A few years ago, I joined a birding tour of Ghana. After several days of enjoying such exotic species as drongos, hornbills, and pratincoles, we encountered a beautiful red and black finch, the Black-bellied Seedcracker (Pyrenestes ostrinus).1 I was delighted to see this species because I had long known, and had described in my textbook of evolutionary biology, a study of this species by Thomas Smith,2 a professor at University of CaliforniaâLos Angeles. Smith had followed the life of members of a population in Cameroon by fitting each individual with a unique combination of colored leg bands. Bill size is highly variable in seedcracker populations; most birds have either small or large bills, although a minority are intermediate (plate 1). Smith found that large-billed birds feed more efficiently on the large, hard seeds of one species of sedge and small-billed birds handle the small seeds of another sedge more efficiently. Large-billed and small-billed birds both had higher rates of survival than intermediate birds: a striking example of natural selection in action. By occupying somewhat different âecological niches,â birds with different genotypes (specific combinations of genes) persist. Years later, when the study of genomes had advanced, Smith and his collaborators determined that the inherited difference in bill size is caused by different forms of a single gene (called IGF1, or insulin-like growth factor 1).3 As we admired the seedcracker, I told my companions this story. One of them exclaimed, âSo thatâs why it doesnât look like the picture in the field guide! I wondered if the book was wrong.â He was intrigued by the idea that different members of a species have different diets and ways of life.
Some birders are focused on seeing and listing species; others are curious about the lives and features of the birds they see. Once in a while a fellow birder, knowing that Iâm a biologist, will ask me a question. Sometimes it is along the lines of âhow can birds fly so fast through dense vegetation without hitting it?â or âhow can a tiny Blackpoll Warbler (Setophaga striata) fly nonstop from New England to Venezuela?â I awkwardly answer that I donât know much about how birds achieve these amazing feats because those are topics studied by biologists who specialize in bird physiology or brain function, and I havenât followed those fields since I was a student. Some other questions, though, tempt me to say more than they may want to hear. (And I can resist anything but temptation.4) Why do some bird species have different color morphs? Why are males more brightly colored than females in some species but not others? Why do albatrosses and many other sea birds lay only one egg? How come I can see more bird species in a two-week birding tour in Peru than in an entire year in eastern North America? Why do they keep changing bird classifications, and how do they know falcons are closer to parrots than to hawks?
Most questions about birds fall into two categoriesâhow and whyâthat correspond to two major kinds of biological research. Much of biology poses âhowâ questions: it aims to understand how organisms functionâhow the molecular, cellular, and organ components of an organism work, here and now, without reference to how they came to be. âWhyâ questions are the province of evolutionary biology. We ask why a Eurasian Golden Oriole (Oriolus oriolus) or an American Baltimore Oriole (Icterus galbula) is brightly colored because we understand that it could have been otherwise: something in its historyâin its evolutionâcaused it to be bright rather than drab. For every characteristic of every species, we can ask âhowâ questions about its functional role (if any) in an organismâs lifetime, complemented by âwhyâ questions about its origin. All species of birds have evolved from a single ancestral species (âcommon ancestorâ), which was one of a great many species of vertebrates that all evolved from a single, more ancient, common ancestor; this, in turn, was a descendant of the ur-ancestor of all animals, from sponges to primates. And so every feature of every bird, from its DNA sequences to its behaviors, has come into existenceâhas evolvedâduring this history of descent.
Evolutionary biologists attempt to develop broad principles that can explain all these features of all species. Evolutionary biology illuminates every area of biological research and every group of organisms. The geneticist Theodosius Dobzhansky, who helped to shape modern evolutionary biology, rightly wrote that ânothing in biology makes sense except in the light of evolution.â5 There are biologists who study biochemical processes within cells and biologists who study how these processes evolvedâand likewise for the structure and function of genomes, brains, and hormones. Among ornithologists, some take a mostly functional approach, and others a more evolutionary approach, to bird physiology, morphology,6 behavior, and life histories. Others are devoted to understanding the history of bird evolutionâhow and when birdsâ form, behavior, habitat use, and geographical distribution diversified during their descent from their common ancestor. The amount of research that bears on bird evolution is immense: when I entered âevolution and bird*â in a search engine (Web of Science), it yielded 73,200 articles in scientific journals.7 Variant search terms would add many more.
So for almost any question we might ask about how birds evolved, there is plenty of research on which to draw. Nevertheless, the known is far less than the unknown. Questions such as âhow do new species form?â and âwhy do female birds prefer flashy males?â are debated and are the subjects of active research. And while we may be able to provide a general answer to a question (e.g., why do birdsâ bills differ in shape?), there may not be a definitive answer for a particular species. (I donât know of any research about why the bill of the Groove-billed Ani [Crotophaga sulcirostris] is grooved.) Evolutionary biologists strive, instead, to develop theories that should apply to a wide range of species but which require detailed information to explain particular cases. For example, there are several models8 to account for genetic polymorphismâthe persistence of two or more genetically different types within a population, such as the color âphasesâ of the Tawny Owl (Strix aluco) and the Eastern Screech Owl (Megascops asio). Information about the survival and reproduction of each form, under several environmental conditions, may be needed to match a particular instance to one of the models.
I can imagine someone thinking, at this point, âI watch birds because Iâm entranced by their beauty and their behavior or because I enjoy the challenge of finding and identifying as many species as I can. Itâs an aesthetic, emotionally rewarding experience. Doesnât looking at a bird with the cold analytical eye of science ruin the experience?â Of course, I canât speak for everyone, but for me, birding certainly has those rewards, and the more I know, the more my appreciation is enhanced. As many as I have seen, I still am overwhelmed by a peacockâs beauty, but it also spurs me to ask why and how it came to be, and having an answer enlarges and makes whole my experience. We integrate intellectual and aesthetic appreciation when we want to know the names of the birds we encounter and to which family or group a species belongs.
With knowledge of their biology, the most common, everyday birds take on new interest. Take the ubiquitous House Sparrow (Passer domesticus).9 When I stop to look at a House Sparrow, I sometimes think of its broader evolutionary context: other species in the genus Passer. For example, the Italian Sparrow (Passer italiae) originated as a hybrid between House and Spanish Sparrows (Passer hispaniolensis) (see chapter 10), and the Eurasian Tree Sparrow (Passer montanus) replaces the House Sparrow as a human associate in southeastern Asia. The House Sparrow itself shows interesting geographical variation in Europe: northern birds are bigger than birds in the south. This is one of many species of birds and mammals that have this pattern due to adaptive evolution: larger bodies lose heat more slowly than smaller ones and are advantageous in colder regions. What is more, since House Sparrows were introduced from Europe into North America in 1851, they have spread widely, and northern populations have evolved larger size. This was one of the first examples of how rapid evolution can be; Darwin never imagined that evolutionary changes could happen within a few human lifetimes.
The Superb Fairywren (Malurus cyaneus) in Australia (plate 2) is another example of a common bird that poses interesting questions. A group usually has two or more bright blue and black males and several brown birds that include both males and females. Biologist Andrew Cockburn and his associates studied the extraordinary breeding behavior of fairywrens for more than twenty-five years.10 The bright-plumaged and brown males all cooperate to rear nestlings. Cooperative breeding is known in many birds, and why it has evolved poses a very interesting question (chapter 7). But there is more: female fairywrens, to a greater extent than any other bird yet known, engage in âextra-pair copulation,â or adultery: they will travel across intervening territories to mate with a âhotshotâ male. The femaleâs male associates dutifully help raise babies that usually arenât their own offspring. Why are females so unfaithful, and why do males stay and rear the offspring?
These are fascinating questions that evolutionary biology can help to answerâas it can shed light on countless other aspects of birds, ranging from their coloration and structure to their geographic distribution and diversity. My aim in this book is to pose such questions and show how insights from evolutionary biology can answer them. Also, research into these topics has revealed features of many species that I think will amaze and delight anyone who likes birds and help them appreciate birds all the more. And if some readers learn more about evolution and how it is studied, the book will have served another purposeâsharing some of the richness of evolutionary science that I have found so rewarding.
By âevolution,â biologists usually mean change in the features of a single species over time (that is, across generations) as well as the division of a single species into two or more descendant species, both of which undergo change. The alterations of a feature must be inherited to count as evolutionary change. Some features can be affected by an individualâs environment, but these changes are generally not inherited. A generation of people might be lighter skinned than their grandparents because they work in offices instead of fields and so are less suntanned, but this doesnât count as evolution. As inheritance is a defining feature of evolution, evolutionary change of organismsâ features (their phenotype) is accompanied by evolution at the level of the genes. There is also evolution at the genetic (DNA) level that may not affect any features of the organism.
In The Origin of Species, Darwin developed two main themes: that all living things have descended, with modification, from common ancestors; and that the chief cause of modification is natural selection of inherited variations. The wealth of insights, hypotheses, and information in Darwinâs writings is staggering. Every time I read a few pages of The Origin of Species, Iâm simply floored by the questions he thought to ask, the possible answers he advanced, and the evidence he found in an extraordinary range of facts, some of them seemingly trivial. During his voyage on the Beagle, he notices, in South America, that a flycatcher, the Great Kiskadee (Pitangus sulphuratus) (figure 1.1), sometimes acts like kestrels and kingfishers when foraging. Later he cites this, in The Origin of Species, to illustrate that species might change and perhaps become adapted to new ways of life. Not everyone can see a world in a grain of sand, but Darwin realized that a coherent explanation or theory must be able to accommodate, and build on, every fact, however trivial it might seem.
Evolutionary biology today is devoted to Darwinâs two great themes: what has happened in the evolution of the worldâs organisms, and what have been the causes of these evolutionary events?
In studying the history of evolutio...