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Coloration evidence for natural selection

From Wikipedia, the free encyclopedia

Natural selection has driven the ptarmigan to change from snow camouflage in winter to disruptive coloration suiting moorland in summer.
Selective breeding transformed teosinte's small spikes (left) into modern maize (right). Darwin argued that evolution worked in a similar way.

Animal coloration provided important early evidence for evolution by natural selection, at a time when little direct evidence was available. Three major functions of coloration were discovered in the second half of the 19th century, and subsequently used as evidence of selection: camouflage (protective coloration); mimicry, both Batesian and Müllerian; and aposematism.

Charles Darwin's On the Origin of Species was published in 1859, arguing from circumstantial evidence that selection by human breeders could produce change, and that since there was clearly a struggle for existence, that natural selection must be taking place. But he lacked an explanation either for genetic variation or for heredity, both essential to the theory. Many alternative theories were accordingly considered by biologists, threatening to undermine Darwinian evolution.

Some of the first evidence was provided by Darwin's contemporaries, the naturalists Henry Walter Bates and Fritz Müller. They described forms of mimicry that now carry their names, based on their observations of tropical butterflies. These highly specific patterns of coloration are readily explained by natural selection, since predators such as birds which hunt by sight will more often catch and kill insects that are less good mimics of distasteful models than those that are better mimics, but the patterns are otherwise hard to explain.

Darwinists such as Alfred Russel Wallace and Edward Bagnall Poulton, and in the 20th century Hugh Cott and Bernard Kettlewell, sought evidence that natural selection was taking place. Wallace noted that snow camouflage, especially plumage and pelage that changed with the seasons, suggested an obvious explanation as an adaptation for concealment. Poulton's 1890 book, The Colours of Animals, written during Darwinism's lowest ebb, used all the forms of coloration to argue the case for natural selection. Cott described many kinds of camouflage, and in particular his drawings of coincident disruptive coloration in frogs convinced other biologists that these deceptive markings were products of natural selection. Kettlewell experimented on peppered moth evolution, showing that the species had adapted as pollution changed the environment; this provided compelling evidence of Darwinian evolution.

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Transcription

Hi, I'm Hank. And I'm a human, but let's pretend for a moment that I'm a moth. And not just any moth, a peppered moth. Now let's pretend that I'm living in London in the early 1800s, right as the industrial revolution is starting. Life's swell. My light-colored body lets me blend in with the light-colored lichens and tree bark, which means birds have a hard time seeing me, which means I get to live. But it's getting noticeably darker around here with all these coal-powered factories spewing soot into the air, and suddenly all the trees have gone from looking like this to looking like this. So thanks to the soot-covered everything, I've got problems. But you know who doesn't have problems? My brother. He looks like this Yeah, he has a different form of the gene that affects pigmentation. Moths like him represent about 2 percent of all the peppered moths at the start of the industrial revolution. But by 1895 it'll be 95 percent! Why? Well, you're probably already guessing, as the environment gets dirtier, darker moths will be eaten less often, and therefore have more opportunities to make baby moths. The white ones will get eaten more, so over time, the black-colored trait will become more common. As for me? [Eaten.] This, my friends, is a wonderful example of natural selection. The process by which certain inherited traits make it easier for some individuals to thrive and multiply, changing the genetic makeup of populations over time. For this revelation, which remains one of the most important revelations in biology, we have to thank Charles Darwin, who first identified this process in his revolutionary 1859 book, On the Origin of Species by Natural Selection. Now lots of factors play a role in how species change over time including mutation, migration and random changes in how frequently some alleles show up, a process known as genetic drift. But natural selection is the most powerful and most important cause of evolutionary change, which is why today we're going to talk about the principles behind it, and the different ways in which it works. Darwin came to understand the process of selection because he spent his adult life, even most of his childhood, obsessed with observing nature. He studied barnacles, earthworms, birds, rocks, tortoises, fossils, fish, insects and to some extent, even his own family. I'll get back to that in a bit. But it was during Darwin's famous voyage on the H.M.S. Beagle in the 1830s, a surveying expedition around the world, that he began to formulate this theory. Darwin was able to study all kinds of organisms, and he kept amazing journals. Looking back on his notes, he hit upon a couple of particularly important factors in species' survival. One of them was the many examples of adaptations he noticed on his journey. The ways in which organisms seemed to be nearly ideally shaped to enhance their survival and reproduction in specific environments. Maybe the most famous example of these were the variations of beaks Darwin observed among the finches in the remote Galapagos Islands off the coast of South America. He observed more than a dozen closely-related finch species, all of which were quite similar to mainland finch species, but each island species had different shaped and sized beaks that were adapted to the food available specifically on each island. If there were hard seeds, the beaks were thick. If there were insects, the beaks were skinny and pointed. If there were cactus fruit, the beaks were sharp to puncture the fruit's skin. These superior inherited traits led Darwin to another idea, the finches' increased fitness for their environment, that is, their relative ability to survive and create offspring. Explaining the effects of adaptation and relative fitness would become central to Darwin's idea of natural selection. And today we often define natural selection, and describe how it drives evolutionary change, by four basic principles, based on Darwin's observations. The first principle is that different members a population have all kinds of individual variations. These characteristics, whether their body size, hair color, blood type, facial markings, metabolisms or reflexes, are called phenotypes. The second is that many variations are heritable and can be passed on to offspring. If a trait happens to be favorable, it does future generations no good if it can't be passed on. Third: this one tends to get glossed over a lot, even though it's probably the most interesting, is Darwin's observation that populations can often have way more offspring than resources, like food and water, can support. This leads to what Darwin called "the struggle for existence." He was inspired here by the work of economist Thomas Malthus, who wrote that when human populations get too big, we get things like plague and famine and wars, and then only some of us survive and continue to reproduce. If you missed the SciShow Infusion that we did on human overpopulation today and Malthus's predictions, you should check it out now. This finally leads to the last principle of natural selection, which is that, given all of this competition for resources, heritable traits that affect individuals' fitness can lead to variations in their survival and reproductive rates. This is just another way of saying that those with favorable traits are more likely to come out on top and will be more successful with their baby-making. So to wrap all these principles together, in order for natural selection to take place, a population has to have variations, some of which are heritable, and when a variation makes an organism more competitive, that variation will tend to be selected. Like with the peppered moth. It survived because there was variation within the species, the dark coloration, which was heritable, and in turn allowed every moth that inherited that trait to better survive the hungry birds of London. But notice how this works. A single variation in a single organism is only the very beginning of the process. The key is that individuals don't evolve. Instead, natural selection produces evolutionary change because it changes the genetic composition of entire populations, and that occurs through interactions between individuals and their environment. Let's get back to Darwin for a minute. In 1870, Darwin wrote to his neighbor and parliamentarian John Lubbock requesting that a question be added to England's census regarding the frequency of cousins marrying and the health of their offspring. His request was denied, but the question was something that weighed heavily on Darwin's mind, because he was married to Emma Wedgwood, who happened to be his first cousin. Her grandfather was Josiah Wedgwood, founder for the company that remains famous for its pottery and china. Oh, and he was also Darwin's grandfather. In fact, much of Darwin's family tree was...complicated. His marriage to Emma was far from the first Wedgewood-Darwin pairing. Darwin's maternal grandparents and mother were also Wedgwoods, and there were several other marriages between cousins in the family, though not always between those two families. So Darwin, and to a greater extent his children, carried more genetic material of Wedgwood origin than Darwininan. And this caused some problems, the likes of which Darwin was all too aware of, thanks to his own scientific research. Darwin of course spent time studying the effects of crossbreeding and inbreeding in plants and animals, noting that consanguineous pairs often resulted in weaker and sickly descendants. And the same was true of his family. Emma and Charles had 10 children, three of whom died in childhood from infectious disease, which is more likely to be contracted by those with high levels of inbreeding. And while none of Darwin's seven other children had any deformities, he noted that they were "not very robust" and three of them were unable to have children of their own, likely another effect of inbreeding. Now, so far we've been talking about natural selection in terms of physical characteristics, like beak shape or coloration. But it's important to understand that it's not just organism's physical form, or its phenotype, that's changing but its essential genetic form, or genotype. The heritable variations we've been talking about are a function of the alleles that organisms are carrying around. And as organisms become more successful, evolutionarily speaking, by surviving in larger numbers for longer and having more kids, that means that the alleles that mark their variation become more frequent. But these changes can come about in different ways. To understand how, let's walk through the different modes of selection. The mode we've been talking about for much of this episode is an example of directional selection, which is when a favored trait is at one extreme end of the range of traits, like from short to tall, or white to black, or blind to having super-night-goggle vision. Over time this leads to distinct changes in the frequency of that expressed trait in a population, when a single phenotype is favored. So our peppered moth is an example of a population's trait distribution shifting toward one extreme, almost all whitish moths, to the other extreme, almost all blackish. Another awesome example is giraffe necks. They've gotten really long over time because there was selection pressure against short necks, which couldn't reach all of those delicious leaves. But there's also stabilizing selection, which selects against extreme phenotypes and instead favors the majority that are well adapted to an environment. An example that's often used is a human's birth weight: Very small babies have a harder time defending themselves from infections and staying warm, but very large babies are too large to deliver naturally. Because of this, the survival rate for babies has historically been higher for those in the middle weight range, which helped stabilize average birth weight. At least, until Cesarian sections became as common as bad tattoos. So what happens when the environment favors extreme traits at both ends of the spectrum, while selecting against the common traits? That's disruptive selection. Now examples of this are rare, but scientists think they found an instance of it in 2008, in a lake full of tiny crustaceans called Daphnia. The population was hit with an epidemic of yeast parasite, and after about a half-dozen generations, a variance had emerged in how the Daphnia responded to the parasite. Some became less susceptible to the yeast, but were smaller and had fewer offspring. The others actually became more susceptible to the parasite, but were bigger and able to reproduce more, at least while they were still alive. So there were two traits that were being selected for, both in extremes and both to the exclusion of each other: susceptibility and fecundity. If you got one, you didn't get the other. An interesting example, of selection being driven by a parasite. Now while these are the main ways that selective pressures can affect populations, those pressures can also come from factors other than environmental ones like food supply or predators or parasites. There's also sexual selection, another concept introduced by Darwin and described in The Origin of Species as depending "not on a struggle for existence, but a struggle between individuals of the same sex, generally the males, for the possession of the other sex." Basically, for individuals to maximize their fitness, they not only need to survive but they also need to reproduce more, and they can do that one or two ways: One, they can make themselves attractive to the opposite sex. Or two, they can go for the upper hand by intimidating, deterring or defeating the same-sex rivals. The first of these strategies is how we ended up with this: I mean, the peacock tail isn't exactly camouflage. But the more impressive the tail, the better chances a male will find a mate and pass its genes to the next generation. Sad-looking peacock tails will diminish over generations, making it a good example of directional sexual selection. The other strategy involves fighting, or at least looking like= you want to fight, for the privilege of mating, which tends to select for bigger or stronger or meaner-looking mates. And finally, thanks to us humans there are also un-natural forms of selection, and we call that artificial selection. People have been artificially selecting plants and animals for thousands of years, and Darwin spent a lot of time in Origin of Species talking about the breeding of pigeons and cattle and plants to demonstrate the principles of selection. We encourage the selection of some traits and discourage others. It's how we got grains that produce all those nutrients. Which is how we managed to turn the gray wolf into domesticated dogs that can look like this or like that, two of my favorite examples of artificial selection. Now these are different breeds of dogs- Oh, where you goin'? No. No. But they're both still dogs. They're the same species. Technically, a corgi and a greyhound could get together and have a baby dog, though it would be a weird looking dog. But, what happens when selection makes populations so different that they can't even be the same species any more? Well, that's what we're going to talk about next episode on Crash Course Biology: how one species can turn into another species. In the meantime, feel free to review what we've gone over today, ask us questions down in the comments below, or on Facebook or Twitter, We'll see you next time. [WOOF]

Context

Charles Darwin published On the Origin of Species in 1859,[1] arguing that evolution in nature must be driven by natural selection, just as breeds of domestic animals and cultivars of crop plants were driven by artificial selection.[2][3] Darwin's theory radically altered popular and scientific opinion about the development of life.[4] However, he lacked evidence and explanations for some critical components of the evolutionary process. He could not explain the source of variation in traits within a species, and did not have a mechanism of heredity that could pass traits faithfully from one generation to the next. This made his theory vulnerable; alternative theories were being explored during the eclipse of Darwinism; and so Darwinian field naturalists like Wallace, Bates and Müller looked for clear evidence that natural selection actually occurred.[5] Animal coloration, readily observable, soon provided strong and independent lines of evidence, from camouflage, mimicry and aposematism, that natural selection was indeed at work.[6][7][8] The historian of science Peter J. Bowler wrote that Darwin's theory "was also extended to the broader topics of protective resemblances and mimicry, and this was its greatest triumph in explaining adaptations".[9]

Camouflage

Snow camouflage

Convergent evolution of snow camouflage in Arctic hare, ermine, and ptarmigan provided early evidence for natural selection.

In his 1889 book Darwinism, the naturalist Alfred Russel Wallace considered the white coloration of Arctic animals. He recorded that the Arctic fox, Arctic hare, ermine and ptarmigan change their colour seasonally, and gave "the obvious explanation", that it was for concealment.[8][a] The modern ornithologist W. L. N. Tickell, reviewing proposed explanations of white plumage in birds, writes that in the ptarmigan "it is difficult to escape the conclusion that cryptic brown summer plumage becomes a liability in snow, and white plumage is therefore another cryptic adaptation." All the same, he notes, "in spite of winter plumage, many Ptarmigan in NE Iceland are killed by Gyrfalcons throughout the winter."[11]

More recently, decreasing snow cover in Poland, caused by global warming, is reflected in a reduced percentage of white-coated weasels that become white in winter. Days with snow cover halved between 1997 and 2007, and as few as 20 percent of the weasels had white winter coats. This was shown to be a result of natural selection by predators making use of camouflage mismatch.[12][13]

Coincident disruptive coloration

Hugh Cott's drawings of 'coincident disruptive coloration' formed "persuasive arguments"[7] for natural selection. Left: active; right: at rest, marks coinciding.

In the words of camouflage researchers Innes Cuthill and A. Székely, the English zoologist and camouflage expert Hugh Cott's 1940 book Adaptive Coloration in Animals provided "persuasive arguments for the survival value of coloration, and for adaptation in general, at a time when natural selection was far from universally accepted within evolutionary biology."[7]

In particular, they argue, "Coincident Disruptive Coloration" (one of Cott's categories) "made Cott's drawings the most compelling evidence for natural selection enhancing survival through disruptive camouflage."[7]

Cott explained, while discussing "a little frog known as Megalixalus fornasinii" in his chapter on coincident disruptive coloration, that "it is only when the pattern is considered in relation to the frog's normal attitude of rest that its remarkable nature becomes apparent... The attitude and very striking colour-scheme thus combine to produce an extraordinary effect, whose deceptive appearance depends upon the breaking up of the entire form into two strongly contrasted areas of brown and white. Considered separately, neither part resembles part of a frog. Together in nature the white configuration alone is conspicuous. This stands out and distracts the observer's attention from the true form and contour of the body and appendages on which it is superimposed".[14]

Cott concluded that the effect was concealment "so long as the false configuration is recognized in preference to the real one".[14] Such patterns embody, as Cott stressed, considerable precision as the markings must line up accurately for the disguise to work. Cott's description and in particular his drawings convinced biologists that the markings, and hence the camouflage, must have survival value (rather than occurring by chance); and further, as Cuthill and Székely indicate, that the bodies of animals that have such patterns must indeed have been shaped by natural selection.[7]

Bernard Kettlewell claimed that changes in the frequencies of light and dark morphs of the peppered moth, Biston betularia were direct evidence of natural selection.

Industrial melanism

Between 1953 and 1956, the geneticist Bernard Kettlewell experimented on peppered moth evolution. He presented results showing that in a polluted urban wood with dark tree trunks, dark moths survived better than pale ones, causing industrial melanism, whereas in a clean rural wood with paler trunks, pale moths survived better than dark ones. The implication was that survival was caused by camouflage against suitable backgrounds, where predators hunting by sight (insect-eating birds, such as the great tits used in the experiment) selectively caught and killed the less well-camouflaged moths. The results were intensely controversial, and from 2001 Michael Majerus carefully repeated the experiment. The results were published posthumously in 2012, vindicating Kettlewell's work as "the most direct evidence", and "one of the clearest and most easily understood examples of Darwinian evolution in action".[15]

Mimicry

The butterfly Papilio polytes (left) mimics the unpalatable Pachliopta aristolochiae (right).

Batesian

Batesian mimicry, named for the 19th century naturalist Henry Walter Bates who first noted the effect in 1861, "provides numerous excellent examples of natural selection"[16] at work. The evolutionary entomologist James Mallet noted that mimicry was "arguably the oldest Darwinian theory not attributable to Darwin."[6] Inspired by On the Origin of Species, Bates realized that unrelated Amazonian butterflies resembled each other when they lived in the same areas, but had different coloration in different locations in the Amazon, something that could only have been caused by adaptation.[6]

Müllerian

Müllerian mimicry, too, in which two or more distasteful species that share one or more predators have come to mimic each other's warning signals, was clearly adaptive; Fritz Müller described the effect in 1879, in an account notable for being the first use of a mathematical argument in evolutionary ecology to show how powerful the effect of natural selection would be.[b][6]

Warning coloration protects poison dart frog Dendrobates leucomelas.

Aposematism

In 1867, in a letter to Darwin, Wallace described warning coloration. The evolutionary zoologist James Mallet notes that this discovery "rather illogically"[6] followed rather than preceded the accounts of Batesian and Müllerian mimicry, which both rely on the existence and effectiveness of warning coloration.[6][c] The conspicuous colours and patterns of animals with strong defences such as toxins are advertised to predators, signalling honestly that the animal is not worth attacking. This directly increases the reproductive fitness of the potential prey, providing a strong selective advantage. The existence of unequivocal warning coloration is therefore clear evidence of natural selection at work.[17]

Defence of Darwinism

Warning coloration of the "Brazilian Skunk" in Edward Bagnall Poulton's The Colours of Animals, 1890

Edward Bagnall Poulton's 1890 book, The Colours of Animals, renamed Wallace's concept of warning colours "aposematic" coloration, as well as supporting Darwin's then unpopular theories of natural selection and sexual selection.[18] Poulton's explanations of coloration are emphatically Darwinian. For example, on aposematic coloration he wrote that

At first sight the existence of this group seems to be a difficulty in the way of the general applicability of the theory of natural selection. Warning Colours appear to benefit the would-be enemies rather than the conspicuous forms themselves, and the origin and growth of a character intended solely for the advantage of some other species cannot be explained by the theory of natural selection. But the conspicuous animal is greatly benefited by its Warning Colours. If it resembled its surroundings like the members of the other class, it would be liable to a great deal of accidental or experimental tasting, and there would be nothing about it to impress the memory of an enemy, and thus to prevent the continual destruction of individuals. The object of Warning Colours is to assist the education of enemies, enabling them to easily learn and remember the animals which are to be avoided. The great advantage conferred upon the conspicuous species is obvious when it is remembered that such an easy and successful education means an education involving only a small sacrifice of life."[19]

Poulton summed up his allegiance to Darwinism as an explanation of Batesian mimicry in one sentence: "Every step in the gradually increasing change of the mimicking in the direction of specially protected form, would have been an advantage in the struggle for existence".[19]

The historian of biology Peter J. Bowler commented that Poulton used his book to complain about experimentalists' lack of attention to what field naturalists (like Wallace, Bates, and Poulton) could readily see were adaptive features. Bowler added that "The fact that the adaptive significance of coloration was (sic) widely challenged indicates just how far anti-Darwinian feeling had developed.[d] Only field naturalists such as Poulton refused to give in, convinced that their observations showed the validity of selection, whatever the theoretical problems."[20]

Notes

  1. ^ In the case of prey animals like the Arctic hare, this is concealment from predators, reducing the chance of being seen and eaten, directly affecting survival. In the case of predators like the Arctic fox, camouflage provides concealment from prey, increasing the change of a successful hunt, and again improving survival chances. Thus whether the camouflage is defensive or aggressive, it would be favoured by natural selection. Edward Bagnall Poulton named the two cases "general protective resemblance" and "general aggressive resemblance" respectively.[10]
  2. ^ If one may ignore, as Mallet comments, Malthus's discussion of the effects of population growth that influenced Darwin.[6]
  3. ^ Edward Bagnall Poulton invented the name aposematism some years later, in 1890, in his book The Colours of Animals.[10]
  4. ^ In the eclipse of Darwinism.

References

  1. ^ Darwin, Charles (1859). On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (1st ed.). John Murray. LCCN 06017473. OCLC 741260650. The book is available from The Complete Work of Charles Darwin Online.
  2. ^ Lewontin, Richard C. (November 1970). "The Units of Selection" (PDF). Annual Review of Ecology and Systematics. 1: 1–18. doi:10.1146/annurev.es.01.110170.000245. JSTOR 2096764.
  3. ^ Hall, Brian K.; Hallgrímsson, Benedikt (2008). Strickberger's Evolution (4th ed.). Jones and Bartlett. pp. 4–6. ISBN 978-0-7637-0066-9. OCLC 796450355.
  4. ^ Bowler, Peter J. (2003). Evolution: The History of an Idea (3rd ed.). University of California Press. pp. 177–223. ISBN 978-0-520-23693-6.
  5. ^ Larson, Edward J. (2004). Evolution: The Remarkable History of a Scientific Theory. New York: Modern Library. pp. 121–123, 152–157. ISBN 978-0-679-64288-6.
  6. ^ a b c d e f g Mallet, James (July 2001). "Mimicry: An interface between psychology and evolution". PNAS. 98 (16): 8928–8930. Bibcode:2001PNAS...98.8928M. doi:10.1073/pnas.171326298. PMC 55348. PMID 11481461.
  7. ^ a b c d e Cuthill, I. C.; Székely, A. (2011). Stevens, Martin; Merilaita, Sami (eds.). Animal Camouflage: Mechanisms and Function. Cambridge University Press. p. 50. ISBN 978-1-139-49623-0.
  8. ^ a b Wallace, Alfred Russel (2015) [1889]. Darwinism - An Exposition Of The Theory Of Natural Selection - With Some Of Its Applications. Read Books. p. 180. ISBN 978-1-4733-7510-9.
  9. ^ Bowler, Peter J. (1983). The Eclipse of Darwinism: anti-Darwinian evolutionary theories in the decades around 1900. Johns Hopkins University Press. p. 29. ISBN 978-0-8018-4391-4.
  10. ^ a b Poulton, Edward Bagnall (1890). The Colours of Animals. p. Foldout after page 339.
  11. ^ Tickell, W. L. N. (March 2003). "White Plumage". Waterbirds: The International Journal of Waterbird Biology. 26 (1): 1–12. JSTOR 1522461.
  12. ^ Knapton, Sarah (24 May 2018). "White-furred animals could die out because of climate change, study suggests". The Daily Telegraph. Retrieved 24 May 2018.
  13. ^ Atmeh, Kamal; Andruszkiewicz, Anna; Zub, Karol (24 May 2018). "Climate change is affecting mortality of weasels due to camouflage mismatch". Scientific Reports. 8 (1): 7648. Bibcode:2018NatSR...8.7648A. doi:10.1038/s41598-018-26057-5. PMC 5967304. PMID 29795400.
  14. ^ a b Cott, Hugh B. (1940). Adaptive Coloration in Animals. Methuen. pp. 68–72.
  15. ^ Cook, L. M.; Grant, B. S.; Saccheri, I. J.; Mallet, James (2012). "Selective bird predation on the peppered moth: the last experiment of Michael Majerus". Biology Letters. 8 (4): 609–612. doi:10.1098/rsbl.2011.1136. PMC 3391436. PMID 22319093.
  16. ^ Rice, Stanley A. (2009). Encyclopedia of Evolution. Infobase Publishing. p. 274. ISBN 978-1-4381-1005-9.
  17. ^ Sherratt, T. N. (2002). "The coevolution of warning signals". Proceedings of the Royal Society B. 269 (1492): 741–746. doi:10.1098/rspb.2001.1944. PMC 1690947. PMID 11934367.
  18. ^ Mallet, Jim. "E.B. Poulton (1890)". University College London. Retrieved 23 November 2012.
  19. ^ a b Poulton, Edward Bagnall (1890). The Colours of Animals, their meaning and use, especially considered in the case of insects. Kegan Paul, Trench & Trübner. pp. 160–161, 220, and whole book.
  20. ^ Bowler, Peter J. (2003) [1989]. Evolution:The History of an Idea. University of California Press. p. 250. ISBN 978-0-520-23693-6.
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