MH PART 1 ORIGINSa, The Diversity of Birds It is easy to understand why so many of us are so fond of birds. They are lively; they are lovely; and they are everywhere [Attenborough 1998, p. 7] illions of years ago, a small, bipedal reptile lived among the dinosaurs. From its skin evolved a novel and em- powering structure—the feather. Feathered insulation enhanced its ability to control its high body temperature, thereby increas ing its activity and endurance. Extensions of feathers on forelimbs and tails led to powerful, graceful flight. Mastery of flight opened a world of ecological opportunities, and a new group of vertebrates—the Class Aves—evolved, and then thrived, The biggest conservation challenge facing the world is that of main taining the global fabric of biodiversity. As author David Quammen writes in his Song of the Dodo (1996), the global tapestry of life on Earth is in danger of unraveling as key ecological threads are broken. Birds are one of those threads. Birds move in vast numbers across the hemispheres, play- ing essential roles as consumers of insects, pollinators of flowers, and dis persers of seeds. Birds are pivotal players in ecosystem dynamics and pro~ vide essential services to human societies. Birds also serve as barometers of the health of ecosystems, pristine and altered, that serve mankind. Un- derstanding their diversity, their ecology, their history, and their future will serve us well. ‘This chapter previews the major features of the diversity of birds: their basic form, function, and biology; their major kinds; and their geograph- ical distribution, Chapter 2 examines the evolutionary history of birds from the original fossil bird, Archaeopteryx litagraphica, and birdlike di- nosaurs to modern birds themselves. Chapter 3 briefly considers the re- construction of evolutionary tees of relationships among birds, now informed by powerful new DNA-comparison techniques.4 (CHAPTER ONE Basic Characters of Birds Birds are two-legged (bipedal) vertebrates—the group of animals with backbones that also includes mammals, amphibians, reptiles, and fishes. Despite their diversity of form, birds are a well-defined group of verte- brates. They are distinguished from other (modern) vertebrates by feath- ers, which are unique modifications of the outer skin, Compared with the scales of reptiles, feathers are filamentous, soft in texture, flexible, light weight structures (Figure 1-1). No comparable structures exist in other living vertebrates. Dead structures that wear easily and must be replaced a FIGURE 1-1 The courtship display of the King Bird-of-Paradise deploys its full array of elaborate feathers, including racquet-tipped wire tail feathers. [From Ingram 1907)w © FIGURE 1-2 The bills of birds correspond to their feeding specialties. (A) Red Crossbills extract seeds from pine cones. (8) Norther Cardinals crack large, hard seeds. (C) Northern Shaveler ducks strain food from the mud. (0) Reddish Egrets spear small fish. (E) Golden Eagles tear apart the flesh of their prey. regularly, feathers are essential for both temperature regulation and flight. They insulate the body and help birds to maintain their high body tem- perature. Lightweight and strong, the long feathers of the wing generate lift and thrust for flighe. All birds have bills, a distinctive ateribute that facilitates instant recog- nition, The avian bill varies greatly in form and function but is always toothless and covered with a horny sheath (Figure 1-2). The avian bill has no exact parallel among other extant vertebrates; it is approximated only by the snout of the duck-billed platypus, a strange, egg-laying mar mal of Australia. Because birds lack teeth that chew food before swallowing, the avian digestive system is specialized co process unmasticated food. The lack of teeth in birds appears to be a weight-reducing adaptation for flight, be- cause teeth require a heavy jawbone for support. Instead of teeth, birds have a gizzard. The avian gizzard—a functional analogue of mammalian molars—is a large, strong, muscular structure used primarily for grinding and digesting cough food. The gizzards of grain caters a such as turkeys, pigeons, and finches, are especially large and have pow- erful layers of striated muscles. Turkey gizzards can pulverize English wal- nuts, steel needles, and surgical lancets. The internal grinding surfaces of id seed eaters, 5 THE DIVERSITY OF BIRDS6 ‘CHAPTER ONE the gizzard are covered with a rough pleated or folded surface with many grooves and ridges. In some pigeons, it has strong, tooth-shaped projec tions. The gizzard can also contain large quantities of grit, which grinds food. The gizzards of moas, extinct ostrichlike birds of New Zealand, have been found to contain as much as 2.3 kilograms of grit. The giz zard is not so muscular in birds that eat softer foods such as meat, insects, or fruit; and, in raptors and herons, it may take the form of a large thin- walled sac. Birds are Feathered flying machines (Figure 1-3). Their wings and eheir ability to fly are familiar attributes but, unlike feathers, are not diagnos- tic features; bats and flying insects also have wings. Hand bones: are fused Wrist reduced to ‘wo bones for lightness Wishbone (urcula) forms elbow arch for strength bo without bulk Uncinate process. Neck has 15 vertebrae Backbone and pelvis ‘act as shock absorbers ‘on landing Wing, muscles attach to keel of sternum Pygostyle suppor tail feathers for braking and steering. Ankle bends forward Knee bends back FIGURE 1-3 Adaptations for flight. Supporting the wings of the Herring Gull is a strong but lightweight skeleton. An enlarged, keeled sternum houses and anchors the large breast muscles that empower the wings. The bones of the hand and wrist, which support and maneuver the primary flight feathers, are reduced in number and fused for extra strength. Similarly, the pygostyle, made of fused tail vertebrae, supports and controls the tail feathers, which are used for braking and steering. Strengthening the body skeleton are fusions of the pelvic bones and associated vertebrae, plus horizontal rib projections called uncinate processes. The furcula, or wishbone, compresses and rebounds like a powerful spring in rhythm to the beat of the wings. (After Pasquier 1983; drawing by Biruta Akerberys]Tibiotarsus Ankle 0 @ © FIGURE 1-4 (A) Log bones of equal lengths contribute to the balance of long-legged birds. When a bird crouches to incubate its egg, for exemple, leg bones of different lengths (B and C) would displace the center of gravity. What appears at first glance to be a backward-bending knee joint is really the ankle joint. in birds, the foot bones (three tarsals) are fused both 10 one another and to the metatarsals, thereby creating 2 long, strong, single leg element, the tarsometatarsus, which enables birds to walk on their toes rather than on the whole foot. [After Storer 1971] Tarsometatarsus The entire avian body is structured for flight. Bird bones, for exan ple, are typically lightweight structures, being spongy, strutted, and hol- low. The skeleton generally is strengthened and reinforced through fi sions of the bones of the hands, head, pelvis, and feet. Horizontal, backward-curved projections—called uncinate processes—on the ribs overlap other ribs and so strengthen the walls of the body. The furcula, or wishbone, compresses and rebounds like a powerful spring in rhythm to the beat of the wings. The wing iself is a highly modified forelimb that, with a few remarkable exceptions, is nearly incapable of functions other than flight. Fused hand bones support and maneuver the large and powerful primary flight feathers. For stable balance on land, a bird's center of gravity is positioned di- rectly over and between its feet, particularly when the bird perches, squats, or rises (Figure 1-4). The equal length of the two main leg bones—the tibiotarsus and tarsometatarsus—of long-legged birds ensures this relation, Foot-propelled diving birds such as loons have sacrificed balance on land for their considerable swimming abilities. For efficient propulsion, they have powerful legs situated at the rear of a streamlined body, which places their center of gravity far forward of their feet when ‘on and. Arboreal—tree-dwelling—species, which constitute the majority of birds, have feet tha tightly grip branches. Among the features of such feet, are long tendons that pass around the backside of the ankle joints. When a bird bends its joints to squat, the tendons automatically flex, locking the z THE DIVERSITY OF BIRDS8 CHAPTER ONE TTendons Ankle joint ‘Tendons FIGURE 1-5 When a perching bird squats, the leg tendons, which are located on the rear side of the ankle, automatically cause the toes to grip. [From Wilson 1980, with permission from Scientific American, Inc.) toes around the branch (Figure 1-5). When a bird stands, the tension re~ laxes and the toes open. The foot of the songbirds—Order Passeriformes— is perhaps the most advanced in this respect. A special system of ridges and pads between the tendons that flex the toes and the insides of the toe pads acts as a natural locking mechanism and permits birds to sleep while perching. The large, opposible single rear toe, or hallux, which enhances the ability of a bird to grip a branch, is unusual among vertebrates, Avian physiology accommodates the extreme metabolic demands of flight and temperature regulation. The red fibers of avian flight muscles have an extraordinary capacity for sustained work and can produce heat by shivering (see Chapter 6). Birds are endothermic; that is, they are warm-blooded and maintain high body temperatures (40°-44°C) over a wide range of ambient temperatures. The circulatory and respiratory sys- tems of birds include a powerful four-chambered heart and efficient lungs, which deliver fuel and remove both waste and heat produced by meta~ bolic activities. The reproductive systems of birds also are unusual, Birds produce large, richly provisioned external eggs, the most elaborate reproductive cells of any animal, No bird species bears live young like those pro- duced by other classes of vertebrates. Nurturing the growth of the em- bryos in the eggs and of the young after they hatch requires dedicatedparental care. Most birds form monogamous pair bonds, some for life; but many, it turns out, engage in additional sexual liaisons. As a result, the eggs in one nest may be of mixed paternities and even maternities. Mating systems, spacing behavior, and cooperation afford varied solu tions to the challenges of successful reproduction in a highly compet- itive social world. Birds have large, well-developed brains, 6 to 11 times as large as those of like-sized reptiles. Bird brains and primate brains exhibit func tional associated with learning and innovation in vocal repertoires. Substantial learning by birds guides the mastery of complex motor tasks, social behavior, and vocalizations. Highly developed neural systems and acute senses in birds mediate feats of communication and navigation. Birds, particularly the song- birds, have the greatest sound-producing capabilities of all vertebrates ‘The syrinx of birds is a unique sound-producing structure. In contrast with the syrinx, the larynx, the analogous structure in mammals, con strains mammalian vocalizations by its position and structural simplic- ity. Birds can navigate by using patterns of Earth’s magnetism, celestial cues, and perhaps polarized light. Their highly developed color vision reaches into the near-ultraviolet range of the spectrum. Their broad hearing range encompasses infrasounds—sounds below the hearing range of humans, teralization, with left hemispheric dominan Adaptive Radiation of Form and Function Roughly 300 billion birds now inhabit the Earth. The variety of birds is the grand result of millions of years of evolutionary change and adap- tation. The current classification of living birds arranges 30 orders, 193, families, 2099 genera, and at least 9700 species (Table 1-1, page 12) Yet this number is only a small fraction of the number of species that, have existed since the age of dinosaurs. The earliest birds in the Meso zoic era more than 150 million years ago had feathers and probabl could fly after a fashion. Responding to ecological opporcunities, sub- sequent birds diversified in form and function, From the fundamental anatomy of their common ancestor evolved perching songbirds such as robins; nocturnal forest hunters such as owls; aquatic divers such as penguins; oceanic mariners such as albatrosses; shoreline waders such as plovers; and large, flightless ground birds such as the ostrich (Figure 1-6). Birds range in size from only 2 grams (hummingbird) to 100,000 grams (ostrich). The diversity of birds is due to the evolution of additional varied species adapted to different ecologies and behaviors, a phenomenon called adap- tive radiation. Bill sizes and bill shapes change in relation to the ty of food eaten. Leg lengths change in relation to habits of perching or 9 THE DIVERSITY OF BIRDS10 (CHAPTER ONE FOREST BIRDS SONGBIRDS Owl Robin UPLAND BIRDS Pheasant WADING BIRDS AND SHOREBIRDS Plover AQUATIC BIRDS OCEANIC BIRDS FLIGHTLESS AND GROUND BIRDS Penguin Albatross Ostrich FIGURE 1-6 Birds have evolved along several major lines, each adapted to a particular mode of life. [From Evolution of Vertebrates by E. H. Colbert. Copyright 1955 Joh Wiley @ Sons, Inc.; reprinted by permission of John Wiley @ Sons, Inc.| terrestrial locomotion, and wing shapes change in relation to patterns of flight. For example, from a single ancestral species of shorebird evolved aerial pirates such as skuas and plunging divers such as terns, as well as a host of waders, including sandpipers, plovers, turnstones, stilts, jacanas, snipes, woodcocks, curlews, and godwits, each with characteristic leg lengths and bill lengths, shapes, and curvatures. As varied as the habitats that they occupy, shorebirds also include aerial pratincoles, gulls, and skim- mers, deep-water divers such as puffins, and the grouselike seedsnipes of South American moorlancs. All these related species are members of the Order Charadriiformes (Figure 1-7). The varied diets of modern birds include buds, fruits, nectar, seeds, inds, including carrion, invertebrates of all sizes, and vertebrates of many ooo FIGURE 1-7 shorebirds, gulls, and allies (Order Charadriiformes): (1) Pheasant-talled Jacana (Jacanidae); (2) Snowy Sheathbill (Chionidae); (3) Eurasian Woodcock (Gcolopacidae); (4) Atlantic Puffin (Alcidae); (5) Blacksmith Lapwing (Charadriidae); (6) Ring-billed Gull (Laridae); (2) Black Skimmer (Rynchopidae)un THE DIVERSITY OF BIRDS[EVABLE 1-1 The orders of the birds of the world Number of Taxa Order Families Genera Species Members Tinamitormes 1 9 47 Tinamous Rheiformes 1 2 2 Rheas ‘Struthioniformes 1 1 1 Ostrich Casuariiformes 2 2 6 Cassowaries, Emu Dinomichiformes 1 1 3 Kiwis Galliformes 5 80 290 Gallinaceous birds: grouse, quails, pheasints, chickens, curasows, guans, chachalacas, guineafowl, moundbuilders | Anseriformes 3 52 162 Waterfowl: ducks, geese, swans, screamers | Sphenisciformes 1 6 17 Penguins Gaviiformes 1 1 5 Loons Procellaiiformes 4 26 112 Tube-nosed seabirds: petrels, shearwaters, albatrosses, storm petrels, diving petrels Podicipediformes 1 6 22° Grebes Phoenicopteriformes 1 3 5 Flamingos | Ciconitformes 3 39 116 Long-legged wading birds: storks, herons, ibises, spoonbills Pelecaniformes 8 10 65 Water birds with totipalmate feet: cormorants, pelicans, anhingas, boobies, gannets,frigatebirds, copicbirds, Hammerkop, Shoebill Falconiformes 3 83 304 Raptors: falcons, caracaras, hawks, eagles, Old World vuleures, kites, Osprey, Secretarybird, New World vultures Gruiformes i 61 212 Diverse terrestrial and marsh birds: rails, coots, sungrebes, cranes, Sunbitcern, Kagu, Limpkin, seriemas, bustards, buetonquails, crumpeters, roatelos Charadbiformes 88 367. Shorebirds and their relatives: sandpiper, plovers, phalaropes, stilts, jacanas, painted-snipes, pratincoles, gulls and cerns, seedsnipes, sheathbills, skimmers, skuas, auks, sandgrouse Columbiformes 1 42308 Pigeons, doves Paittacitormes 1 85-364 Parrots, macaws, lories, cockatoos Opisthocomiformes 1 1 1 Hoatzin Musophagiformes i 6 23 Turacos, plaintain-eaters Cuculiformes 1 35.138 Cuckoos Strigiformes 2 29 180 Owls, bam owls Caprimalgiformes 5 22 118 Nightjars, potoos, frogmouths, Oitbird, owlet- nightjars Apodiformes 3 124 429.—_Swifts, crested swifts, hummingbirds Coliiformes 1 2 6 Mousebirds ‘Trogoniformes 1 6 39 Trogons, quetzals Coraciiformes ul 51 209 Kingfishers and allies: todies, motmots, bee-eaters, rollers, Cuckoo Roller, hoopoes, wood hoopoes, hornbills Piciformes 5 8 398 Woodpeckers and allies: wrynecks, piculets, barbets, toucans, honeyguides, jacamars, puffbirds Passeriformes 96 12185753. Perching birds, songbirds, passerines Totals 193-2099 9702 Family clasification and totals from Dickinson 2003. 12Fruits, seeds, and insects nourish the majority of birds, especially the passer- ine land birds, whose adaptive radiation was coupled to those of flower- ing plants and their associated insects. Few birds are specialized herbivores; apparently, mammals have usurped most of the terrestrial grazing and browsing niches. In the absence of mammals in New Zealand, numerous species of flightless, herbivorous moas evolved. The long, complex diges- tive cracts required for green leaves and the weight of slowly digesting plant matter also may limit the fying abilities of avian herbivores or favor flightlessness, as in ostriches. Corresponding to a diversity of diets is a diversity of bills (sce Figure 1-2). A bird’s bill is its key adaptation for feeding, The size, shape, and strength of the bill prescribe the potential diet. The land carnivores— eagles, hawks, falcons, and owls—have strong, hooked beaks with which they tear flesh and sinew. Other bill types tear meat, spear fish, crack seeds, probe crevices, or strain microscopic food from the mud. The broad, flat bill of a duck is suitable for straining mud, whereas the chisel-like bill of a woodpecker is suitable for digging into trees to reach insects. Ma- rine predators, such as penguins and cormorants, have bills with curved projections that direct fish toward the esophagus. The varied lengths and curvatures of shorebird bills determine which prey they can reach by prob- ing into the mud (Figure 1-8), Nectar feeders, such as hummingbirds, probe their long, thin bills into floral nectar chambers and draw up nec tar through tubed tongue tips, Their bill forms tend co match the lengths and curvatures of preferred flowers, which, in turn, depend on the birds turasion Lite Eurasian Bartailed Oysters” Common Red Gray Ringed Ruddy Curlew” Godwit™ Giteher Redshank Knot Plover Plover Tumstone FIGURE 1-8 Varied bill lengths enable shorebirds to probe to various depths in the ‘mud and sand for food. Ptovers feed on small invertebrates, mainly by surface pecking with their short bill. Common Redshanks and other species of waders with moderate bill lengths probe the top 4 centimeters of the substratum, which contain many worms, bivalves, and erustacea. Only the longbilled birds such as curlews and godwits can reach deep-burtowing prey such as lugworms. [After Goss-Custard 1975] 13 THE DIVERSITY OF BIRDS14 CHAPTER ONE Green Violet-Ear OO Fiery-throated Hummingbird aS Volcano Hummingbird FIGURE 1-9 The lengths and curvatures of hummingbird bills match those of their preferred flowers. [After Wolf etal. 1976] ‘Trabeculae FIGURE 1-10 The form of their large bills enables finches such as the Northern Cardinal to bite hard seeds without straining the nasofrontal hinge (located between bill and skull) with excessive shear forces. Shown here is a cross section of a cardinal skull, revealing the bony struts (trabeculae) in the upper jaw and forehead. The deeper, nontrabecular areas of the upper jaw are shown in fine stippling: other nontrabecular bone is shown in heavier stippling, Lower jaw is not shown. [From Bock 1966]for pollination (Figure 1-9). Even slight differences in bill dimensions in- fluence the rate at which food can be consumed. The bone configurations that constitute the bill, jaws, and palatal re- gion are an engineer’s delight. The avian bill is not rigid; birds can flex or bend the upper half of the bill, an ability called cranial kinesis (see Zusi 1984). The upper mandible, or maxilla, is a flattened, hollow, bony cone reinforced internally by a complex system of bony struts called wabeculae (Figure 1-10). These struts make the bill much stronger than a hollow Dill but add little weight. Covering both jaws is a horny sheath, or sham- photheca, which may have sharp cutting edges (as in boobies), numerous toothlike serrations (as in mergansers), or well-developed notches (as in falcons and toucans). Woodcocks can open just the tip of their bills co grasp earthworms deep in the mud. The variety of bill forms that can evolve in the process of adaptive radiation is seen in the Hawaiian honeycreepers, which apparently evolved from a flock of small finches that strayed out over the Pacific Ocean from Asia or North America millions of years ago. The finches made landfall on one of the Hawaiian Islands, then flourished and spread throughout the archipelago. Isolated populations changed in genetic composition and appearance, at first imperceptibly and then conspicuously. Subtle changes in bill shapes and bill sizes led to a pro- liferation of bill types and their related feeding behaviors: from heavy grosbeak-like bills for cracking large legume seeds to long sicklelike bills for sipping nectar from flowers or probing bark crevices for in- sects (Figure 1-11). Different modes of locomotion further expand the ecological oppor- tunities of birds, Shorebirds, as already mentioned, include aerial, wad- ing, and diving species. Birds soar through the sky, scurry and stride across the land, hop agilely from branch to branch, hitch up tree trunks, and swim powerfully co great depths in the sea. The combination of forelimbs adapted for flight and hindlimbs for bipedal locomotion gives birds a tremendous range of ecological options. There are specialized flying birds, as well as specialized swimmers, run ners, waders, climbers, and perchers. Wing shapes and modes of flight range from the long, narrow wings of the albatross, adapted for soaring over the oceans, to the short, round wings of wrens, adapted for agile fluttering through dense vegetation. At another extreme are the adapta- tions of wing-propelled diving birds, such as penguins, which use their flipperlike wings to move underwater (Figure 1-12). Like the structures of bills and wings, the anatomy of feet and legs corresponds to different life styles (Figure 1-13, page 18). At one extreme are the long, powerfal legs of wading and cursorial, or running, birds such as storks and ostriches. At the other extreme are the tiny feet and short legs of aerial specialists such as swifts. The long toes of herons and jacanas spread the bird’s weight over a large surface area and facilitate walking on soft surfaces. Sandgrouse scurry on soft desert sands, and ptarmigan can walk on snow by virtue of snowshoe-like adaptations of their feet. Lobes (Text continues on page 18.) "THE DIVERSITY OFa CHAPTER ONE re @ a a= 7 e <> ong ate FIGURE 1-11 A classic example of adaptive radiation: Hawaiian honeycreepers have evolved bills that range from thin warblerlike bills to long sicklelike bills to heavy grosbeak-like bills, [From Raikow 1976; drawing by H, Douglas Pratt]THE DIVERS FIGURE 1-12 Penguins: (1) Chinstrap Penguin; (2) Rockhopper Penguin; (3) Jackass Penguin; (4) King Penguin, juvenile (left), adult (right)18 (0) Pheasant (G) Woodpecker (6) Prarmigan (6) Heron (H) Eagle () Warbler FIGURE 1-13 The feet of birds reveal their ecological habits. Water bieds have (A) webbed or (B) lobed toes for swimming; terrestrial birds have toes specialized for (C) running, (0) scratching in dirt, (E) walking on snow, or () wading. Other land birds have feet designed for (G) climbing, (H) holding prey, or (1) perching. [From Wilson 1980, with permission from Scientific American, Inc.| on the toes of coots and webbing between the toes of ducks aid swim- ming. Climbing birds such as woodpeckers have large, sharply curved claws; nuthatches climb downward by gripping a tree’s bark with a large claw on the hind toe. Life Histories In addition to their form and function, birds have diversified in all aspects of their seasonal and social biology. Reproductive rate, adult life span, and age at maturity, differ more than tenfold among species (Ricklef 2000a). Albatrosses are long-lived and lay only one egg at a time. Small songbirds, instead, tend to have short life spans and to raise many young together. Some species lay large eggs for their body size; other species lay small and lightly provisioned eggs. Newly hatched young may be help-less or agile. Parental care may be minimal or prolonged. Such attributes of lite history contribute to an individual bird’s lifetime reproductive sue~ cess. They are as subject to evolutional by population dynamics as are the shapes of bills and wings nolding by the environment and Natural Selection and Convergence We accept that the close fit benween form and function portrayed so vividly by the diversity of birds and their life-history traits is due (0 evo- Jutionary adaptation through natural selection, As set forth by Charles Darwin in 1859 and confirmed subsequently through experiment and independent observation, natural selection is simply the predictable pre- dominance of individual organisms with advantageous traits. Healthy individuals leave more offspring than do sickly individuals, Camouflaged chicks are more likely to escape predation and co reproduce themselves than are boldly colored chicks. To be favored by natural selection, how= ever, traits need not be dramatically better. Subcle or slight advantages in egg quality, camouflage, or agi population, Adaptation by natural selection is a process without plan or purpose, a process that gradually transforms the appearances and abilities of organisr A large finch Dill can be so advantageous in times of food shortage that the average bill size in a population increases from one year tw the next (Boag and Grant 1981), In 1976 and 1977, a severe drought gripped Daphne Island in the Galipagos archipelago. Plants failed to produce new crops of sveds, and seed densities dropped sharply, especially the densities of small seeds. Many finches starved. In regard to the Medium Ground Finch, individual birds wich large, deep bills survived in_ greater numbers than diel those with small bills because the large-billed birds could crack the remaining larger, harder seeds. The result was a dramatic increase in average bill size over only one year’s time, due to natural selection (F ure I-14), This intense natural selection was later reversed by the im= proved survival of small-billed birds during wet years, when small seeds were again plentiful, The power of natural selection is perhaps best demonstrated through convergence—the independent evolution of similar adaptations in unre: lated organisms, Adaptation to similar ecological roles causes unrelated species of birds to become superficially similar (ie.. «© converge) in de- tails of appearance and behavior. For example, the meadowlarks of North American grasslands and the unrelated longelaws of the African grasslands are classic cases of convergence in color pattern, Both have streaked brown, backs, bright yellow underparts with a black Von the neck, and white outer tail feathers, The meadowlark is related co the blackbirds (eteri- dae), the longelaws to pipits and wagtails (Motacillidae), Another classic case of convergence is that of the northem ocean auks and the southern ocean penguins. From differen aerial ancestors, species of compact ¥y increase in prevalence in a local 19 THE DIVERSITY OF BIRDS20 (CHAPTER ONE —— seed hardness —- SNJ|MMJSNJMMJSNJMMJ)SN4 1975] 1978 | 1977 | 1978 FIGURE 1-14 increase in bill size (top) in the Medium Ground Finch during a period Of drought that resulted in intense natural selection. Failure of the usual seed crop fon Daphne Island favored individual birds with large bills able to crack the more abundant, large, hard seeds (boitom). Abbreviations, starting at left on x-axis: J, June; 'S September; N, November; J, January: M, March; M, May. [After Boag and Grant 1981] black-and-white seabirds have evolved in both ocean regions. including flightless forms that use their wings to propel themselves underwater to capture maine crustaceans and fish (Figure 1-15), Biogeography Biogeography is the study of the geographical distributions of plant animals. For more than a century, biogeographers have divided Earth into six major faunal regions corresponding roughly to the major continental21 Southern Hemisphere Adaptive Northern Hemisphere Petrel~Penguin stock stage ‘GullAuk stock Wings used for — submarine flight only e Penguins ’ Stage C Tt Great Auk m | Wings used for ‘both submarine so and aerial flight, we Diving pete Suoge 8 TSE razorit ) Wings used for "rial | flight only stage A ” 7 Gu petrels FIGURE 1-15 Convergent evolution of wing-propelled divers, Adaptive stages in the paralle! evolution of two stocks of wing-propelled diving birds, the petrels to penguins and the gulls to auks, respectively. [From Storer 1960] areas (Figure 1-16). Each endemic taxa or species, which are found nowhere else, and other birds that represent major adaptive radiations of more widespread taxa. ‘Waxwings and loons are restricted to North America and Eurasia, the Nearctic and Palearctic regions, respectively. The birds that are endemic to Aftica, or the Ethiopian region, include ostriches, mousebirds, and turacos. Australia and New Guinea, the Australasian region, have emus. honeyeaters, and birds-of-paradise. South America, the Neotropical region, has toucans. tinamous, and trumpeters. Most avifaunas—regional assemblages of bird species—are mixtures of species of varied ages and origins, Some species trace back more than 60 million years to the rearrangement of the continents from the early land- mass called Gondwanaland. Other species arose in recent colonizations of new islands or continents, The history of bird distributions can be viewed a8 a series of waves of adaptive radiations, moving north, south, east, and west from their ancestral origins. New groups of birds replaced older ones, nd in tum produced complex mosaies of ancient. recent, and new colonists from different regions. The birds of North America include old and new colonists from Asia and South America, remnants of ancient avi- faunas, plus diverse species groups that evolved only on that continent— for example, the colorful wood warblers ly avian colonists on each continent or major group of islands di- y locally in response to the ecological opportunities available to them. The diversity of finches on the Gakipagos Islands and of honeyereepers anal region has its characteristic birds: so-called THE DIVERSITY OF BIRDS7 So? PER [A | k aun Lol hab’ ah | FT, MNT2B THE DIVERSITY OF BIRDS FIGURE 1-16 The six major faunal regions (A) and some of their bi (8) toucans (Neotropical region); (C) loans (Nearctic and Palearctic 1 and (E) fainywrens (Australasian region); (F) turacos and (G) mousebi rd specialties: gions); (D) honeyesters ids (Ethiopian region) FF24 CHAPTER ONE on the Hawaiian Islands inspired the evolutionary theories of Charles Dar- win, Just as dramatic is the diversity of the tyrant flycatchers of South America, which radiated to include new species that look and function like shrikes, wheatears, tits, warblers, pipits, or thrushes from other parts of the world. Australian land birds, related genetically most closely to one another, matched so well the external attributes of shrikes, flycatchers, and small insect-eating warblers that, until 1990, they were misclassified wich superficially similar species native to Europe and Asia, Summary Characterized as vertebrates with feathers, birds have distinctive bills, maintain high body temperatures, produce large external eggs, and have elaborate parcntal behavior and extraordinary vocal abilities. The anatomy and physiology of most birds are adapted for flight. The diversity of birds is due to millions of years of divergence and adaptation by natural selection, The process of adaptive radiation is well illustrated by the members of the Order Charadriiformes, which include terrestrial waders, aerial plungers. and wing-propelled divers. The adap- tive radiation of Hawaiian honeycreepers illustrates the way in which bill forms can evolve in relation to feeding habits, In addition to their phys- ical features. birds have diverse life-history traits, mating systems, and reproductive strategies. ‘The birds of the world constitute geographical assemblages of species: these large groups are called avifiunas. The six major avifiunas are the Nearetie (North America), Neotropical (Central and South America), Palearctic (Europe and Asia), Ethiopian (Africa south of the Sahara), Ori- ental (Southeast Asia), and Australasian (Australia and New Guinea). Each region has its chai teristic birds.