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CHAPTER

19

CLASS AVES-BIRDS

ESSENTIAL QUESTIONS: 1. UNDERSTAND THE INTERNAL AND EXTERNAL ANATOMY OF BIRDS PHYSIOLOGICAL RELATIONSHIPS THAT PERTAIN TO THE ABILITY TO PRODUCE FLIGHT. 2. DISTINGUISH BETWEEN THE DIFFERENT PRIMARY ORDERS OF COMMON BIRDS. 3. DESCRIBE THE FLIGHT MECHANISM OF BIRDS INCLUDING DIRECT AND INDIRECT FLIGHT MUSCLE INVOLVEMENT AND DIFFERENT WING FORMS (ELLIPTICAL, HIGH SPEED, SOARING, AND HIGH LIFT). 4. CLARIFY REPRODUCTIVE STRATEGIES AND MATERNAL/PATERNAL CARE IN BIRD SPECIES. CHAPTER OUTLINE
19.1 Long Trip to a Summer Home A. Birds have conquered the air, escaping predators by reaching distant, difficult-to-reach locations. B. Permits them to utilize ever-changing environments. C. Summer locations have an abundance of insects, space for breeding, and favors homeostasis. D. Birds avoid climate extremes in migration. Diversity A. Profile 1. Over 9,900 species have been described worldwide; only fishes have more species among vertebrates. 2. Birds live in all biomes, from mountains to prairies, on all oceans, and from the North to the South Pole. 3. Some live in dark caves, and some dive to 45 meters depth. 4. The feather is the unique and essential feature or hallmark of birds (and its immediate ancestral group). 5. Uniformity in Structure a. Despite 150 million years of evolution, birds are still readily recognized. b. Forelimbs are modified as wings, although not all are capable of flight. c. Hind limbs are adapted for walking, swimming or perching. d. All birds have horny beaks. e. All birds lay eggs. f. The driving force for this uniformity appears to be the adaptations necessary for flight . 1) Wings are present for support and propulsion. 2) The respiratory system must meet high oxygen demands and cool the body. 3) Bones must provide a light but rigid airframe. 4) Digestion and circulation must meet the high-energy demands of flight. 5) And the nervous system must have superb sensory systems for high-velocity flight. g. Mammals, the endothermic evolutionary peers, have developed far more diverse forms. Origin and Relationships A. History 1. The discovery of the fossil of Archaeopteryx lithographica in 1861 linked birds and reptiles. a. The skull resembled modern birds but it had teeth rather than a beak. b. The skeleton was reptilian with clawed fingers, abdominal ribs and a long bony tail. c. Feathers were unmistakably imprinted along the wings. 2. Zoologists had long recognized that birds and reptiles shared many similarities. a. Both have skulls that abut the first neck vertebra by a single ball-and-socket joint. b. Both have a single middle ear bone, the stapes. c. The lower jaw in both is composed of five or six bones; in mammals there is one mandibular bone. d. Both birds and reptiles excrete nitrogenous wastes as uric acid; mammals excrete urea. e. Both lay similar yolked eggs; the embryo develops on the surface by shallow cleavage patterns. 3. Thomas Henry Huxley classified birds with theropod dinosaurs.
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19.4

a. This group of dinosaurs has a long, mobile, S-shaped neck. b. Therapods belong to the lineage of diapsid reptiles, the archosaurians, which includes crocodiles. c. Some dromeosaur-like fossils contained filaments or feathers. 1) These feathered dinosaurs could not fly. 2) Filaments and feathers may have been used for thermoregulation and colorful social displays. d. More derived fossils than Archaeopteryx from Spain and Argentina exhibited: 1) Keeled sternums and alulas. 2) Loss of teeth and fusion of bones. B. Relationships 1. Modern birds include Paleognathae with a flat sternum and Neognathae with a keeled sternum. 2. Original theories were based on the Paleognathae (or ratite) lineage never having attained flight. 3. This is now rejected; flightlessness has evolved many times among many bird groups. 4. Smaller birds can revert to flightlessness on islands that lack terrestrial predators. 5. Larger flightless birds such as the ostrich and emu can outrun predators. 6. Flightless birds are free from the weight restrictions of flight and some evolved to very large sizes. 7. Flightlessness has evolved many times in many different lineages of birds. Structural and Functional Adaptations for Flight 1. Flying requires more power and less weight for birds. 2. They must also find food, escape predators, repair themselves, thermoregulate, and reproduce. A. Feathers 1. Structure a. The feather is a special bird adaptation that contributes to more power or less weight. b. The hollow quill emerges from the skin follicle and continues as a shaft or rachis. c. The rachis bears numerous barbs. d. Up to several hundred barbs are arranged to form a flat, webbed surface, the vane. e. Each barb resembles a miniature feather; numerous parallel filaments or barbules spread laterally. f. With up to 600 barbules in each side of a barb, there may be over one million in the whole feather. g. Barbules from two neighboring barbs overlap; they zip together with tiny hooks. h. When separated, they are zipped back together by preening. 2. Types of Feathers a. Contour feathers provide the form of the bird; flight feathers extend off the wing in flight. b. Down feathers are under contour feathers; their barbules lack hooks and they function as insulation. c. Filoplume feathers are hair-like, degenerate feathers with a weak shaft and tuft of short barbs. d. Powder-down feathers on herons and their relatives disintegrate and release a talc-like powder to waterproof feathers. 3. Origin and Development a. The bird feather is homologous to the reptile scale. b. The feather develops from an epidermal elevation over a nourishing dermal core. c. Rather than flattening, the feather bud rolls into a cylinder. d. During growth, pigments (lipochromes and melanin) are added to the epidermal cells. e. Near the end of its growth, the soft rachis and barbs transform into hard structures of keratin. 4. Molting a. The fully-grown feather is a dead structure; shedding or molting is an orderly process. b. Molting is a gradual process that avoids leaving bare spots. c. Flight and tail feathers are lost in pairs, one on each side, to maintain balance. d. In some species, replacement is continuous; therefore flight is unimpaired. e. In many water birds, primary feathers are molted all at once and the birds are temporarily grounded. f. Most birds molt once a year, usually in late summer after the nesting season.
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5. Color a. Feather color may be due to pigments or to structural color. b. Pigments, or lipochromes, color red, orange and yellow feathers. c. Black, brown, red-brown, and gray colors are from the pigment melanin. d. The blue color of the blue jay, indigo bunting and bluebird is from scattering of light by structure. B. Skeleton 1. Bone Weight a. Compared with the Archeopteryx, modern birds have light, delicate bones laced with air cavities. b. These are termed pneumatized bones; they are nevertheless strong. c. The total weight of a birds feathers may outweigh its skeleton. 2. Bird Skull a. As archosaurs, birds evolved from ancestors with diapsid skulls. b. Bird skulls are so specialized that it is difficult to see the diapsid condition. c. The skull is fused into one piece; the braincase and orbits are large to hold a larger brain and eyes. 3. Jaws a. In Archeopteryx, both jaws contained teeth set in sockets. b. Modern birds have a horny keratinous beak molded around bony jaws. c. Most birds have kinetic skulls; in some, the upper jaw is hinged to the skull. 4. Vertebral Column and Appendages a. The bird vertebral column is very rigid; vertebrae are fused except for the cervical vertebrae. b. Additional bony structures called uncinate processes are fused with the pelvic girdle to support legs and provide rigidity for flight. c. Ribs are mostly fused with the vertebrae, pectoral girdle and sternum. d. Except in flightless birds, the sternum bears a large keel for anchorage of flight muscles . e. An elastic furcula stores energy as it flexes during wingbeats. f. Archaeopteryx with its small sternum was probably not a strong flier. g. Bones in the forelimbs are highly modified for flight, with some bones reduced in number or fused. h. All of the elements of the basic vertebrate limb are represented in modified form. C. Muscular System 1. The pectoralis muscles depress the wing in flight and are attached to the keel. 2. The supracoracoideus muscle raises the wing, is also attached to the keel, lies under the pectoralis muscles, and pulls the wing up from below by way of a rope-and-pulley type of arrangement. 3. Having both these muscles low in the body provides aerodynamic stability. 4. The main leg muscle mass is in the thigh with connections by long tendons to the feet and toes. a. This mass is also located near the birds center of gravity and permits great agility to the feet. 5. Feet are composed of bone, tendon, and tough scaly skin structures resistant to freezing. 6. A toe-locking mechanism prevents a perching bird from falling off a branch while asleep. D. Digestive System 1. Birds have rapid and efficient digestive systems. a. A shrike can digest a mouse in three hours. b. A thrush will pass berries through the tract in just 30 minutes. c. Insects are found in many bird diets. 2. Birds lack teeth, a gizzard grinds food. 3. Components: horn-covered tongue; short pharynx; elastic esophagus; stomach; and crop in many species 4. Salivary glands are poorly developed but do secrete mucus to lubricate food. 5. Many birds have a crop that stores food at the lower end of the esophagus. a. The crop of pigeons, doves and some parrots, also produces lipid- and protein-rich milk.
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E.

F.

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6. Stomach proper consists of a proventriculus, which secretes gastric juice and a muscular gizzard. a. Grain-eating birds swallow gritty objects or pebbles to grind up food in the gizzard. b. Owl pellets are composed of bones and fur formed in the proventriculus. c. At the end of the intestine with the rectum are paired ceca (well-developed in herbivorous birds), which serve as fermentation chambers. d. The cloaca is the terminal digestive organ that also receives genital ducts and ureters. 7. A great variety of beaks have evolved from the strong, pointed beaks of crows to the highly specialized beaks of flamingos and pelicans . a. A woodpeckers skull partially absorbs the shock of drilling into wood. b. The woodpecker beak is straight, hard, and chisel-like. c. The tail braces a woodpeckers body against a tree while it delivers rapid blows. d. The long, flexible, barbed tongue of woodpecker seeks insects in drilled holes and crevices. Circulatory System 1. The four-chambered heart is large, with strong ventricular walls. 2. Birds share with mammals a complete separation of respiratory and systemic circulations. 3. The heartbeat is relatively fast compared to mammals and is inversely proportional to size. a. A turkey heart beats 93 times per minute. b. A chicken heart beats 250 times per minute. c. A small black-capped chickadee heart beats 500 times per minute. 4. Bird red blood cells (erythrocytes) are nucleated and biconvex. 5. Mobile phagocytes are active and efficient in repairing wounds and destroying microbes. Respiratory System 1. The bird respiratory system differs radically from the lungs of both reptiles and mammals. 2. Bird Lungs a. The finest branches of the bronchi do not terminate in alveoli but are tube-like parabronchi. b. Nine interconnecting air sacs extend into the thorax, abdomen, and even the long bones. c. A large portion of the air bypasses the lungs and flows directly to the air sacs on inspiration. d. On expiration, this oxygenated air flows through the lungs; therefore there is continuous air flow. 3. The air sacs extend into bones, legs and wings, providing considerable buoyancy to the bird . Excretory System 1. A pair of large metanephric kidneys is composed of many thousands of nephrons. 2. Each nephron has a renal corpuscle and a nephric tubule. 3. Birds use the vertebrate pattern of glomerular filtration and selective resorption. 4. Urine flows through ureters to the cloaca. 5. Uric Acid a. Birds also use the reptilian adaptation of excreting nitrogenous wastes as uric acid. b. In shelled eggs, all excretory products remain within the eggshell; uric acid is stored harmlessly. c. Since uric acid has low solubility, a bird can use far less water to excrete wastes. d. Concentration of uric acid occurs almost entirely in the cloaca where water is absorbed. e. A bird kidney is less efficient than a mammal kidney in removing ions of sodium, etc. f. Marine birds must excrete larger salt loads due to the food they eat and seawater they drink; salt glands located above each eye excrete highly concentrated solutions . g. This salt solution runs out the nostrils; thus gulls and other sea birds have a perpetual runny nose. Nervous and Sensory Systems 1. A birds nervous and sensory system must accommodate the problems of flight and a visual lifestyle. 2. The birds brain has well-developed cerebral hemispheres, cerebellum and optic lobes. a) The cerebral cortex, a chief coordinating center in mammals, is thin, unfissured and poorly developed.
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b) The core of the cerebrum, the dorsal ventricular ridge, is enlarged into the principal integrating center. c) The size of the cerebral hemisphere is directly related to the intelligence of the bird. d) The cerebellum is where muscle-position sense (proprioception), equilibrium sense and visual cues are assembled. e) The optic lobes bulge to each side of the midbrain and form a visual association apparatus. 3. Sense of smell is poorly developed except in flightless birds, ducks and vultures . 4. Birds have good hearing and superb vision, the best in the animal kingdom. 5. The bird ear is similar to the ear of mammals. a. An inner ear has a short cochlea; it allows birds to hear about the same range of sound as humans. b. Bird ears do not hear as high a frequency as do humans but surpass us in ability to distinguish differences in pitch and intensities. 6. The bird eye is similar to the mammal eye but it is relatively larger for a given body size. a. A bird eye is less spherical and almost immobile; a bird turns its head rather than its eyes. b. The light-sensitive retina has both rods and cones. c. Diurnal birds have more cones; nocturnal birds have more rods. d. A pecten is a highly vascularized organ attached to the retina and it juts into the vitreous humor; it may provide oxygen and nutrients to the eye but its function is not certain . e. Many birds have two foveae or regions of detailed vision; this provides both sharp monocular and binocular vision. f. A hawk has eight times the visual acuity of a human and can see a rabbit over a kilometer away. g. An owls ability to see in dim light is more than ten times that of a human. h. Many birds can see partially into the ultraviolet spectrum, seeing flower nectar guides. I. Flight 1. History a. There are two hypotheses on the evolution of bird flight. 1) The ground-up hypothesis is based on running birds with primitive wings to snare insects. 2) The trees-down hypothesis has birds passing through tree-climbing, leaping, parachuting, gliding, and finally powered flight. b. Feathers preceded flight and arose for thermoregulatory purposes. 2. Bird Wing as a Lift Device a. Lift is provided by the more medial part of the wing and secondary feathers of the forearm . b. A wing is streamlined with a concave lower surface. c. The leading edge of the wing has small tight-fitting feathers. d. Over two-thirds of the total lift comes from negative pressure from the airstream flowing a longer distance over the top of the wing (the cambered surface), than over the convex surface. e. Lift-to-drag ratio is determined by the angle of tilt and the airspeed. f. At a point near 15o, the angle of attack becomes too steep and stalling occurs. g. Stalling is delayed or prevented by a wing slot along the leading edge to direct rapidly moving air across the leading surface. 1) In some birds the alula, or group of small feathers on the thumb, provides a midwing slot. 2) Slotting between the primary feathers provides a wing-tip slot. 3. Flapping Flight a. Flapping flight requires a vertical lifting force and a horizontal thrusting force. b. Thrust is provided by primaries at the wing tips and lift is provided by the secondaries. c. Greatest power is provided by the downstroke. d. Primary feathers are bent upward and twist to a steep angle of attack. e. On the upstroke, the primary feathers bend so that their upper surfaces twist to produce thrust. f. The powered upstroke is essential for hovering and fast, steep takeoffs.
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4. Basic Forms of Bird Wings a. Elliptical Wings 1) Found in sparrows, warblers, doves, and woodpeckers. 2) Birds that must maneuver in forested habitats have elliptical wings. 3) This wing type has a low aspect ratio (ratio of length to average width). 4) Elliptical wings are slotted between primary feathers to prevent stalling at low speeds, etc. 5) The small chickadee can change its course within 0.03 seconds. b. High-Speed Wings 1) Found in swallows, hummingbirds, swifts, plovers, sandpipers, gulls, and terns. 2) Birds that feed on the wing or make long migrations have high-speed wings. 3) These wings sweep back and taper to a slender tip; this reduces tip vortex turbulence. 4) They are flat in section and lack wing-tip slotting, and have a moderately high aspect ratio. c. Soaring Wings 1) Albatrosses and other oceanic soaring birds have wings with long, narrow wings. 2) The high-aspect ratio of long, narrow wings lacking wing slots permits high speed, high lift, and dynamic soaring. 3) They have the highest aerodynamic efficiency of any design, but are less maneuverable. d. High-Lift Wings 1) Vultures, hawks, eagles, owls and other birds of prey that carry heavy loads have wings with slotting, alulas, and pronounced camber. 2) This wing design produces high lift at slow speed. 3) Many are land soarers; their broad, slotted wings allow sensitive response for static soaring. 19.5 Migration and Navigation A. Migration 1. About half of all bird species migrate. 2. They generally move between southern wintering regions and northern summer breeding regions. 3. It also favors homeostasis, birds avoid climatic extremes and food shortages. B. Migration Routes 1. Most migratory birds follow established north-south routes. 2. Some use different routes in the fall and spring. C. Stimulus for Migration 1. The long days of late winter and early spring stimulate development of gonads and fat. 2. Long day length stimulates the anterior lobe of the pituitary. D. Direction Finding in Migration 1. Experiments suggest birds navigate chiefly by sight. 2. Birds recognize topographical landmarks and follow familiar migratory routes. 3. Birds also have a highly accurate innate sense of time and an innate sense of direction. 4. Research indicates they can navigate by the earths magnetic field; this may be related to magnetite (Fe3O4) found in the neck musculature of pigeons. 5. Sun-azimuth Orientation a. German ornithologists used special cages to show birds navigate by sun at day and stars at night. b. Planetarium experiments revealed the sun as a compass; an internal clock tracks position. c. The North Star may be used as an axis at night. 6. Migration involves a combination of environmental and innate cues. 19.6 Social Behavior and Reproduction A. Cooperative Behavior 1. Sea birds often gather in huge colonies to nest and to rear young.
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2. Land birds, except for birds such as starlings and rooks, tend to seek isolation for rearing their brood. 3. Birds that isolate during breeding may congregate for migration or feeding. 4. There are many advantages for flocking together: a. Mutual protection from enemies. b. Greater ease in finding mates. c. Less opportunity for an individual straying during migration. d. Mass huddling for protection against low night temperatures during migration. 5. Pelicans use organized cooperative behavior to feed. B. Reproductive System 1. Bird testes are very small until the approach of the breeding season, when they may enlarge 300 times. 2. Males of most species lack a penis; mating involves bringing cloacal surfaces in contact. 3. In most birds, the left ovary and oviduct develop and the right ovary and oviduct degenerate. 4. Special glands add albumin or egg white to the egg as it passes down the oviduct. 5. Sperm remain alive in the oviduct for many days after a single mating. C. Mating Systems 1. Over 90% of bird species are monogamous; they only mate with one partner each breeding season. a. Extrapair copulations may improve the genetic quality or fitness of the offspring. b. Males father offspring that are taken care of by other males. 2. In a few species, such as swans and geese, partners are chosen for life. 3. A smaller number are polygamous; individuals mate with two or more partners each breeding season. 4. Because of their high monogamy rate, compared to mammals, birds lack a built-in food supply and require the parental care from both parents to provision the young. 5. Bird Territories a. A male sings often to announce his presence to females and drive away males. 6. Polygyny a. The most common form of polygamy is polygyny where one male mates with many females. b. Competition for females is intense and females appear to choose the dominant male for mating. D. Nesting and Care of Young 1. Cuckoos and cowbirds are nest parasites; they lay eggs in other birds nests. 2. Precocial birds are covered with down and feed and run or swim as soon as they hatch. 3. Altricial birds are naked and helpless at birth and must be fed in the nest for a week or more. 4. Altricial birds have advantages too: a. The mother makes a smaller reproductive investment with smaller eggs and yolk supplies. b. Eggs lost to predation and weather can be easily replaced. c. Altricial young grow faster. Bird Populations A. Factors 1. Humans have introduced birds to new regions; the starling and the house sparrow are both abundant now in the United States. 2. Since the dodo went extinct in 1695, more than 80 bird species have also become extinct due to human influence. 3. Causes of bird extinction include habitat destruction, hunting, and competition with introduced species. 4. Recent Decline of Songbirds a. Some songbird species that were abundant 40 years ago are in decline. b. The loss of tropical forests also deprives about 250 migratory songbirds of their wintering homes. c. Lead poisoning of waterfowl is a side effect of hunting. Nonlead shot is used today, but lead is still in the environment and ingested by game birds.
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CHAPTER 19: BIRDS VOCABULARY ____contour feather____ ____supracoracoideus____ ____down feather____ ____crop____ ____filoplume feather____ ____gizzard____ ____powder down feather____ ____air sacs____ ____syrinx____ ____salt glands____ ____pneumatized____ ____alula____ ____pectoralis____ ____precocial____

____high aspect ratio____ ____low aspect ratio____ ____sun-azimuth orientation____ ____fovea____ ____wing slot____ ____altricial____

LEARN THE FOLLOWING 15 ORDERS. YOU WILL HAVE A QUIZ OVER THESE WITH NO NOTES!!! Order Struthioniformes (ostrich form)-15 species of flightless birds of Africa, S.America, and Australia. The ostrich of Africa, is the largest of living birds. Ex: ostrich, rheas, emus, & kiwis. Order Sphenisciformes (wedge form)-17 species of web-footed marine swimmers from Antarctica to the Galapagos. Ex: penguins. Order Procellariiformes (tempest form)-115 species worldwide. Marine birds with hooked beak Albatrosses are the larges of flying birds. Ex: albatrosses, petrels, shearwaters. Order Pelecaniformes (pelican form)-65 species worldwide especially tropics. Colonial fisheaters with throat pouch. Ex: pelicans, cormorants, boobies. Order Ciconiiformes (stork form)-120 species worldwide. Long-necked, long-legged mostly colonial waders and vultures. Ex: herons, bitterns, storks, flamingos & vultures. Order Falconiformes (falcon form)-310 species worldwide. Diurnal birds of prey. All are strong fliers with keen vision and sharp curved talons. Ex: eagles, hawks, falcons, condors, & buzzards. Order Anseriformes (goose form)- 160 species worldwide. Broad bills with filtering ridges at margins, long breastbone with low keel. Ex: swans, geese, & ducks. Order Galliformes (cock form)-290 species worldwide distribution. Chicken-like ground-nesting herbivores with strong beaks and heavy feet. Ex: quail, pheasants, turkeys & domestic fowl. Order Columbiformes (dove form)-320 species worldwide. Short necks, short legs and slender bill. Ex: pigeons, doves and the extinct dodo. Order Psittaciformes (parrot form)-3709 species pantropical distribution. Birds with hinged and movable upper beak and fleshy tongue. Ex: parrots & parakeets. Order Cuculiformes (cuckoo form)-150 species worldwide. Some lay their eggs in nests of smaller birds that raise the young cuckoos. Ex: cuckoo & roadrunners. Order Strigiformes (screech owl form) 185 species worldwide. Nocturnal predators with large eyes, powerful beaks and feet and silent flight. Ex: owls. Order Apodiformes (footless form)-435 species worldwide. Small birds with short legs and rapid wingbeat. Most hummingbirds are found in tropics, but there are 14 species in the U.S. Ex: swifts & hummingbirds. Order Piciformes (woodpecker form)-410 species worldwide. Birds with highly specialized bills and two two toes extending forward and two backwards. All nest in cavities. Ex.: woodpeckers, toucans, puffbirds. Order Passeriformes (sparrow form)-5900 species worldwide. Largest order containing 60% of birds. Most have a highly developed syrinx. Feet adapted for perching on stems. Young are altricial. Ex: mockingbird, jays, swallows, crows, nuthatches and creepers.

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