisis

In the Cretaceous (last period of the Mesozoic), the climate became cooler and more variable, and mass extinctions occurred.

Twenty-five percent of the families of marine invertebrates disappeared-
Nearly all dinosaurs disappeared by the end of the Mesozoic (65 million years ago) over a period of only five to ten million years-
The few reptilian groups to survive gave rise to extant forms.

C. Reptiles of Today

The largest and most diverse extant orders are the: Chelonia (turtles), Squamata (lizards and snakes), and Crocodilia (alligators and crocodiles).

Turtles of the order Chelonia evolved from anapsids during the Mesozoic.

They show little change from the earliest forms.
They are protected from predators by a hard shell-
All turtles, even aquatic species, lay their eggs on land.

Lizards and snakes are classified in the order Squamata.

Lizards are the most numerous and diverse group of extant reptiles.

They evolved from the diapsid lineage.
Most are small.
Many nest in crevices and decrease activity during cold periods.

Snakes probably descended from burrowing lizards.

They are limbless and most live above ground.
Vestigial pelvic and limb bones present in primitive snakes (boas) are evidence of a limbed ancestor.

Snakes are carnivorous and have a number of adaptations for hunting prey.

They have acute chemical sensors.
They are sensitive to ground vibrations (although lacking eardrums).
Pit vipers have sensitive heat-detecting organs between their eyes and nostrils.
Flicking tongue helps transmit odors toward olfactory organs on roof of mouth.
Poisonous snakes inject a toxin through a pair of sharp, hollow teeth and loosely articulated jaws allow them to swallow large prey.

Crocodiles and alligators of the order Crocodilia are among the largest living reptiles. (See Campbell, Figure 30.2ld)

Crocodilians also evolved from the diapsid lineage.
The spend most of their time in the water, breathing air through upturned nostrils.
They are confined to warm regions of Africa, China, Indonesia, India, Australia, South America and the southeastern United States.
They are the living reptiles most closely related to dinosaurs.

XII. Birds began as flying reptiles

The class Aves contains the birds which evolved from reptiles during the Mesozoic era.

They possess distinct reptilian characteristics such as the amniotic egg and scales on the legs.

A. Characteristics of Birds

Each part of the bird's anatomy is modified in some way that enhances flight-

The bones have a honeycombed internal structure that provides strength while reducing weight.
Some organ systems are reduced ( only one ovary in females).
Birds have no teeth (reduces weight) and food is ground in the gizzard.

=> The beak is made of keratin and evolution has produced many shapes in relation to the bird's diet.

Flying requires much energy production from an active metabolism.

Birds are endothermic with insulation provided by feathers and a fat layer .
Birds have efficient circulatory system with a four-chambered heart that segregates oxygenated blood from unoxygenated blood.
They have efficient lungs with tubes connecting to elastic air sacs that help dissipate heat and reduce the body density.

Birds also have a very well developed nervous system.

Acute vision and well-developed visual and coordinating areas of the brain aid in Flying.
They show complex behavior especially during breeding season when elaborate courtship rituals are performed.

Birds are dioecious with sexual reproduction and internal fertilization.

Sperm are transferred from the cloaca of the male to the cloaca of the female (males of most species lack a penis) during copulation.
Eggs are laid and must be kept warm through brooding by the female, male or both depending on the species.

Wings are airfoils, formed by the shape and arrangement of the feathers, that illustrate the same aerodynamic principles as airplane wings.

Power is supplied to the wings by contraction of the large pectoral (breast) muscles which are anchored to a keel on the sternum (breastbone).
Some birds have wings adapted for soaring (hawks) while others must beat their wings continuously to stay aloft (hummingbirds).

Feathers are made of keratin and are extremely light and strong. (See Campbell, Figure

30.23)

Feathers evolved from the scales of reptiles and may have first functioned as insulation.
Feathers also function to control air movements around the wing.

Radical alteration of body form was necessary for evolution of flight, but flight provides

many benefits.

Allows aerial reconnaissance that enhances hunting and scavenging.
Birds can exploit flying insects as an abundant, highly nutritious food resource.
Flight provides an escape mechanism from land-bound predators.
Flight also allows migration to utilize different food resources and seasonal breeding areas.

B. The Origin of Birds

Birds shared a common ancestor with Archaeopteryx lithographica.

Fossils of Archaeoptery.x have been recovered from limestone dated to the Jurassic period (150 million years ago).
Archaeopteryx had clawed forelimbs, teeth, a long tail containing vertebrae and feathers.
Archaeopteryx is not considered the ancestor to modern birds, but a side branch of the avian lineage.

=> The skeleton indicates a weak flyer which may have been a tree-dwelling glider.

C. Modern Birds

There are about 8,600 extant species in 28 orders. Most birds can fly but several are flightless (ostrich, kiwi, and emu).

Flightless birds are called ratites because the breastbone lacks a keel and large breast muscles used for flying are absent.
Flying birds are referred to as carinates due to the presence of a sternal keel (carina) that supports the large breast muscles used in flying.
Carinate birds exhibit a variety of feather colors, beak and foot shape, behavior and flight ability .

=> Penguins are also carinate birds which do not fly but use powerful breast muscles in swimming.

Almost 60% of extant species belong to one order of carinate birds (the Passeriformes, or perching birds) which includes the jays, swallows, sparrows, warblers and many others.

XIII. Mammals diversified extensively in the wake of the Cretaceous extinctions

There are about 4,500 species of extant mammals-

Extinction of the dinosaurs opened new adaptive zones at the end of the Mesozoic era.
Mammals underwent a massive radiation to fill these vacant zones.

A. Mammalian Characteristics

Species in the class Mammalia have the following characteristics:

Possess hair which is composed of keratin but is not believed to have evolved from reptilian scales.

=> The hair provides insulation.

Mammals are endothermic with an active metabolism.

=> An efficient respiratory system which utilizes a diaphragm for ventilation supports the metabolism.

=> A four-chambered heart segregates oxygenated from unoxygenated blood.

Mammary glands are present and produce milk to nourish the young.
Teeth are differentiated into various sizes and shapes which are adapted to chewing many types of food.

=> The jaw apparatus of the ancestral reptiles was also modified during evolution with two of the jaw bones becoming incorporated into the middle ear.

Mammals are dioecious with sexual reproduction and internal fertilization.

Most are viviparous with the developing embryo receiving nutrients from the female across the placenta.
A few are oviparous.

Mammals have large brains in comparison to other vertebrate groups and are capable of learning.

Parental care of long duration helps young learn from the parents.

B. Evolution of Mammals

Mammals evolve!i from therapsid ancestors (part of the synapsid branch) during the Triassic period.

The oldest fossil mammals are dated to 220 million years ago.
Early mammals coexisted with dinosaurs throughout the Mesozoic era.
Most Mesozoic mammals were small, probably insectivorous and nocturnal.

Mammals continued to diversify during the Cenozoic.

During the Cretaceous period (last of the Mesozoic), mass extinctions and mass radiations transformed the flora and fauna of Earth.

=> Angiosperms replaced gymnosperms as the dominant plants in many areas-

By the beginning of the Cenozoic, mammals were undergoing an adaptive radiation, and their diversity is represented today by three major groups: monotremes. marsupials and placental mammals.

C. Monotremes

The Monotremes include the platypuses and echidnas which are characterized by: Oviparity.

A reptilian-like egg with large amounts of yolk that nourishes the developing embryos.
Hair.
Milk production from specialized glands on the belly of females.

=> After hatching, young suck milk from the fur of the mother who lacks nipples.

The mixture of ancestral reptilian and derived mammalian traits suggests that monotremes descended from an early branch of the mammalian lineage.

Extant monotremes are found in Australia and New Guinea.

D. Marsupials

The Marsupials include opossums, kangaroos, koalas, and other mammals that complete their development in a marsupium (maternal pouch).

Marsupial eggs contain a moderate amount of yolk that nourishes the embryo during early development in the mother's reproductive tract-

Young are born in an early stage of development and are small (about the size of a honeybee in kangaroos).
The hind legs are simple buds, but the forelimbs are strong enough for the young to climb from the female reproductive tract exit to the marsupium.
In the marsupium, the young attaches to a teat and completes its development while nursing.

Convergent evolution in Australian marsupials has produced a diversity of forms which resemble placental counterparts in all ecological roles.

Opossums are the only extant marsupials outside of the Australian region.
South America had an extensive marsupial fauna during the Tertiary period as seen in the fossil record.

Plate tectonics and continental drift provide a mechanism which explains the distribution of fossil and modern marsupials.

Fossil evidence indicates marsupials probably originated in what is now North America and spread southward while the land masses were joined.
The breakup of Pangaea produced two island continents: South America and Australia.

=> With isolation, their marsupial faunas diversified away from the placental mammals that began adaptive radiation on the northern continents.

Australia has remained isolated from other continents for about 65 million years, thus isolating its developing fauna.
When North and South America joined at the isthmus of Panama, extensive migrations took place over the land bridge in both directions.

=> The most important migrations occurred about 12 million years ago and again about 3 million years ago.

E. Placental Mammals

In placental mammals, embryonic development is completed within the uterus where the embryo is joined to the mother by the placenta. (See Campbell, Figure 30.27c)

Adaptive radiation during the late Cretaceous and early Tertiary periods (about 70 -45 million years ago) produced the orders of extant placental mammals. (See Campbell, Table 30.2)

Fossil evidence indicates that placentals and marsupials diverged from a common ancestor about 80 -100 million years ago; thus, they are more closely related than either is to the monotremes.

Most mammalogists favor a genealogy that recognizes at least four main evolutionary lines of placental mammals.

One lineage consists of the orders Chiroptera (bats) and Insectivora (shrews) which resemble early mammals.

=> The modified forelimbs which serve a wings in bats probably evolved from

insectivores that fed on flying insects.

=> Some bats feed on fruits while others bite mammals and lap the blood.

=> Most bats are nocturnal.

A second lineage consists of medium-sized herbivores that underwent a massive adaptive radiation during the Tertiary period.

=> This led to such modern orders as the Lagomorpha (rabbits), Perissodactyla (odd-toed ungulates), Artiodactyla (even-toed ungulates), Sirenia (sea cows),

Proboscidea (elephants) and Cetacea (whales, porpoises).

The third evolutionary lineage produced the Order Carnivora which probably first appeared during the Cenozoic.

=> Included in this order are the cats, dogs, raccoons, skunks, and pinnipeds (seals,

sea lions, walruses).

=> Seals and their relatives evolved from middle Cenozoic carnivores that became adapted for swimming.

The fourth lineage had the greatest adaptive radiation and produced the primate- rodent complex.

=> Includes the orders Rodentia (rats, squirrels, beavers) and Primates (monkeys, apes, humans).

XIV .Primate evolution provides a context for understanding human origins

A. Evolutionary Trends in Primates

The first primates were small arboreal mammals.

Dental structure suggests they descended from insectivores in the late Cretaceous.
Purgatorius unio, found in Montana, is considered to be the oldest primate.

Primates have been present for 65 million years (end of Mesozoic era) and are defined by characteristics shaped by natural selection for living in trees. These characteristics include:

Limber shoulder joints which make it possible to brachiate (swing from one hold to the next).
Dexterous hands for hanging on branches and manipulating food.
Sensitive fingers with nails, not claws.
Eyes are close together on the front of the face, giving overlapping fields of vision for enhanced depth perception (necessary for brachiating).
Excellent eye-hand coordination.
Parental care with usually single births and long nurturing of offspring.

B. Modern Primates

Modem primates are divided into two suborders: Prosimii (premonkeys) and Anthropoidea (monkeys, apes, humans).

Prosimians (lemurs, lorises, pottos, tarsiers) probably resemble early arboreal primates.

There is a question as to which early prosimian lineage is ancestral to the anthropoids.

Two groups of prosimian fossils are recognized by paleontologists.

=> One ancestral to the tarsiers, the other to lemurs, lorises, and pottos.

The divergence of these two groups occurred at least 50 million years ago and it has been debated as to which of these two groups was also ancestral to the anthropoids.
Recently discovered fossils raise another possibility.

=> Fossils found in Asia and Africa which date at least 50 million years ago appear to be more similar to anthropoids than to either groups of prosimian fossils.

=> These fossils indicate an early divergence of prosimians into three lineages with the third being ancestral to the anthropoids.

Fossils of monkeylike primates indicate anthropoids were established in Africa and Asia by 40 million years ago in Africa or Asia (South America and Africa had already separated).

Ancestors of New World monkeys may have reached South America by rafting from Africa or migration southward from North America.

New World monkeys and Old World monkeys have evolved along separate pathways for many millions of years. (See Campbell, Figure 30.30)

All New World monkeys are arboreal-
Old World monkeys include arboreal and ground-dwelling forms.
Most monkeys, both New and Old World, are diurnal and usually live in social bands.

There are also four genera of apes included in the anthropoid suborder: Hylobates

(gibbons), Pongo (orangutans), Gorilla (gorillas) and Pan (chimpanzees). (See Campbell,

Figure 30.31)

Apes are confined to the tropical regions of the Old World.
They are larger than monkeys (except the gibbons) with relatively long legs, short arms and no tails-
Only gibbons and orangutans are primarily arboreal although all are capable of brachiation.
Social organization varies with the gorillas and chimpanzees being highly social.
Apes have larger brains than monkeys and thus exhibit more adaptable behavior.

XV. Humanity is one very young twig on, the vertebrate tree

Paleoanthropology concentrates on the small span of geological time during which humans and chimpanzees diverged from a common ancestor .

Paleoanthropology = The study of human origins and evolution.

Competition between researchers has often clouded the field of paleoanthropology .

Researchers gave new names to fossil forms which were actually the same species recovered by others. (This practice ended about 20 years ago.)
Theories were proposed on insufficient evidence; often a few teeth or a jawbone fragment.
Such actions resulted in many persistent misconceptions about human evolution even though fossil discoveries have disproved many of the myths.

A. Some Common Misconceptions

Our ancestors were chimpanzees or other modern apes.

Humans and chimpanzees represent two divergent branches of the anthropoid lineage which evolved from a common, less specialized ancestor .

Human evolution represents a ladder with a series of steps leading directly from an ancestral anthropoid to Homo sapien.

This progression is usually shown as a line of fossil hominids becoming progressively more modem.
Human evolution included many branches which led to dead ends with several different human species coexisting at times (See Campbell, Figure 30.32).
If punctuated equilibrium applies to humans, most evolutionary change occurred with the appearance of new hominid species, not phyletic (anagenic) change in an unbranched lineage.

Various human characteristics like upright posture and an enlarged brain evolved in unison.

Mosaic evolution occurred with different features evolving at different times.

=> Some ancestral forms walked upright but had small brains.

Present understanding of our ancestry remains unclear even after dismissing many of these myths.

B. Early Anthropoids

The oldest known fossils of apes are of Aegyptopithecus, the "dawn ape," which was a cat-sized tree-dweller about 35 million years ago.

About 25 million years ago (during the Miocene epoch) descendants of the first apes diversified and spread to Eurasia.

About 20 million years ago the Indian plate collided with Asia and the Himalayan range formed.

The climate became drier and the African and Asian forests contracted.

=> This isolated these regions of anthropoid evolution from each other .

Most anthropologists believe that humans and apes diverged from a common African anthropoid ancestor 6-8 million years ago.

Evidence from the fossil record and DNA comparisons between humans and chimpanzees supports this conclusion.

C. Australopithecines: The First Humans

Australopithecus africanus was discovered by Raymond Dart in 1924.

Additional fossils proved that Australopithecus was a hominid that walked fully erect and had humanlike teeth and hands-
The brain was about one-third the size of modem humans.
Various species of Australopithecus began appearing about 4.4 million years ago and existed for over 3.0 million years.

"Lucy", an Australopithecus skeleton, was discovered in 1974 in the Afar region of Ethiopia by paleoanthropoligists.

Lucy is 3.18 million years old.
The skeleton was 40% complete and small, about one meter tall with a head the size of a softball.

=> The structure indicates an upright posture.

It was different enough to be placed in a different species, Australopithecus afarensis.
Similar fossils have been discovered which indicate the species existed for about one million years.

The history of hominids is now believed to extend back to 4.4 million years.

Bone fragments from a hominid were recovered in 1994 in an area 75 km from the site of Lucy's discovery.

=> These fragments were so different from A. afarensis that a new hominid species was named, Australopithecus ramidus.

ramidus, which represents the oldest known hominid, exhibited some interesting characteristics.

=> Skull fragments indicated the head balanced on top of the spinal column - evidence of the early evolution of upright posture.

=> The skeletons of forest-dwelling animals were found among the bones –this challenges the view that bipedal ism evolved when humans began living on the savanna.

The discovery of A. ramidus and A. afarensis has raised several questions.

Is A. ramidus the ancestor of A. afarensis or an extinct evolutionary branch?
Is A. afarensis ancestral to other hominids or did it share a common ancestor with Homo?

=> A. afarensis underwent little change during its one million year span.

=> Several new hominid species resulted from an adaptive radiation which began about 3 million years ago.

The new species included A. africanus, several heavy-boned species of Australopithecus, and Homo habilis (appeared about 2.5 million years ago).

While the phylogeny of early hominids is uncertain, one fact is clear: hominids walked upright for two million years without a substantial increase in brain size.

This posture may have freed the hands for other things such as gathering food or caring for infants.

D. Homo habilis

Enlargement of the human brain is first evident in fossils dating to about 2.5 million years ago.

Skulls with brain capacities of about 650 cubic centimeters have been found compared with the 500 cc capacity of A. africanus.
Simple stone tools have been found at times with the larger-brained fossils.
Most paleoanthropologists believe these advances warrant placing the larger-brained fossils in the genus Homo and naming them Homo habilis.
It is clear from the fossil record that after walking upright for more than two million years, hominids began to use their brains and hands to fashion tools.

Homo habilis and other new hominids were a part of a larger speciation event among African mammals.

About 2.5 million years ago, Africa's climate began to become drier and savannas started to replace forests.
The fauna began to adapt to these new conditions.
Homo habilis coexisted with the smaller-brained Australopithecus for nearly one million years.
One hypothesis is that Australopithecus africanus (and other australopithecines) and Homo habilis were two distinct lines ofhominids.

=> Australopithecus africanus was an evolutionary dead end with Homo habilis leading first to Homo erectus which later gave rise to Homo sapiens.

E. Homo erectus and Descendants

Homo erectus was the first hominid to migrate out of Africa into Europe and Asia.

Fossils known as Java Man and Beijing Man are examples if H. erectus.

Homo erectus lived from about 1.8 million years ago unti1300,000 years ago.

Fossils found in Africa cover the entire span of H. erectus’ existence.
These populations existed during the same period as H. erectus populations on other continents.
The spread to new continents may have resulted from a gradual range expansion associated with a shift in diet to include a larger portion of meaL

=> In general, carnivores need a larger range than herbivores.

Homo erectus was taller and had a larger brain than H. habilis.

The El erectus brain capacity increased to as large as 1200 cc during the 1.5 million years of its existence.

=>This overlaps the normal range of modern humans.

The intelligence that evolved in H. habilis allowed early humans to survive in the colder climates to which they migrated.

Homo erectus lived in huts or caves, built fires, wore clothes of skins, and designed more refined stone tools than H. habilis.
Homo erectus was poorly equipped in a physical sense to live outside of the tropics but made up for the deficiencies with intelligence and social cooperation.

Some descendants of H. erectus developed larger brain capacities and exhibited regional diversity in populations.

The Neanderthals are the best known descendants of H. erectus.
Neanderthals lived in Europe, the Middle East, and Asia from 130,000 to 35,000 years ago.
They had heavier brows, less pronounced chins, and slightly larger brain capacities than modem man.
They were skilled tool makers who participated in burials and other rituals requiring abstract thought.

Many paleoanthropologists group the African post-Homo erectus fossils with Neanderthals and other descendants from Asia and Australasia. They believe these fossils represent the earliest forms of Homo sapiens.

Some post-H. erectus fossils date to 300,000 years ago.

F. The Emergence of Homo sapiens: Out of Africa...But When? Science as a Process

The debate over the origin of modem humans continues unabated with two widely divergent models currently being discussed. These are the Multiregional Model and the Monogenesis Model.

The Multiregional Model proposes: 1) Neanderthals and other post-Homo erectus hominids were ancestors to modem humans; and 2) modem humans evolved along the same lines in different parts of the world.

If this model is correct, the geographic diversity of humans originated between one and two million years ago when Homo erectus spread from Africa to other continents.
Supporters feel that interbreeding among neighboring populations provided opportunities for gene flow over the entire range and resulted in the genetic similarity of modem humans.

During the 1980's, some paleoanthropologists who interpreted the fossil record in a different way began to develop an alternative to the Multiregional Model. This alternative became known as the "Out of Africa" or Monogenesis Model.

The Monogenesis Model proposes: 1) Homo erectus was the ancestor to modern humans who evolved in Africa; and 2) modern humans dispersed from Africa, displacing the Neanderthals and other post-H erectus hominids.

If this model is correct, the diversity of modem humans has developed from geographic diversification within the last 100,000 years.
To supporters of this model, an exclusively African genesis for modem humans is strongly indicated by the fact that the complete transition from archaic Homo sapiens to modem humans is found only in African fossils.
The focus of their interpretations is on the relationship between Neanderthals and modem humans in Europe and the Middle East.
The oldest fossils of modem Homo sapiens are about 100,000 years old. These were found in Africa and similar fossils have been recovered from caves in Israel.
The fossils from Israel were found in caves near other caves containing Neanderthal- like fossils which date from 120,000 to 60,000 years ago -overlapping H sapiens by about 40,000 years.
Supporters of the Monogenesis Model interpret this information to mean that no interbreeding occurred during the time of coexistence since the two types of hominids persisted as distinct forms.
This interpretation means the Neanderthals were not ancestors of modem humans since they coexisted and were probably evolutionary dead ends along with other dispersed post-H erectus hominids.

Modem molecular techniques are being used to examine the question of human origins.

In the late 1980's, a group of geneticists compared the mitochondrial DNA (mtDNA) from a multiethnic group of more than lOO people from four different continents.
The premise for this analysis is that the greater the differences in the mtDNA of two people, the longer the time of divergence from a common source.
Analysis of the mtDNA comparisons resulted in the tracing of the source of all human mtDNA back to Africa with the divergence from that common source beginning about 200,000 years ago.
These results appeared to support the Monogenesis Model in that the divergence from the common source was too late to represent dispersal of Homo erectus, but supported a later dispersal of modem humans.

Several researchers have challenged the interpretation of the mtDNA study, especially the methods used to construct evolutionary trees from this type of data and the reliability of

mtDNA as a biological clock.

This criticism has encouraged advocates of the Multiregional Model to argue that the fossil evidence supports the multiregional evolution of humans more strongly than African monogenesis.
These scientists also consider certain fossils from different geographical regions to be links between that region's archaic Homo sapiens and the modern humans currently on that continent.

New evidence from comparisons of nuclear DNA and new methods for using the mtDNA data to trace the relationships of populations are stimulating further debate.

This new information strengthens the Monogenesis Model.
Greater genetic diversity is found in African populations south of the Sahara than in other parts of the world.

=> If modern humans evolved in southern Africa, these populations would have the longest history of genetic diversification.

=> If populations of humans in other parts of the world resulted from migrations out of Africa, the smaller genetic diversity could be the result of the founder effect and genetic drift.

Debate continues about whether the Multiregional Model or the Monogenesis Model is more accurate.

New evidence obtained by attempts to extract DNA from fossilized Neanderthal and other archaic Homo sapiens skulls may shed new light on this question.

G. Cultural Evolution: A New Force in the History of Life

Erect stance was a very radical anatomical change in our evolution and required major changes in the foot, pelvis and vertebral column.

Enlargement of the brain was a secondary alteration made possible by prolonging the growth period of the skull and its contents.

The brains of nonprimate mammalian fetuses grow rapidly, but growth slows and stops soon after birth.

The brains of primates continue to grow after birth and the period is longer for a human than other primates.
Parental care is lengthened due to this extended development and this contributes to the child's learning.

Learning from the experiences of earlier generations is the basis of culture (transmission of accumulated knowledge over generations); the transmission is by written and spoken language.

A cultural evolution is a continuum, but three stages are recognized:

Nomads of the African grasslands made tools, organized communal activities and divided labors about 2 million years ago.
The development of agriculture in Africa, Eurasia, and the Americas about 10,000 to 15,000 years ago encouraged permanent settlements.

The Industrial Revolution began in the eighteenth century.

=> Since then, new technology and the human population have escalated

exponentially.

No significant biological change in humans has occurred from the beginning to now.

Evolution of the human brain may have been anatomically simpler than acquiring an upright stance, but the consequences of cerebral growth have been enormous.

Cultural evolution resulted in Homo sapiens becoming a species that could change the environment to meet its needs and not have to adapt to an environment through natural selection.

=> Humans are the most numerous and widespread of large animals.

Cultural evolution outpaces biological evolution and we may be changing the world faster than many species can adapt.

=> The rate of extinctions this century is 50 times greater than the average for the past 100,000 years.

=> The overwhelming rate of extinction is due primarily to habitat destruction and chemical pollution, both functions of human cultural changes and overpopulation.

=> Global temperature increase and alteration of world climates are a result of escalating fossil fuel consumption.

=> Destruction of tropical rain forests, which playa role in maintenance of atmospheric gas balance and moderating global weather, is startling.

The effect of Homo sapiens is the latest and may be the most devastating crisis in the history of life.