ORIGINS

The Diversity of Bi rds

It is easy to understand why so man)' 4 us are so fond oj birds. They afe lively; they are lovely; and they are

everywhere.

IAttenbol"Ough 1998. p 7]

illions of years ago, a small, bipedal reptile lived among the dinosaurs. From its skin evolved a novel and empowering structure-the feather. Feathered insulation enhanced its ability to control its high body temperature, th ere by increasing its activity and endurance. Extensions of feathers on forelimbs and tails led to powerful, gracef . .il £light. 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 mainraining the global fabric of biodiversity. As author David Quammen writes 10 1115 Song cif the Dodo (1996), the global tapestry of life on Earth IS in danger of unravelmg as key ecological threads are broken. Birds are one of those threads. Birds move 111 vast numbers across the hemispheres, playing essential roles as consumers of insects, pollinators of flowers, and dispersers of seeds. Birds are pivotal players Il1 ecosystem. dynamics and provide essential services to human societies. BIrds also serve as barometers of the health of ecosystems, pristine and altered, that serve mankin d. Understanding 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 geographical distribution. Chapter 2 examines the evolutionary history of birds from the origmal fossil bird, Archaeopteryx litllOgraphica, and birdlike dinosaurs to modern birds themselves. Chapter 3 briefly considers the reconstruction of evolutionary trees of relationships among birds, now informed by powerful new DN A-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 vertebrates. They are distinguished from other (modem) vertebrates by feathers, which are unique modifications of the outer skin. Compared with the scales of reptiles, feathers are filamentous, soft in texture, flexible, lightweight structures (Figure 1-1). No comparable structures exist ill other living vertebrates. Dead structures that wear easily and must be replaced

FIGURE 1-1 The courtship display of the King Bird-of-Paradise deploys its full array of elaborate feathers, including racquet-tipped wire tai I feathers. [From Ingram 1907]

(C)

FIGURE 1-2 The bills of birds correspond to their feeding specialties. (A) Red Crossbills extract seeds from pine cones .. (8) Northern Cardinals crack large. hard seeds. (C) Northern Shoveler ducks strain food from the mud. (D) 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 temperature. Lighrweight and strong, the long feathers of the wing generate lift and thrust for flight.

AJ] birds have bills, a distinctive attribute that facilitates instant recognition. The avian bill varies greatly in form and function bur 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 mammal of Australia.

Because birds lack teeth that chew food before swallowing, the avian digestive system IS specialized to process unmasticated food. The lack of teeth in birds appears to be a weight-reducing adaptation for flight, because teeth reqUlre a heavy jawbone for support. Instead of teeth, birds have a gizzard. The avian gizzard-a functional analogue of mammalian molars-v-is a large, strong, muscular structure used primarily for grinding and digesting tough food. The gizzards of grain eaters and seed eaters, such as turkeys, pigeons, and finches, are especially large and have powerfullayers of striated muscles. Turkey gizzards can pulverize English walnuts, steel needles, and surgical Iancets. The internal grinding surfaces of

5

THE DIVERSITY OF BIRDS

6

the gizzard are covered with a rough pleated or folded surface with many grooves and ridges. In some pigeons, it has strong, tooth-shaped projections. 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 gizzard is not so muscular in birds that eat softer foods such as meat, insects, or fruit; and, in Tap tars and herons, it may take the form of a large thinwalled sac.

Birds are feathered flying machines (Figure 1-3). Their wings and their ability to fly are familiar attributes but, unlike feathers, are not diagnostic features; bats and flying in. ects also have wings,

CHAPTER ONE

Wrist reduced to two bones for lightness

Wishbone (furcula) forms arch for strength without bulk

Backbone and pelvis act as shock absorbers on landing

Pygostyle supports tail feathers for braking and steering

Ankle bends forward

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 pygosryle. 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 a nd 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 A/<erbergs]

7

THE DIVERSITY OF BIRDS

(A)

(B)

(C)

FIGURE 1-4 (A) Leg bones of equal lengths contribute to the bala nee of long-legged birds. When a bird crouches to incubate Its egg, for example, leg bones of different lengths (6 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 to one another and to the metatarsals, [hereby creating a long, strong, single leg element, the ta rsometatarsus, which enables birds to walk on their toes rather than on the whole foot [After Storer 1971]

The entire avian body is structured tor flight Bird bones, for example, are typically lightweight structures, being spongy, strutted, and hollow. The skeleton generally is strengthened and reinforced through fuSIOns 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 itself is a higllly 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 011 land, a bird's center of gravIty IS positioned directly 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 rarsometatarsus-v-of long-legged birds ensures this relation, Foot-propelled diving birds such as loons have sacrificed balance on land for their considerable SWil1l111ll1g 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 then feet when on land,

Arboreal-tree-dwelling-specles, which constitute the majority of birds, have feet that tightly grip branches. }\mong 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, lockmg tbe

8

CHAPTER ONE

Hallux

Ankle joint

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 relaxes and the toes open. The foot of the songbirds-Order Passeriformesis 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 (Q sleep while perching. The large, opposable 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 ilight 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 (40D_44DC) over a wide range of ambient temperatures. The circulatory and respiratory systems of birds include a powerful four-chambered heart and efficient lungs, which deliver fuel and remove both waste and heat produced by metabolic 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 produced by other classes of vertebrates. Nurturing the growth of the embryos 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 Iiaisoris. As a result, the eggs In one nest may be of mixed paternities and even maternities. Mating systems, spacing behavior, and cooperation afford varied solutions to the challenges of successful reproduction in a highly competitive social world.

Birds have large,\vell-developed brains, 6 to 11 times as large as those of like-sized reptiles. Bird brains and primate brains exhibit functional lateralization, with left: hemispheric dominance associated with learning and innovation in vocal repertoires. Substantial learning by birds guides the mastery of complex motor tasks, social behavior, and vocahzations .

Highly developed neural systems and acute senses in birds mediate feats of communication and navigation. Birds, particularly the songbirds, have the greatest sound-producing capabilities of aJJ vertebrates. The syrinx of birds is a Ll11lgue sound-producing structure. In contrast with the syrinx, the larynx, the analogous structure in mammals, constrains mammalian vocalizations by its position and structural simplicrty. 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 infi·asounds-sounds below the hearing range of humans.

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 adaptation. The current classification of living buds 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 111 the Mesozoic era more than 150 million years ago had feathers and probably could fly after a fashion. Responding co ecological opportunities, subsequent 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 ostnch (Figure 1-6). Birds range in size ficrn only 2 grams (hummingbird) to 100,000 grams (ostrich).

The diversity ofbn·ds is due to the evolution of additional varied species adapted to different ecologies and behaviors, a phenomenon called adaptive radiation, Bill sizes and bill shapes change in relation to the types of food eaten. Leg lengths change in relation to habits of perching or

9··

THE DIVERSITY OF BIRDS

CHAPTER ONE

FOREST BIRDS Owl

AQUATIC BIRDS Penguin

OCEANIC BIRDS Albatross

WADING BIRDS AND SHOREBIRDS Plover

FLIGHTLESS AND GROUND BIRDS Ostrich

FIGURE 1-6 Birds have evolved along several major lines, each adapted to a particular mode of life. [From EJJolutioll of Vertebrates by E. H. Colbert. Copyright 19S5 John Wiley 80 Sons, inc; reprinted by permission of john Wiley G 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, turns tones, 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, gul1s, and skimmers, deep-water divers such as puffins, and the grouseEke seedsnipes of South American moorlands. All these related species are members of the Order Charadmformes (Figure 1-7).

The varied diets of modern birds include buds, fruits, nectar, seeds, invertebrates of all sizes, and vertebrates of many kinds, including carrion.

FIGURE 1-7 Shorebirds, gulls, and allies (Order Charadriiformes): (1) Pheasant-tailed [acana (jacanidae): (2) Snowy Sheathbill (Chionidae); (3) Eurasian Woodcock (Scolopacidae); (4) Atlantic Puffin (Alcidae); (5) Blacksmith Lapwing (Charadriidae);

(6) Ring-billed Gull (Laridae); (7) Black Skimmer (Rynchopidae).

11

THE DIVERSITY OF BIRDS

\" \

'-

2

./

(I:'

-'?1-~- 7

Number of Taxa
Order Families Genera Species
Tinamitormcs 1 Sl 47
IZheiformes 1 2 2
Struthioniformes 1 1 1
Casuariiformes 2 2 6
Dinomithiformes 1 1 3
Galliformes 5 80 290
Anseriforrnes " 52 162
_)
Sphcnisciforrnes 1 6 n
Caviifonues 1 1 5
Procellariiformes 4 26 112
Podicipediformes 1 6 22
Phoenicopreriforrnes 1 3 5
Ciconiiforrnes 3 39 116
Pelecaniformes 8 10 65
Falcon i form es 3 83 304
Cruiformes 11 61 212 Charadriiformes

17

88

Columbiforrnes 1 42 308
Psi ttaciformes 1 85 364
Opisthocorniformes 1 1 1
Musophagiformes 1 6 23
C L1 culi form es 1 35 138
Srri gi [onnes 2 29 180
Caprimulgiformes 5 22 118
Ap odiform cs 3 124 429
Coliiforrnes 1 2 6
Trogoniformes 1 6 39
Coraciifonnes 11 51 209
Piciforrnes _, 8 398
Passerifo rrnes 96 '1218 5753
Totals 193 2099 9702
Family classificacion and weals from Dickinson 2003.
12 Members

Tinarnous

Rheas

Ostrich Cassowaries, Emu Kiwis

Gallinaceous birds: grouse, quails, pheasants, chickens, curassows, guans, chachalacas, guineafowl, moundbuilders

Wate[fovvl: ducks, geese, swans, screamers Penguins

Loons

Tube-nosed seabirds: petrels, shearwaters,

albatrosses, storm petrels, diving petrels Grebes

Flamingos

Long-legged wading birds: storks, herons, ibises, spoon bills

Water birds with totipalrnate teet: cormorants, pelicans, anhingas, boobies, gannets, frigatebirds, tropicbirds, Harnmerkop, Shoebill

Raptors: falcons, caracaras, hawks, eagles,

Old Wodd vultures, kites, Osprey, Secretarybird, New Wodd vultures

Diverse terrestrial and marsh birds: rails, coots, sungrebes, cranes, Sunbittern, Kagu, Limpkin, seriemas, bustards, buttonquails, trumpeters, roatelos

367 Shorebirds and [heir relatives: sandpipers, plovers,

phalaropes, stilts, jacanas, painted-snipes, pratincoles, gulls and terns, seedsnipes,

shearhbills, skimmers, skuas, auks, sandgrouse Pigeons, doves

Pa~rots, macaws, lories, cockatoos Hoatzin

Turacos, plaintain-eaters Cuckoos

Owls, barn owls

Nightjars, potoos, frogmouths, Oilbird, owlet-

night jars

Swifts, crested swifts, hummingbirds Mousebirds

Trogons, quetzals

Kingfishers and allies: todies, motmots, be-eaters, rollers, Cuckoo Roller, hoopoes,

"mod hoopoes, hom bills

Woodpeckers and allies: wrynecks, piculets, barbers, tou cans, honeyguides, jacamars, puftbirds Perching birds, songbirds, passennes

Fruits, seeds, and insects nourish the majority of birds , especially the passerine land birds, whose adaptive radiation was coupled to those of flO\:vering 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, nurnerous species of flightless, herbivorous m03S evolved. The long, complex digestive tracts required for green leaves and the weight of slowly digesting plant matter also may limit the Hying abilities of- avian herbivores or favor flighrlessness, as in ostriches.

Corresponding to a diversity of diets is a diversity of bills (see Figure 1-2). A bini's bill is Its key adaptation for feeding. The size, shape, and strength of the bill prescribe the potential diet. The land carnivoreseagles, hawks, falcons, and owls-s-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 fi:oJIl the mud. The broad, flat bill of a duck is suitable for straimng mud, whereas the chisel-Eke bill of a woodpecker is suitable for digging 111m trees to reach insects. Marine 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 probll1g into the mud (Pigure 1-8). Nectar feeders, such as hummingbirds, probe their long, thin bills into floral nectar chambers and draw up nectar through tubed tongue tips. Their bill f0l1115 tend to match the lengths and curvatures of preferred flowers, which, in turn, depend on the birds

Eurasian

Eurasian Bar-tailed Ovster- Common Red

Curlew Godwit catcher Redshank Knot

Little

Gray Ringed Ruddy

Plover Plover Tu rnstone

FIGURE 1-8 Varied bill lengths enable shorebirds to probe to various depths in the mud and sand for food. Plovers feed on small invertebrates, mainly by surface pecking with their short bills. Common Redshanks and other species of waders with moderate bill lengths probe the top 4 centimeters of the substratum, which contain mallY worms, bivalves, and crustacea. Only the long-billed birds such as curlews and godwits can reach deep-burrowing prey such as lugworms. [A/tel' Cess-Custard 7975]

13

THE DIVERSITY OF BIRDS

14

CHAPTER ONE

Magnificent Hummingbird

Green Violet-Ear

Fiery-throated Hummingbird

1 cm 1----------1

.. ~ ... " .

. - I.:;

Volcano Hummingbird .:. ·1;~ . .

FIGURE 1-9 The lengths and curvatures of hummingbird bills match thOSE of their preferred flowers. [After Wolf et at. 1976]

FIGUR.E 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 sku II, revea ling the bony struts (trabeculae) in the upper Jaw and forehead The deeper, non trabecular areas of the upper jaw are shown in fine stippling; other nontrabecular bone is shown in heavier stippling. Lower jaw is not shown [From Bocl: 1966]

for pollination (Figure 1-9). Even slight differences in bill dimensions 111- fluence the race at which food can be consumed.

The bone configurations that constitute the bill, jaws, and palatal region 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 trabeculae (Figure 1-10). These struts make the bill much stronger than a hollow bill but add little weight. Covering both jaws is a horny sheath, or dumphotheca, 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 to 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 horieycreepers, 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 bi1J shapes and bill sizes led to a proIiferation 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 frorn flowers or probing bark crevices for insects (Figure 1-11).

Different modes of locomotion further expand the ecological opportunities of birds. Shorebirds, as already mentioned, include aerial, wadmg, and diving species. Birds soar through the sky, scurry and stride across the land, hop agilely frorn branch to branch, hitch up tree trunks, and swim powerfully to 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, runners, 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 tluttering through dense vegetation. At another extreme are the adaptations 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, powerful legs of wading and cursorial, or running, birds such as storks and ostriches. At the other extreme are the tiny teet and short legs of aerial specialists such as swifts. The long toes of herons and jacanas spread the bini's weight over a large surface area and facilitate walking 011 soft surfaces. Sandgrouse scurry on soft desert sands, and ptarmigan can walk on snow by virtue of snowshoe-like adaptations of their feet. Lobes

(Tex/ elm/ill lies (111 P'1ge J 8.)

15

THE DIVERSITY OF BIRDS

16

CHAPTER ONE

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 Railww 1916; dmwtng by 1-1. Douglas Pratt]

-

FIGURE 1-12 Penguins: (1) Chi nstrap Pengu In; (2) Rockllopper Pengui n; (3) Jackass Penguin; (4) King Penguin. juvenile (left). adult (dght).

17

THE DIVERSITY OF BIRDS

(G) Woodpecker

(E) Pta rmigan

(H) Eagle

eC) Ostrich

(F) Heron

(I) Warbler

FIGURE 1-13 The feet of birds reveal their ecological habits. Water birds have

CA) webbed or (S) lobed toes 1'01· swimm ing; terrestrial birds have toes specialized for (C) running, (D) scratching in dirt, (E.) walking on snow, or (F) wading. Other la\ld birds have feet designed for (G) climbing, (H) holding prey, or (I) perching. [From Wilson 1980, with permission from Scientific American, Inc.]

on the toes of coots and webbing between the toes of ducks aid swimmmg. 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 (Ricklefs 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 111.any 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-

18

less or agile. Parental care may be minimal or prolonged. Such attributes of lite history contribute to an individual bird's lifetime reproductive success. They arc as subject to evolutionary molding by the environment and by population dynamics as are the shapes of bills and wings.

Natural Selection and Convergence

We accept that the close fit between form and function portrayed so vividly by the diversity of birds and their life-history traits is due to evolutionary 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 predominance of individual organisms with advantageous traits. Healthy individuals leave more offspring than do sickly individuals. Camouflaged chicks an: more likely to escape predation and to reproduce themselves than are boldly colored chicks. To be flvored by natural selection, however, traits need not be dramatically better. Subtle or slight advantages in egg quality, camouflage, or agility increase in prevalence in a local population. Adaptation by natural selection is a process without plan or purpose, a process that gradually transforms the appearances and abilities of organisms.

A large finch bill can be so advantageous in times of food shortage that the average bill size in a population increases from one year to the next (Boag and Grant 1981). In 1976 and 1977, a severe drought gripped Daphne Island in the Cal.ipagos archipelago. Plants tailed to produce new crops of seeds, and seed densities dropped sharply, especially the densities of small seeds. Many finches starved. I n regard to the Medium C round Finch, individual birds with large, deep bills survived in greater numbers than did 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 (Figure 1-14). This intense natural selection was later reversed by the improved survival of small-billed birds during wet years, when small seeds vvere again plentiful.

The power of natural selection is perhaps best demonstrated through convergence-the independent evolution of similar adaptations in unrelated organisms. Adaptation to similar ecological roles causes unrelated species of birds to become superficially similar (i.e., to converge) ill details of appearance and behavior. For example, the meadowlarks of North American grasslands and the unrelated longclaws of the African grasslands are classic cases of convergence in color pattern. Both have streaked brown backs, bright yellow underparts with a black V OIl the neck, and white outer tail feathers. The meadowlark is related to the blackbirds (Icteridae), the longclaws to pipits and wagtails (Motacillidac), Another classic case of convergence is that of the northern ocean auks and the southern ocean penguins. From different aerial ancestors, species of compact

19

THE DIVERSITY OF BIRDS

20

CHAPTER ONE

Q) N

'",

co '" Q)

c

"2

rn s:

"0 Q)

3\

I

S N 1975

M M J S N 1976

M M J S N 1977

M M J S N 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

on Daphne Island favored individual birds with large bills able to crack the more abundant, large, hard seeds (bottom), Abbreviations, starting at left on x-axis: J, June;

S, September; N, November; j, January; M, March; M, May, [After Botu; 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 marine crustaceans and fish (Figure 1-15),

Biogeography

Biogeography is the study of the geographical distributions of plants and animals. For more than a century, biogeographers have divided Earth into six major faunal regions corresponding roughly to the major continental

Southern Hemisphere Adaptive Northern Hemisphere
Petrel-Penguin stock stage Gull-Auk stock
*,1" Wings used for ~
( submarine
flight only \
~.~ ~~=t;;tAuk
Penguins Stage C
I ~ Wings used for ~
I both r-,
I ( submarine
I ~ and aerial flight ~
I
I ~ Stage B ~
I Diving petrels Razorbill
I ~ ~
I Wings used For
I
:V aerial 1\
flight only
r ~ ~ \
Stage A ~UIIS
Petrels FIGURE 1-15 Convergent evolution of wing-propelled divers. Adaptive stages in the parallel evolution of two stocks of wing-propelled diving birds, the petrels to penguins and the gulls to auks, respectively. r From Storer 1960]

areas (Figure 1-16). Each faunal region Ius its characteristic birds: so-called endemic taxa or species, which arc 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 Africa. or the Ethiopian region, include ostriches. mousebirds, and turacos, Australia and New Guinea, the Australasian region, have emus. honeyeaters, and birds-or-paradise. South America, the Ncotropical region, has toucans, tinamous, and trumpeters,

Most avifaunas-v-rcgional assemblages of bird species-arc 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 landmass called Gondwanaland. Other species arose in recent colonizations of new islands or continents. The history of bird distributions can be viewed as a series of waves of adaptive radiations, moving north, south. east, and west from their ancestral origins. New groups of birds replaced older ones and in turn produced complex mosaics 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 avifaunas, plus diverse species groups that evolved only on that continentfor example, the colorful wood warblers.

Early avian colonists on each continent or major group of islands diversify locally in response to the ecological opportunities available to them. The diversity of finches on the GaLipagos Islands and of honeycreepers

21

THE DIVERSITY OF BIRDS

22

23

THE DIVERSITY OF BIRDS

FIGURE 1-16 The six major faunal regions (A) and some of their bird specialties:

(B) toucans (Neotropical region); (C) loons (Nearctic and Palearctic regions); (D) honeyeaters and (E) fairywrer-s (Australasian region); (F) turacos and (G) mousebirds (Ethiopian region).

24

CHAPTER ONE

Oil the Hawaiian Islands inspired the evolutionary theories of Charles Darwin. 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. wheatcars, 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 rnisclassified with 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 parental behavior and extraordinary vocal abilities. The anatomy and physiology of 1110St 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, whicb include terrestrial waders, aerial plungers. and wing-propelled divers. The adaptive radiation of Hawaiian honeycreepers illustrates the way in which bill forms can evolve in relation to feeding habits. In addition to their physical features. birds have diverse lite-history traits, mating systems, and reproductive strategies.

The birds of the world constitute geographical assemblages or- species: these large groups are called avifaunas. The six major avifaunas are the Neurctic (North America), Neotropical (Central and South America), Palearctic (Europe and Asia), Ethiopian (Africa south of the Sahara). Oriental (Southeast Asia), and Australasian (Australia and New Guinea). Each region has its characteristic birds.