Essay Nature Selected

Mechanism of evolution by differential survival and reproduction of individuals

For other uses, see natural selection (disambiguation).

Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritabletraits characteristic of a population over generations. Charles Darwin popularised the term "natural selection", contrasting it with artificial selection, which is intentional, whereas natural selection is not.

Variation exists within all populations of organisms. This occurs partly because random mutations arise in the genome of an individual organism, and offspring can inherit such mutations. Throughout the lives of the individuals, their genomes interact with their environments to cause variations in traits. The environment of a genome includes the molecular biology in the cell, other cells, other individuals, populations, species, as well as the abiotic environment. Because individuals with certain variants of the trait tend to survive and reproduce more than individuals with other, less successful, variants, the population evolves. Other factors affecting reproductive success include sexual selection (now often included in natural selection) and fecundity selection.

Natural selection acts on the phenotype, the characteristics of the organism which actually interact with the environment, but the genetic (heritable) basis of any phenotype that gives that phenotype a reproductive advantage may become more common in a population. Over time, this process can result in populations that specialise for particular ecological niches (microevolution) and may eventually result in speciation (the emergence of new species, macroevolution). In other words, natural selection is a key process in the evolution of a population.

Natural selection is a cornerstone of modern biology. The concept, published by Darwin and Alfred Russel Wallace in a joint presentation of papers in 1858, was elaborated in Darwin's influential 1859 book On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. He described natural selection as analogous to artificial selection, a process by which animals and plants with traits considered desirable by human breeders are systematically favoured for reproduction. The concept of natural selection originally developed in the absence of a valid theory of heredity; at the time of Darwin's writing, science had yet to develop modern theories of genetics. The union of traditional Darwinian evolution with subsequent discoveries in classical genetics formed the modern synthesis of the mid-20th century. The addition of molecular genetics has led to evolutionary developmental biology, which explains evolution at the molecular level. While genotypes can slowly change by random genetic drift, natural selection remains the primary explanation for adaptive evolution.

Historical development[edit]

Main article: History of evolutionary thought

Pre-Darwinian theories[edit]

Several philosophers of the classical era, including Empedocles[1] and his intellectual successor, the Roman poet Lucretius,[2] expressed the idea that nature produces a huge variety of creatures, randomly, and that only those creatures that manage to provide for themselves and reproduce successfully persist. Empedocles' idea that organisms arose entirely by the incidental workings of causes such as heat and cold was criticised by Aristotle in Book II of Physics.[3] He posited natural teleology in its place, and believed that form was achieved for a purpose, citing the regularity of heredity in species as proof.[4][5] Nevertheless, he accepted in his biology that new types of animals, monstrosities (τερας), can occur in very rare instances (Generation of Animals, Book IV).[6] As quoted in Darwin's 1872 edition of The Origin of Species, Aristotle considered whether different forms (e.g., of teeth) might have appeared accidentally, but only the useful forms survived:

So what hinders the different parts [of the body] from having this merely accidental relation in nature? as the teeth, for example, grow by necessity, the front ones sharp, adapted for dividing, and the grinders flat, and serviceable for masticating the food; since they were not made for the sake of this, but it was the result of accident. And in like manner as to the other parts in which there appears to exist an adaptation to an end. Wheresoever, therefore, all things together (that is all the parts of one whole) happened like as if they were made for the sake of something, these were preserved, having been appropriately constituted by an internal spontaneity, and whatsoever things were not thus constituted, perished, and still perish.

— Aristotle, Physics, Book II, Chapter 8[7]

But Aristotle rejected this possibility in the next paragraph, making clear that he is talking about the development of animals as embryos with the phrase "either invariably or normally come about", not the origin of species:

... Yet it is impossible that this should be the true view. For teeth and all other natural things either invariably or normally come about in a given way; but of not one of the results of chance or spontaneity is this true. We do not ascribe to chance or mere coincidence the frequency of rain in winter, but frequent rain in summer we do; nor heat in the dog-days, but only if we have it in winter. If then, it is agreed that things are either the result of coincidence or for an end, and these cannot be the result of coincidence or spontaneity, it follows that they must be for an end; and that such things are all due to nature even the champions of the theory which is before us would agree. Therefore action for an end is present in things which come to be and are by nature.

— Aristotle, Physics, Book II, Chapter 8[8]

The struggle for existence was later described by the Islamic writer Al-Jahiz in the 9th century.[9][10][11]

The classical arguments were reintroduced in the 18th century by Pierre Louis Maupertuis[12] and others, including Darwin's grandfather, Erasmus Darwin.

Until the early 19th century, the prevailing view in Western societies was that differences between individuals of a species were uninteresting departures from their Platonic ideals (or typus) of created kinds. However, the theory of uniformitarianism in geology promoted the idea that simple, weak forces could act continuously over long periods of time to produce radical changes in the Earth's landscape. The success of this theory raised awareness of the vast scale of geological time and made plausible the idea that tiny, virtually imperceptible changes in successive generations could produce consequences on the scale of differences between species.[13]

The early 19th-century zoologist Jean-Baptiste Lamarck suggested the inheritance of acquired characteristics as a mechanism for evolutionary change; adaptive traits acquired by an organism during its lifetime could be inherited by that organism's progeny, eventually causing transmutation of species.[14] This theory, Lamarckism, was an influence on the Soviet biologist Trofim Lysenko's antagonism to mainstream genetic theory as late as the mid 20th century.[15]

Between 1835 and 1837, the zoologist Edward Blyth worked on the area of variation, artificial selection, and how a similar process occurs in nature. Darwin acknowledged Blyth's ideas in the first chapter on variation of On the Origin of Species.[16]

Darwin's theory[edit]

Main articles: Inception of Darwin's theory and Development of Darwin's theory

Further information: Coloration evidence for natural selection

In 1859, Charles Darwin set out his theory of evolution by natural selection as an explanation for adaptation and speciation. He defined natural selection as the "principle by which each slight variation [of a trait], if useful, is preserved".[17] The concept was simple but powerful: individuals best adapted to their environments are more likely to survive and reproduce. As long as there is some variation between them and that variation is heritable, there will be an inevitable selection of individuals with the most advantageous variations. If the variations are heritable, then differential reproductive success leads to a progressive evolution of particular populations of a species, and populations that evolve to be sufficiently different eventually become different species.[18][19]

Darwin's ideas were inspired by the observations that he had made on the second voyage of HMS Beagle (1831–1836), and by the work of a political economist, Thomas Robert Malthus, who, in An Essay on the Principle of Population (1798), noted that population (if unchecked) increases exponentially, whereas the food supply grows only arithmetically; thus, inevitable limitations of resources would have demographic implications, leading to a "struggle for existence".[20] When Darwin read Malthus in 1838 he was already primed by his work as a naturalist to appreciate the "struggle for existence" in nature. It struck him that as population outgrew resources, "favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species."[21] Darwin wrote:

If during the long course of ages and under varying conditions of life, organic beings vary at all in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the high geometrical powers of increase of each species, at some age, season, or year, a severe struggle for life, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of all organic beings to each other and to their conditions of existence, causing an infinite diversity in structure, constitution, and habits, to be advantageous to them, I think it would be a most extraordinary fact if no variation ever had occurred useful to each being's own welfare, in the same way as so many variations have occurred useful to man. But if variations useful to any organic being do occur, assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life; and from the strong principle of inheritance they will tend to produce offspring similarly characterised. This principle of preservation, I have called, for the sake of brevity, Natural Selection.

— Darwin summarising natural selection in the fourth chapter of On the Origin of Species[22]

Once he had his theory, Darwin was meticulous about gathering and refining evidence before making his idea public. He was in the process of writing his "big book" to present his research when the naturalist Alfred Russel Wallace independently conceived of the principle and described it in an essay he sent to Darwin to forward to Charles Lyell. Lyell and Joseph Dalton Hooker decided to present his essay together with unpublished writings that Darwin had sent to fellow naturalists, and On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection was read to the Linnean Society of London announcing co-discovery of the principle in July 1858.[23] Darwin published a detailed account of his evidence and conclusions in On the Origin of Species in 1859. In the 3rd edition of 1861 Darwin acknowledged that others—like William Charles Wells in 1813, and Patrick Matthew in 1831—had proposed similar ideas, but had neither developed them nor presented them in notable scientific publications.[24]

Darwin thought of natural selection by analogy to how farmers select crops or livestock for breeding, which he called "artificial selection"; in his early manuscripts he referred to a "Nature" which would do the selection. At the time, other mechanisms of evolution such as evolution by genetic drift were not yet explicitly formulated, and Darwin believed that selection was likely only part of the story: "I am convinced that Natural Selection has been the main but not exclusive means of modification."[25] In a letter to Charles Lyell in September 1860, Darwin regretted the use of the term "Natural Selection", preferring the term "Natural Preservation".[26]

For Darwin and his contemporaries, natural selection was in essence synonymous with evolution by natural selection. After the publication of On the Origin of Species,[27] educated people generally accepted that evolution had occurred in some form. However, natural selection remained controversial as a mechanism, partly because it was perceived to be too weak to explain the range of observed characteristics of living organisms, and partly because even supporters of evolution balked at its "unguided" and non-progressive nature,[28] a response that has been characterised as the single most significant impediment to the idea's acceptance.[29] However, some thinkers enthusiastically embraced natural selection; after reading Darwin, Herbert Spencer introduced the phrase survival of the fittest, which became a popular summary of the theory.[30][31] The fifth edition of On the Origin of Species published in 1869 included Spencer's phrase as an alternative to natural selection, with credit given: "But the expression often used by Mr. Herbert Spencer of the Survival of the Fittest is more accurate, and is sometimes equally convenient."[32] Although the phrase is still often used by non-biologists, modern biologists avoid it because it is tautological if "fittest" is read to mean "functionally superior" and is applied to individuals rather than considered as an averaged quantity over populations.[33]

The modern synthesis[edit]

Main article: Modern synthesis (20th century)

Natural selection relies crucially on the idea of heredity, but developed before the basic concepts of genetics. Although the Moravian monk Gregor Mendel, the father of modern genetics, was a contemporary of Darwin's, his work lay in obscurity, only being rediscovered in 1900.[34] With the early 20th century integration of evolution with Mendel's laws of inheritance, the so-called modern synthesis, scientists generally came to accept natural selection.[35][36] The synthesis grew from advances in different fields. Ronald Fisher developed the required mathematical language and wrote The Genetical Theory of Natural Selection (1930).[37]J. B. S. Haldane introduced the concept of the "cost" of natural selection.[38][39]Sewall Wright elucidated the nature of selection and adaptation.[40] In his book Genetics and the Origin of Species (1937), Theodosius Dobzhansky established the idea that mutation, once seen as a rival to selection, actually supplied the raw material for natural selection by creating genetic diversity.[41][42]

A second synthesis[edit]

Main article: Evolutionary developmental biology § History

Ernst Mayr recognised the key importance of reproductive isolation for speciation in his Systematics and the Origin of Species (1942).[44]W. D. Hamilton conceived of kin selection in 1964.[45][46] This synthesis cemented natural selection as the foundation of evolutionary theory, where it remains today. A second synthesis was brought about at the end of the 20th century by advances in molecular genetics, creating the field of evolutionary developmental biology ("evo-devo"), which seeks to explain the evolution of form in terms of the genetic regulatory programs which control the development of the embryo at molecular level. Natural selection is here understood to act on embryonic development to change the morphology of the adult body.[47][48][49][50]

Terminology[edit]

The term natural selection is most often defined to operate on heritable traits, because these directly participate in evolution. However, natural selection is "blind" in the sense that changes in phenotype can give a reproductive advantage regardless of whether or not the trait is heritable. Following Darwin's primary usage, the term is used to refer both to the evolutionary consequence of blind selection and to its mechanisms.[27][37][51][52] It is sometimes helpful to explicitly distinguish between selection's mechanisms and its effects; when this distinction is important, scientists define "(phenotypic) natural selection" specifically as "those mechanisms that contribute to the selection of individuals that reproduce", without regard to whether the basis of the selection is heritable.[53][54][55] Traits that cause greater reproductive success of an organism are said to be selected for, while those that reduce success are selected against.[56]

Mechanism[edit]

Heritable variation, differential reproduction[edit]

Main article: Genetic variation

Natural variation occurs among the individuals of any population of organisms. Some differences may improve an individual's chances of surviving and reproducing such that its lifetime reproductive rate is increased, which means that it leaves more offspring. If the traits that give these individuals a reproductive advantage are also heritable, that is, passed from parent to offspring, then there will be differential reproduction, that is, a slightly higher proportion of fast rabbits or efficient algae in the next generation. Even if the reproductive advantage is very slight, over many generations any advantageous heritable trait becomes dominant in the population. In this way the natural environment of an organism "selects for" traits that confer a reproductive advantage, causing evolutionary change, as Darwin described.[57] This gives the appearance of purpose, but in natural selection there is no intentional choice. Artificial selection is purposive where natural selection is not, though biologists often use teleological language to describe it.[58]

The peppered moth exists in both light and dark colours in Great Britain, but during the industrial revolution, many of the trees on which the moths rested became blackened by soot, giving the dark-coloured moths an advantage in hiding from predators. This gave dark-coloured moths a better chance of surviving to produce dark-coloured offspring, and in just fifty years from the first dark moth being caught, nearly all of the moths in industrial Manchester were dark. The balance was reversed by the effect of the Clean Air Act 1956, and the dark moths became rare again, demonstrating the influence of natural selection on peppered moth evolution.[59]

Fitness[edit]

Main article: Fitness (biology)

The concept of fitness is central to natural selection. In broad terms, individuals that are more "fit" have better potential for survival, as in the well-known phrase "survival of the fittest", but the precise meaning of the term is much more subtle. Modern evolutionary theory defines fitness not by how long an organism lives, but by how successful it is at reproducing. If an organism lives half as long as others of its species, but has twice as many offspring surviving to adulthood, its genes become more common in the adult population of the next generation. Though natural selection acts on individuals, the effects of chance mean that fitness can only really be defined "on average" for the individuals within a population. The fitness of a particular genotype corresponds to the average effect on all individuals with that genotype.[60]

Competition[edit]

Main article: Competition (biology)

In biology, competition is an interaction between organisms in which the fitness of one is lowered by the presence of another. This may be because both rely on a limited supply of a resource such as food, water, or territory.[61] Competition may be within or between species, and may be direct or indirect.[62] Species less suited to compete should in theory either adapt or die out, since competition plays a powerful role in natural selection, but according to the "room to roam" theory it may be less important than expansion among larger clades.[62][63]

Competition is modelled by r/K selection theory, which is based on Robert MacArthur and E. O. Wilson's work on island biogeography.[64] In this theory, selective pressures drive evolution in one of two stereotyped directions: r- or K-selection.[65] These terms, r and K, can be illustrated in a logistic model of population dynamics:[66]

where r is the growth rate of the population (N), and K is the carrying capacity of its local environmental setting. Typically, r-selected species exploit empty niches, and produce many offspring, each with a relatively low probability of surviving to adulthood. In contrast, K-selected species are strong competitors in crowded niches, and invest more heavily in much fewer offspring, each with a relatively high probability of surviving to adulthood.[66]

Types of selection[edit]

Natural selection can act on any heritable phenotypic trait,[67] and selective pressure can be produced by any aspect of the environment, including sexual selection and competition with members of the same or other species.[68][69] However, this does not imply that natural selection is always directional and results in adaptive evolution; natural selection often results in the maintenance of the status quo by eliminating less fit variants.[57]

Selection can be classified in several different ways, such as by its effect on a trait, on genetic diversity, by the life cycle stage where it acts, by the unit of selection, or by the resource being competed for.

Selection has different effects on traits. Stabilizing selection acts to hold a trait at a stable optimum, and in the simplest case all deviations from this optimum are selectively disadvantageous. Directional selection favours extreme values of a trait. The uncommon disruptive selection also acts during transition periods when the current mode is sub-optimal, but alters the trait in more than one direction. In particular, if the trait is quantitative and univariate then both higher and lower trait levels are favoured. Disruptive selection can be a precursor to speciation.[57]

Alternatively, selection can be divided according to its effect on genetic diversity. Purifying or negative selection acts to remove genetic variation from the population (and is opposed by de novo mutation, which introduces new variation.[70][71] In contrast, balancing selection acts to maintain genetic variation in a population, even in the absence of de novo mutation, by negative frequency-dependent selection. One mechanism for this is heterozygote advantage, where individuals with two different alleles have a selective advantage over individuals with just one allele. The polymorphism at the human ABO blood group locus has been explained in this way.[72]

Another option is to classify selection by the life cycle stage at which it acts. Some biologists recognise just two types: viability (or survival) selection, which acts to increase an organism's probability of survival, and fecundity (or fertility or reproductive) selection, which acts to increase the rate of reproduction, given survival. Others split the life cycle into further components of selection. Thus viability and survival selection may be defined separately and respectively as acting to improve the probability of survival before and after reproductive age is reached, while fecundity selection may be split into additional sub-components including sexual selection, gametic selection, acting on gamete survival, and compatibility selection, acting on zygote formation.[73]

Selection can also be classified by the level or unit of selection. Individual selection acts on the individual, in the sense that adaptations are "for" the benefit of the individual, and result from selection among individuals. Gene selection acts directly at the level of the gene. In kin selection and intragenomic conflict, gene-level selection provides a more apt explanation of the underlying process. Group selection, if it occurs, acts on groups of organisms, on the assumption that groups replicate and mutate in an analogous way to genes and individuals. There is an ongoing debate over the degree to which group selection occurs in nature.[74]

Finally, selection can be classified according to the resource being competed for. Sexual selection results from competition for mates. Sexual selection typically proceeds via fecundity selection, sometimes at the expense of viability. Ecological selection is natural selection via any means other than sexual selection, such as kin selection, competition, and infanticide. Following Darwin, natural selection is sometimes defined as ecological selection, in which case sexual selection is considered a separate mechanism.[75]

Sexual selection[edit]

Main article: Sexual selection

Sexual selection as first articulated by Darwin (using the example of the peacock's tail)[76] refers specifically to competition for mates,[78] which can be intrasexual, between individuals of the same sex, that is male–male competition, or intersexual, where one gender chooses mates, most often with males displaying and females choosing.[79] However, in some species, mate choice is primarily by males, as in some fishes of the family Syngnathidae.[80][81]

Phenotypic traits can be displayed in one sex and desired in the other sex, causing a positive feedback loop called a Fisherian runaway, for example, the extravagant plumage of some male birds such as the peacock.[77] An alternate theory proposed by the same Ronald Fisher in 1930 is the sexy son hypothesis, that mothers want promiscuous sons to give them large numbers of grandchildren and so choose promiscuous fathers for their children. Aggression between members of the same sex is sometimes associated with very distinctive features, such as the antlers of stags, which are used in combat with other stags. More generally, intrasexual selection is often associated with sexual dimorphism, including differences in body size between males and females of a species.[79]

Natural selection in action[edit]

Further information: Antimicrobial resistance

Natural selection is seen in action in the development of antibiotic resistance in microorganisms. Since the discovery of penicillin in 1928, antibiotics have been used to fight bacterial diseases. The widespread misuse of antibiotics has selected for microbial resistance to antibiotics in clinical use, to the point that the methicillin-resistant Staphylococcus aureus (MRSA) has been described as a "superbug" because of the threat it poses to health and its relative invulnerability to existing drugs.[82] Response strategies typically include the use of different, stronger antibiotics; however, new strains of MRSA have recently emerged that are resistant even to these drugs.[83] This is an evolutionary arms race, in which bacteria develop strains less susceptible to antibiotics, while medical researchers attempt to develop new antibiotics that can kill them. A similar situation occurs with pesticide resistance in plants and insects. Arms races are not necessarily induced by man; a well-documented example involves the spread of a gene in the butterfly Hypolimnas bolina suppressing male-killing activity by Wolbachia bacteria parasites on the island of Samoa, where the spread of the gene is known to have occurred over a period of just five years[84][85]

Evolution by means of natural selection[edit]

Main articles: Evolution and Darwinism

A prerequisite for natural selection to result in adaptive evolution, novel traits and speciation is the presence of heritable genetic variation that results in fitness differences. Genetic variation is the result of mutations, genetic recombinations and alterations in the karyotype (the number, shape, size and internal arrangement of the chromosomes). Any of these changes might have an effect that is highly advantageous or highly disadvantageous, but large effects are rare. In the past, most changes in the genetic material were considered neutral or close to neutral because they occurred in noncoding DNA or resulted in a synonymous substitution. However, many mutations in non-coding DNA have deleterious effects.[86][87] Although both mutation rates and average fitness effects of mutations are dependent on the organism, a majority of mutations in humans are slightly deleterious.[88]

Some mutations occur in "toolkit" or regulatory genes. Changes in these often have large effects on the phenotype of the individual because they regulate the function of many other genes. Most, but not all, mutations in regulatory genes result in non-viable embryos. Some nonlethal regulatory mutations occur in HOX genes in humans, which can result in a cervical rib[89] or polydactyly, an increase in the number of fingers or toes.[90] When such mutations result in a higher fitness, natural selection favours these phenotypes and the novel trait spreads in the population. Established traits are not immutable; traits that have high fitness in one environmental context may be much less fit if environmental conditions change. In the absence of natural selection to preserve such a trait, it becomes more variable and deteriorate over time, possibly resulting in a vestigial manifestation of the trait, also called evolutionary baggage. In many circumstances, the apparently vestigial structure may retain a limited functionality, or may be co-opted for other advantageous traits in a phenomenon known as preadaptation. A famous example of a vestigial structure, the eye of the blind mole-rat, is believed to retain function in photoperiod perception.[91]

Speciation[edit]

Main article: Speciation

Speciation requires a degree of reproductive isolation—that is, a reduction in gene flow. However, it is intrinsic to the concept of a species that hybrids are selected against, opposing the evolution of reproductive isolation, a problem that was recognised by Darwin. The problem does not occur in allopatric speciation with geographically separated populations, which can diverge with different sets of mutations. E. B. Poulton realized in 1903 that reproductive isolation could evolve through divergence, if each lineage acquired a different, incompatible allele of the same gene. Selection against the heterozygote would then directly create reproductive isolation, leading to the Bateson–Dobzhansky–Muller model, further elaborated by H. Allen Orr[92] and Sergey Gavrilets.[93] With reinforcement, however, natural selection can favor an increase in pre-zygotic isolation, influencing the process of speciation directly.[94]

Genetic basis[edit]

Genotype and phenotype[edit]

Main article: Genotype–phenotype distinction

Natural selection acts on an organism's phenotype, or physical characteristics. Phenotype is determined by an organism's genetic make-up (genotype) and the environment in which the organism lives. When different organisms in a population possess different versions of a gene for a certain trait, each of these versions is known as an allele. It is this genetic variation that underlies differences in phenotype. An example is the ABOblood typeantigens in humans, where three alleles govern the phenotype.[95]

Some traits are governed by only a single gene, but most traits are influenced by the interactions of many genes. A variation in one of the many genes that contributes to a trait may have only a small effect on the phenotype; together, these genes can produce a continuum of possible phenotypic values.[96]

Directionality of selection[edit]

Main article: Directional selection

When some component of a trait is heritable, selection alters the frequencies of the different alleles, or variants of the gene that produces the variants of the trait. Selection can be divided into three classes, on the basis of its effect on allele frequencies: directional, stabilizing, and purifying selection.[97] Directional selection occurs when an allele has a greater fitness than others, so that it increases in frequency, gaining an increasing share in the population. This process can continue until the allele is fixed and the entire population shares the fitter phenotype.[98] Far more common is stabilizing selection, which lowers the frequency of alleles that have a deleterious effect on the phenotype – that is, produce organisms of lower fitness. This process can continue until the allele is eliminated from the population. Purifying selection conserves functional genetic features, such as protein-coding genes or regulatory sequences, over time by selective pressure against deleterious variants.[99]

Some forms of balancing selection do not result in fixation, but maintain an allele at intermediate frequencies in a population. This can occur in diploid species (with pairs of chromosomes) when heterozygous individuals (with just one copy of the allele) have a higher fitness than homozygous individuals (with two copies). This is called heterozygote advantage or over-dominance, of which the best-known example is the resistance to malaria in humans heterozygous for sickle-cell anaemia. Maintenance of allelic variation can also occur through disruptive or diversifying selection, which favours genotypes that depart from the average in either direction (that is, the opposite of over-dominance), and can result in a bimodal distribution of trait values. Finally, balancing selection can occur through frequency-dependent selection, where the fitness of one particular phenotype depends on the distribution of other phenotypes in the population. The principles of game theory have been applied to understand the fitness distributions in these situations, particularly in the study of kin selection and the evolution of reciprocal altruism.[100][101]

Modern biology began in the nineteenth century with Charles Darwin's work on evolution by natural selection.
Aristotle considered whether different forms could have appeared, only the useful ones surviving.
Selection in action: resistance to antibiotics grows though the survival of individuals less affected by the antibiotic. Their offspring inherit the resistance.

—by Asa Gray

NATURAL SELECTION NOT INCONSISTENT WITH NATURAL THEOLOGY.

Atlantic Monthly for JulyAugust, and October, 1860, reprinted in 1861.

I

Novelties are enticing to most people; to us they are simply annoying. We cling to a long-accepted theory, just as we cling to an old suit of clothes. A new theory, like a new pair of breeches (the Atlantic still affects the older type of nether garment), is sure to have hard-fitting places; or, even when no particular fault can be found with the article, it oppresses with a sense of general discomfort. New notions and new styles worry us, till we get well used to them, which is only by slow degrees.

Wherefore, in Galileo’s time, we might have helped to proscribe, or to burn—had he been stubborn enough to warrant cremation—even the great pioneer of inductive research; although, when we had fairly recovered our composure, and had leisurely excogitated the matter, we might have come to conclude that the new doctrine was better than the old one, after all, at least for those who had nothing to unlearn.

Such being our habitual state of mind, it may well be believed that the perusal of the new book ‘On the Origin of Species by Means of Natural Selection’ left an uncomfortable impression, in spite of its plausible and winning ways. We were not wholly unprepared for it, as many of our contemporaries seem to have been. The scientific reading in which we indulge as a relaxation from severer studies had raised dim forebodings. Investigations about the succession of species in time, and their actual geographical distribution over the earth’s surface, were leading up from all sides and in various ways to the question of their origin. Now and then we encountered a sentence, like Prof. Owen’s ‘axiom of the continuous operation of the ordained becoming of living things,’ which haunted us like an apparition. For, dim as our conception must needs be as to what such oracular and grandiloquent phrases might really mean, we felt confident that they presaged no good to old beliefs. Foreseeing, yet deprecating, the coming time of trouble, we still hoped that, with some repairs and makeshifts, the old views might last out our days.Apres nous le deluge. Still, not to lag behind the rest of the world, we read the book in which the new theory is promulgated. We took it up, like our neighbors, and, as was natural, in a somewhat captious frame of mind.

Well, we found no cause of quarrel with the first chapter. Here the author takes us directly to the barn-yard and the kitchen-garden. Like an honorable rural member of our General Court, who sat silent until, near the close of a long session, a bill requiring all swine at large to wear pokes was introduced, when he claimed the privilege of addressing the house, on the proper ground that he had been ‘brought up among the pigs, and knew all about them’—so we were brought up among cows and cabbages; and the lowing of cattle, the cackle of hens, and the cooing of pigeons, were sounds native and pleasant to our ears. So ‘Variation under Domestication’ dealt with familiar subjects in a natural way, and gently introduced ‘Variation under Nature,’ which seemed likely enough. Then follows ‘Struggle for Existence’—a principle which we experimentally know to be true and cogent—bringing the comfortable assurance, that man, even upon Leviathan Hobbes’s theory of society, is no worse than the rest of creation, since all Nature is at war, one species with another, and the nearer kindred the more internecine—bringing in thousandfold confirmation and extension of the Malthusian doctrine that population tends far to outrun means of subsistence throughout the animal and vegetable world, and has to be kept down by sharp preventive checks; so that not more than one of a hundred or a thousand of the individuals whose existence is so wonderfully and so sedulously provided for ever comes to anything, under ordinary circumstances; so the lucky and the strong must prevail, and the weaker and ill-favored must perish; and then follows, as naturally as one sheep follows another, the chapter on ‘Natural Selection,’ Darwin’s cheval de bataille, which is very much the Napoleonic doctrine that Providence favors the strongest battalions—that, since many more individuals are born than can possibly survive, those individuals and those variations which possess any advantage, however slight, over the rest, are in the long-run sure to survive, to propagate, and to occupy the limited field, to the exclusion or destruction of the weaker brethren. All this we pondered, and could not much object to. In fact, we began to contract a liking for a system which at the outset illustrates the advantages of good breeding, and which makes the most ‘of every creature’s best.’

Could we ‘let by-gones be by-gones,’ and, beginning now, go on improving and diversifying for the future by natural selection, could we even take up the theory at the introduction of the actually existing species, we should be well content; and so, perhaps, would most naturalists be. It is by no means difficult to believe that varieties are incipient or possible species, when we see what trouble naturalists, especially botanists, have to distinguish between them—one regarding as a true species what another regards as a variety; when the progress of knowledge continually increases, rather than diminishes, the number of doubtful instances; and when there is less agreement than ever among naturalists as to what is the basis in Nature upon which our idea of species reposes, or how the word is to be defined. Indeed, when we consider the endless disputes of naturalists and ethnologists over the human races, as to whether they belong to one species or to more, and, if to more, whether to three, or five, or fifty, we can hardly help fancying that both may be right—or rather, that the uni-humanitarians would have been right many thousand years ago, and the multi-humanitarians will be several thousand years later; while at present the safe thing to say is, that probably there is some truth on both sides.

‘Natural selection,’ Darwin remarks, ‘leads to divergence of character; for the more living beings can be supported on the same area, the more they diverge in structure, habits, and constitution’ (a principle which, by-the-way, is paralleled and illustrated by the diversification of human labor); and also leads to much extinction of intermediate or unimproved forms. Now, though this divergence may ‘steadily tend to increase,’ yet this is evidently a slow process in Nature, and liable to much counteraction wherever man does not interpose, and so not likely to work much harm for the future. And if natural selection, with artificial to help it, will produce better animals and better men than the present, and fit them better to the conditions of existence, why, let it work, say we, to the top of its bent There is still room enough for improvement. Only let us hope that it always works for good: if not, the divergent lines on Darwin’s lithographic diagram of ‘Transmutation made Easy,’ ominously show what small deviations from the straight path may come to in the end.

The prospect of the future, accordingly, is on the whole pleasant and encouraging. It is only the backward glance, the gaze up the long vista of the past, that reveals anything alarming. Here the lines converge as they recede into the geological ages, and point to conclusions which, upon the theory, are inevitable, but hardly welcome. The very first step backward makes the negro and the Hottentot our blood-relations—not that reason or Scripture objects to that, though pride may. The next suggests a closer association of our ancestors of the olden time with ‘our poor relations’ of the quadrumanous family than we like to acknowledge. Fortunately, however—even if we must account for him scientifically —man with his two feet stands upon a foundation of his own. Intermediate links between the Bimana and the Quadrumana are lacking altogether; so that, put the genealogy of the brutes upon what footing you will, the four-handed races will not serve for our forerunners—at least, not until some monkey, live or fossil, is producible with great-toes, instead of thumbs, upon his nether extremities; or until some lucky geologist turns up the bones of his ancestor and prototype in France or England, who was so busy ‘napping the chuckie-stanes’ and chipping out flint knives and arrow-heads in the time of the drift, very many ages ago—before the British Channel existed, says Lyell —and until these men of the olden time are shown to have worn their great-toes in the divergent and thumblike fashion. That would be evidence indeed: but, until some testimony of the sort is produced, we must needs believe in the separate and special creation of man, however it may have been with the lower animals and with plants.

No doubt, the full development and symmetry of Darwin’s hypothesis strongly suggest the evolution of the human no less than the lower animal races out of some simple primordial animal—that all are equally ‘lineal descendants of some few beings which lived long before the first bed of the Silurian system was deposited.’ But, as the author speaks disrespectfully of spontaneous generation, and accepts a supernatural beginning of life on earth, in some form or forms of being which included potentially all that have since existed and are yet to be, he is thereby not warranted to extend his inferences beyond the evidence or the fair probability. There seems as great likelihood that one special origination should be followed by another upon fitting occasion (such as the introduction of man), as that one form should be transmuted into another upon fitting occasion, as, for instance, in the succession of species which differ from each other only in some details. To compare small things with great in a homely illustration: man alters from time to time his instruments or machines, as new circumstances or conditions may require and his wit suggest. Minor alterations and improvements he adds to the machine he possesses; he adapts a new rig or a new rudder to an old boat: this answers to Variation. ‘Like begets like,’ being the great rule in Nature, if boats could engender, the variations would doubtless be propagated, like those of domestic cattle. In course of time the old ones would be worn out or wrecked; the best sorts would be chosen for each particular use, and further improved upon; and so the primordial boat be developed into the scow, the skiff, the sloop, and other species of water-craft—the very diversification, as well as the successive improvements, entailing the disappearance of intermediate forms, less adapted to any one particular purpose; wherefore these go slowly out of use, and become extinct species: this is Natural Selection. Now, let a great and important advance be made, like that of steam navigation: here, though the engine might be added to the old vessel, yet the wiser and therefore the actual way is to make a new vessel on a modified plan: this may answer to Specific Creation. Anyhow, the one does not necessarily exclude the other. Variation and natural selection may play their part, and so may specific creation also. Why not?

This leads us to ask for the reasons which call for this new theory of transmutation. The beginning of things must needs lie in obscurity, beyond the bounds of proof, though within those of conjecture or of analogical inference. Why not hold fast to the customary view, that all species were directly, instead of indirectly, created after their respective kinds, as we now behold them—and that in a manner which, passing our comprehension, we intuitively refer to the supernatural? Why this continual striving after ‘the unattained and dim?’ why these anxious endeavors, especially of late years, by naturalists and philosophers of various schools and different tendencies, to penetrate what one of them calls ‘that mystery of mysteries,’ the origin of species?

To this, in general, sufficient answer may be found in the activity of the human intellect, ‘the delirious yet divine desire to know,’ stimulated as it has been by its own success in unveiling the laws and processes of inorganic Nature; in the fact that the principal triumphs of our age in physical science have consisted in tracing connections where none were known before, in reducing heterogeneous phenomena to a common cause or origin, in a manner quite analogous to that of the reduction of supposed independently originated species to a common ultimate origin—thus, and in various other ways, largely and legitimately extending the domain of secondary causes. Surely the scientific mind of an age which contemplates the solar system as evolved from a common revolving fluid mass—which, through experimental research, has come to regard light, heat, electricity, magnetism, chemical affinity, and mechanical power as varieties or derivative and convertible forms of one force, instead of independent species—which has brought the so-called elementary kinds of matter, such as the metals, into kindred groups, and pertinently raised the question, whether the members of each group may not be mere varieties of one species—and which speculates steadily in the direction of the ultimate unity of matter, of a sort of prototype or simple element which may be to the ordinary species of matter what the Protozoa or what the component cells of an organism are to the higher sorts of animals and plants—the mind of such an age cannot be expected to let the old belief about species pass unquestioned. It will raise the question, how the diverse sorts of plants and animals came to be as they are and where they are and will allow that the whole inquiry transcends its powers only when all endeavors have failed. Granting the origin to be supernatural or miraculous even, will not arrest the inquiry. All real origination, the philosophers will say, is supernatural; their very question is, whether we have yet gone back to the origin and can affirm that the present forms of plants and animals are the primordial, the miraculously created ones. And, even if they admit that, they will still inquire into the order of the phenomena, into the form of the miracle. You might as well expect the child to grow up content with what it is told about the advent of its infant brother. Indeed, to learn that the new-comer is the gift of God, far from lulling inquiry, only stimulates speculation as to how the precious gift was bestowed. That questioning child is father to the man—is philosopher in short-clothes.

Since, then, questions about the origin of species will be raised, and have been raised—and since the theorizings, however different in particulars, all proceed upon the notion that one species of plant or animal is somehow derived from another, that the different sorts which now flourish are lineal (or unlineal) descendants of other and earlier sorts—it now concerns us to ask, What are the grounds in Nature, the admitted facts, which suggest hypotheses of derivation in some shape or other? Reasons there must be, and plausible ones, for the persistent recurrence of theories upon this genetic basis. A study of Darwin’s book, and a general glance at the present state of the natural sciences, enable us to gather the following as among the most suggestive and influential. We can only enumerate them here, without much indication of their particular bearing. There is—

1. The general fact of variability, and the general tendency of the variety to propagate its like—the patent facts that all species vary more or less; that domesticated plants and animals, being in conditions favorable to the production and preservation of varieties, are apt to vary widely; and that, by interbreeding, any variety may be fixed into a race, that is, into a variety which comes true from seed. Many such races, it is allowed, differ from each other in structure and appearance as widely as do many admitted species; and it is practically very difficult, even impossible, to draw a clear line between races and species. Witness the human races, for instance. Wild species also vary, perhaps about as widely as those of domestication, though in different ways. Some of them apparently vary little, others moderately, others immoderately, to the great bewilderment of systematic botanists and zoologists, and increasing disagreement as to whether various forms shall be held to be original species or strong varieties. Moreover, the degree to which the descendants of the same stock, varying in different directions, may at length diverge, is unknown. All we know is, that varieties are themselves variable, and that very diverse forms have been educed from one stock.

2. Species of the same genus are not distinguished from each other by equal amounts of difference. There is diversity in this respect analogous to that of the varieties of a polymorphous species, some of them slight, others extreme. And in large genera the unequal resemblance shows itself in the clustering of the species around several types or central species, like satellites around their respective planets. Obviously suggestive this of the hypothesis that they were satellites, not thrown off by revolution, like the moons of Jupiter, Saturn, and our own solitary moon, but gradually and peacefully detached by divergent variation. That such closely-related species may be only varieties of higher grade, earlier origin, or more favored evolution, is not a very violent supposition. Anyhow, it was a supposition sure to be made.

3. The actual geographical distribution of species upon the earth’s surface tends to suggest the same notion. For, as a general thing, all or most of the species of a peculiar genus or other type are grouped in the same country, or occupy continuous, proximate, or accessible areas. So well does this rule hold, so general is the implication that kindred species are or were associated geographically, that most trustworthy naturalists, quite free from hypotheses of transmutation, are constantly inferring former geographical continuity between parts of the world now widely disjoined, in order to account thereby for certain generic similarities among their inhabitants; just as philologists infer former connection of races, and a parent language, to account for generic similarities among existing languages. Yet no scientific explanation has been offered to account for the geographical association of kindred species, except the hypothesis of a common origin.

4. Here the fact of the antiquity of creation, and in particular of the present kinds of the earth’s inhabitants, or of a large part of them, comes in to rebut the objection that there has not been time enough for any marked diversification of living things through divergent variation—not time enough for varieties to have diverged into what we call species.

So long as the existing species of plants and animals were thought to have originated a few thousand years ago, and without predecessors, there was no room for a theory of derivation of one sort from another, nor time enough even to account for the establishment of the races which are generally believed to have diverged from a common stock. Not so much that five or six thousand years was a short allowance for this; but because some of our familiar domesticated varieties of grain, of fowls, and of other animals, were pictured and mummified by the old Egyptians more than half that number of years ago, if not earlier. Indeed, perhaps the strongest argument for the original plurality of human species was drawn from the identification of some of the present races of men upon these early historical monuments and records.

But this very extension of the current chronology, if we may rely upon the archaeologists, removes the difficulty by opening up a longer vista. So does the discovery in Europe of remains and implements of prehistoric races of men, to whom the use of metals was unknown—men of thestone age, as the Scandinavian archaeologists designate them. And now, ‘axes and knives of flint, evidently wrought by human skill, are found in beds of the drift at Amiens (also in other places, both in France and England), associated with the bones of extinct species of animals.’ These implements, indeed, were noticed twenty years ago; at a place in Suffolk they have been exhumed from time to time for more than a century; but the full confirmation, the recognition of the age of the deposit in which the implements occur, their abundance, and the appreciation of their bearings upon most interesting questions, belong to the present time. To complete the connection of these primitive people with the fossil ages, the French geologists, we are told, have now ‘found these axes in Picardy associated with remains of Elephas primigeniusRhinoceros tichorhinusEquus fossilis, and an extinct species of Bos.’ In plain language, these workers in flint lived in the time of the mammoth, of a rhinoceros now extinct, and along with horses and cattle unlike any now existing—specifically different, as naturalists say, from those with which man is now associated. Their connection with existing human races may perhaps be traced through the intervening people of the stone age, who were succeeded by the people of the bronze age, and these by workers in iron. Now, various evidence carries back the existence of many of the present lower species of animals, and probably of a larger number of plants, to the same drift period. All agree that this was very many thousand years ago. Agassiz tells us that the same species of polyps which are now building coral walls around the present peninsula of Florida actually made that peninsula, and have been building there for many thousand centuries.

5. The overlapping of existing and extinct species, and the seemingly gradual transition of the life of the drift period into that of the present, may be turned to the same account. Mammoths, mastodons, and Irish elks, now extinct, must have lived down to human, if not almost to historic times. Perhaps the last dodo did not long outlive his huge New Zealand kindred. The aurochs, once the companion of mammoths, still survives, but owes his present and precarious existence to man’s care. Now, nothing that we know of forbids the hypothesis that some new species have been independently and supernaturally created within the period which other species have survived. Some may even believe that man was created in the days of the mammoth, became extinct, and was recreated at a later date. But why not say the same of the aurochs, contemporary both of the old man and of the new? Still it is more natural, if not inevitable, to infer that, if the aurochs of that olden time were the ancestors of the aurochs of the Lithuanian forests, so likewise were the men of that age the ancestors of the present human races. Then, whoever concludes that these primitive makers of rude flint axes and knives were the ancestors of the better workmen of the succeeding stone age, and these again of the succeeding artificers in brass and iron, will also be likely to suppose that the Equus and Bos of that time, different though they be, were the remote progenitors of our own horses and cattle. In all candor we must at least concede that such considerations suggest a genetic descent from the drift period down to the present, and allow time enough—if time is of any account— for variation and natural selection to work out some appreciable results in the way of divergence into races, or even into so-called species. Whatever might have been thought, when geological time was supposed to be separated from the present era by a clear line, it is now certain that a gradual replacement of old forms by new ones is strongly suggestive of some mode of origination which may still be operative. When species, like individuals, were found to die out one by one, and apparently to come in one by one, a theory for what Owen sonorously calls ‘the continuous operation of the ordained becoming of living things’ could not be far off.

That all such theories should take the form of a derivation of the new from the old seems to be inevitable, perhaps from our inability to conceive of any other line of secondary causes in this connection. Owen himself is apparently in travail with some transmutation theory of his own conceiving, which may yet see the light, although Darwin’s came first to the birth. Different as the two theories will probably be, they cannot fail to exhibit that fundamental resemblance in this respect which betokens a community of origin, a common foundation on the general facts and the obvious suggestions of modern science. Indeed—to turn the point of a pungent simile directed against Darwin—the difference between the Darwinian and the Owenian hypotheses may, after all, be only that between homoeopathic and heroic doses of the same drug.

If theories of derivation could only stop here, content with explaining the diversification and succession of species between the teritiary period and the present time, through natural agencies or secondary causes still in operation, we fancy they would not be generally or violently objected to by the savants of the present day. But it is hard, if not impossible, to find a stopping-place. Some of the facts or accepted conclusions already referred to, and several others, of a more general character, which must be taken into the account, impel the theory onward with accumulated force. Vires (not to say virusacquirit eundo. The theory hitches on wonderfully well to Lyell’s uniformitarian theory in geology—that the thing that has been is the thing that is and shall be—that the natural operations now going on will account for all geological changes in a quiet and easy way, only give them time enough, so connecting the present and the proximate with the farthest past by almost imperceptible gradations—a view which finds large and increasing, if not general, acceptance in physical geology, and of which Darwin’s theory is the natural complement.

So the Darwinian theory, once getting a foothold, marches boldly on, follows the supposed near ancestors of our present species farther and yet farther back into the dim past, and ends with an analogical inference which ‘makes the whole world kin.’ As we said at the beginning, this upshot discomposes us. Several features of the theory have an uncanny look. They may prove to be innocent: but their first aspect is suspicious, and high authorities pronounce the whole thing to be positively mischievous. In this dilemma we are going to take advice. Following the bent of our prejudices, and hoping to fortify these by new and strong arguments, we are going now to read the principal reviews which undertake to demolish the theory—with what result our readers shall be duly informed.

II

‘I can entertain no doubt, after the most deliberate study and dispassionate judgment of which I am capable, that the view which most naturalists entertain, and which I formerly entertained, namely, that each species has been independently created, is erroneous. I am fully convinced that species are not immutable; but that those belonging to what are called the same genera are lineal descendants of some other and generally extinct species, in the same manner as the acknowledged varieties of any one species are the descendants of that species. Furthermore, I am convinced that Natural Selection has been the main, but not exclusive, means of modification.’

This is the kernel of the new theory, the Darwinian creed, as recited at the close of the introduction to the remarkable book under consideration. The questions, ‘What will he do with it?’ and ‘How far will he carry it?’ the author answers at the close of the volume:

‘I cannot doubt that the theory of descent with modification embraces all the members of the same class.’ Furthermore, ‘I believe that all animals have descended from at most only four or five progenitors, and plants from an equal or lesser number.’

Seeing that analogy as strongly suggests a further step in the same direction, while he protests that ‘analogy may be a deceitful guide,’ yet he follows its inexorable leading to the inference that—

‘Probably all the organic beings which have ever lived on this earth have descended from some one primordial form, into which life was first breathed.’

In the first extract we have the thin end of the wedge driven a little way; in the last, the wedge driven home.

We have already sketched some of the reasons suggestive of such a theory of derivation of species, reasons which gave it plausibility, and even no small probability, as applied to our actual world and to changes occurring since the latest tertiary period. We are well pleased at this moment to find that the conclusions we were arriving at in this respect are sustained by the very high authority and impartial judgment of Pictet, the Swiss paleontologist. In his review of Darwin’s book — the fairest and most admirable opposing one that has appeared—he freely accepts that ensemble of natural operations which Darwin impersonates under the now familiar name of Natural Selection, allows that the exposition throughout the first chapters seems ‘a la fois prudent et fort,’ and is disposed to accept the whole argument in its foundations, that is, so far as it relates to what is now going on, or has taken place in the present geological period—which period he carries back through the diluvial epoch to the borders of the tertiary. Pictet accordingly admits that the theory will very well account for the origination by divergence of nearly-related species, whether within the present period or in remoter geological times; a very natural view for him to take, since he appears to have reached and published, several years ago, the pregnant conclusion that there most probably was some material connection between the closely-related species of two successive faunas, and that the numerous close species, whose limits are so difficult to determine, were not all created distinct and independent. But while thus accepting, or ready to accept, the basis of Darwin’s theory, and all its legitimate direct inferences, he rejects the ultimate conclusions, brings some weighty arguments to bear against them, and is evidently convinced that he can draw a clear line between the sound inferences, which he favors, and the unsound or unwarranted theoretical deductions, which he rejects. We hope he can.

This raises the question, Why does Darwin press his theory to these extreme conclusions? Why do all hypotheses of derivation converge so inevitably to one ultimate point? Having already considered some of the reasons which suggest or support the theory at its outset—which may carry it as far as such sound and experienced naturalists as Pictet allow that it may be true—perhaps as far as Darwin himself unfolds it in the introductory proposition cited at the beginning of this article—we may now inquire after the motives which impel the theorist so much farther. Here proofs, in the proper sense of the word, are not to be had. We are beyond the region of demonstration, and have only probabilities to consider. What are these probabilities? What work will this hypothesis do to establish a claim to be adopted in its completeness? Why should a theory which may plausibly enough account for the diversification of the species of each special type or genus be expanded into a general system for the origination or successive diversification of all species, and all special types or forms, from four or five remote primordial forms, or perhaps from one? We accept the theory of gravitation because it explains all the facts we know, and bears all the tests that we can put it to. We incline to accept the nebular hypothesis, for similar reasons; not because it is proved—thus far it is incapable of proof —but because it is a natural theoretical deduction from accepted physical laws, is thoroughly congruous with the facts, and because its assumption serves to connect and harmonize these into one probable and consistent whole. Can the derivative hypothesis be maintained and carried out into a system on similar grounds? If so, however unproved, it would appear to be a tenable hypothesis, which is all that its author ought now to claim. Such hypotheses as, from the conditions of the case, can neither be proved nor disproved by direct evidence or experiment, are to be tested only indirectly, and therefore imperfectly, by trying their power to harmonize the known facts, and to account for what is otherwise unaccountable. So the question comes to this: What will an hypothesis of the derivation of species explain which the opposing view leaves unexplained?

Questions these which ought to be entertained before we take up the arguments which have been advanced against this theory. We can barely glance at some of the considerations which Darwin adduces, or will be sure to adduce in the future and fuller exposition which is promised. To display them in such wise as to indoctrinate the unscientific reader would require a volume. Merely to refer to them in the most general terms would suffice for those familiar with scientific matters, but would scarcely enlighten those who are not. Wherefore let these trust the impartial Pictet, who freely admits that, ‘in the absence of sufficient direct proofs to justify the possibility of his hypothesis, Mr. Darwin relies upon indirect proofs, the bearing of which is real and incontestable;’ who concedes that ‘his theory accords very well with the great facts of comparative anatomy and zoology—comes in admirably to explain unity of composition of organisms, also to explain rudimentary and representative organs, and the natural series of genera and species—equally corresponds with many paleontological data—agrees well with the specific resemblances which exist between two successive faunas, with the parallelism which is sometimes observed between the series of paleontological succession and of embryonal development,’ etc.; and finally, although he does not accept the theory in these results, he allows that ‘it appears to offer the best means of explaining the manner in which organized beings were produced in epochs anterior to our own.’

What more than this could be said for such an hypothesis? Here, probably, is its charm, and its strong hold upon the speculative mind. Unproven though it be, and cumbered prima facie with cumulative improbabilities as it proceeds, yet it singularly accords with great classes of facts otherwise insulated and enigmatic, and explains many things which are thus far utterly inexplicable upon any other scientific assumption.

We have said that Darwin’s hypothesis is the natural complement to Lyell’s uniformitarian theory in physical geology. It is for the organic world what that is for the inorganic; and the accepters of the latter stand in a position from which to regard the former in the most favorable light. Wherefore the rumor that the cautious Lyell himself has adopted the Darwinian hypothesis need not surprise us. The two views are made for each other, and, like the two counterpart pictures for the stereoscope, when brought together, combine into one apparently solid whole.

If we allow, with Pictet, that Darwin’s theory will very well serve for all that concerns the present epoch of the world’s history—an epoch in which this renowned paleontologist includes the diluvial or quaternary period—then Darwin’s first and foremost need in his onward course is a practicable road from this into and through the tertiary period, the intervening region between the comparatively near and the far remote past. Here Lyell’s doctrine paves the way, by showing that in the physical geology there is no general or absolute break between the two, probably no greater between the latest tertiary and the quaternary period than between the latter and the present time. So far, the Lyellian view is, we suppose, generally concurred in. It is largely admitted that numerous tertiary species have continued down into the quaternary, and many of them to the present time. A goodly percentage of the earlier and nearly half of the later tertiary mollusca, according to Des Hayes, Lyell, and, if we mistake not, Bronn, still live. This identification, however, is now questioned by a naturalist of the very highest authority. But, in its bearings on the new theory, the point here turns not upon absolute identity so much as upon close resemblance. For those who, with Agassiz, doubt the specific identity in any of these cases, and those who say, with Pictet, that ‘the later tertiary deposits contain in general the debris of speciesvery nearly related to those which still exist, belonging to the same genera, but specifically different,’ may also agree with Pictet, that the nearly-related species of successive faunas must or may have had ‘a material connection.’ But the only material connection that we have an idea of in such a case is a genealogical one. And the supposition of a genealogical connection is surely not unnatural in such cases—is demonstrably the natural one as respects all those tertiary species which experienced naturalists have pronounced to be identical with existing ones, but which others now deem distinct. For to identify the two is the same thing as to conclude the one to be the ancestor of the other. No doubt there are differences between the tertiary and the present individuals, differences equally noticed by both classes of naturalists, but differently estimated. By the one these are deemed quite compatible, by the other incompatible, with community of origin. But who can tell us what amount of difference is compatible with community of origin? This is the very question at issue, and one to be settled by observation alone. Who would have thought that the peach and the nectarine came from one stock? But, this being proved, is it now very improbable that both were derived from the almond, or from some common amygdaline progenitor? Who would have thought that the cabbage, cauliflower, broccoli, kale, and kohlrabi are derivatives of one species, and rape or colza, turnip, and probably ruta-baga, of another species? And who that is convinced of this can long undoubtingly hold the original distinctness of turnips from cabbages as an article of faith? On scientific grounds may not a primordial cabbage or rape be assumed as the ancestor of all the cabbage races, on much the same ground that we assume a common ancestry for the diversified human races? If all our breeds of cattle came from one stock why not this stock from the auroch, which has had all the time between the diluvial and the historic periods in which to set off a variation perhaps no greater than the difference between some sorts of domestic cattle?

That considerable differences are often discernible between tertiary individuals and their supposed descendants of the present day affords no argument against Darwin’s theory, as has been rashly thought, but is decidedly in its favor. If the identification were so perfect that no more differences were observable between the tertiary and the recent shells than between various individuals of either, then Darwin’s opponents, who argue the immutability of species from the ibises and cats preserved by the ancient Egyptians being just like those of the present day, could triumphantly add a few hundred thousand years more to the length of the experiment and to the force of their argument.

As the facts stand, it appears that, while some tertiary forms are essentially undistinguishable from existing ones, others are the same with a difference, which is judged not to be specific or aboriginal; and yet others show somewhat greater differences, such as are scientifically expressed by calling them marked varieties, or else doubtful species; while others, differing a little more, are confidently termed distinct, but nearly-related species. Now, is not all this a question of degree, of mere gradation of difference? And is it at all likely that these several gradations came to be established in two totally different ways—some of them (though naturalists can’t agree which) through natural variation, or other secondary cause, and some by original creation, without secondary cause? We have seen that the judicious Pictet answers such questions as Darwin would have him do, in affirming that, in all probability, the nearly-related species of two successive faunas were materially connected, and that contemporaneous species, similarly resembling each other, were not all created so, but have become so. This is equivalent to saying that species (using the term as all naturalists do, and must continue to employ the word) have only a relative, not an absolute fixity; that differences fully equivalent to what are held to be specific may arise in the course of time, so that one species may at length be naturally replaced by another species a good deal like it, or may be diversified into two, three, or more species, or forms as different as species. This concedes all that Darwin has a right to ask, all that he can directly infer from evidence. We must add that it affords a locus standi, more or less tenable, for inferring more.

Here another geological consideration comes in to help on this inference. The species of the later tertiary period for the most part not only resembled those of our days—many of them so closely as to suggest an absolute continuity—but also occupied in general the same regions that their relatives occupy now. The same may be said, though less specially, of the earlier tertiary and of the later secondary; but there is less and less localization of forms as we recede, yet some localization even in palaeozoic times. While in the secondary period one is struck with the similarity of forms and the identity of many of the species which flourished apparently at the same time in all or in the most widely-separated parts of the world, in the tertiary epoch, on the contrary, along with the increasing specialization of climates and their approximation to the present state, we find abundant evidence of increasing localization of orders, genera and species, and this localization strikingly accords with the present geographical distribution of the same groups of species Where the imputed forefathers lived their relatives and supposed descendants now flourish All the actual classes of the animal and vegetable kingdoms were represented in the tertiary faunas and floras and in nearly the same proportions and the same diversities as at present The faunas of what is now Europe, Asia America and Australia, differed from each other much as they now differ: in fact—according to Adolphe Brongniart, whose statements we here condense 1 —the inhabitants of these different regions appear for the most part to have acquired, before the close of the tertiary period, the characters which essentially distinguish their existing faunas. The Eastern Continent had then, as now, its great pachyderms, elephants, rhinoceros, hippopotamus; South America, its armadillos, sloths, and anteaters; Australia, a crowd of marsupials; and the very strange birds of New Zealand had predecessors of similar strangeness. Everywhere the same geographical distribution as now, with a difference in the particular area, as respects the northern portion of the continents, answering to a warmer climate then than ours, such as allowed species of hippopotamus, rhinoceros, and elephant, to range even to the regions now inhabited by the reindeer and the musk-ox, and with the serious disturbing intervention of the glacial period within a comparatively recent time. Let it be noted also that those tertiary species which have continued with little change down to our days are the marine animals of the lower grades, especially mollusca. Their low organization, moderate sensibility, and the simple conditions of an existence in a medium like the ocean, not subject to great variation and incapable of sudden change, may well account for their continuance; while, on the other hand, the more intense, however gradual, climatic vicissitudes on land, which have driven all tropical and subtropical forms out of the higher latitudes and assigned to them their actual limits, would be almost sure to extinguish such huge and unwieldy animals as mastodons, mammoths, and the like, whose power of enduring altered circumstances must have been small.

This general replacement of the tertiary species of a country by others so much like them is a noteworthy fact. The hypothesis of the independent creation of all species, irrespective of their antecedents, leaves this fact just as mysterious as is creation itself; that of derivation undertakes to account for it. Whether it satisfactorily does so or not, it must be allowed that the facts well accord with that hypothesis. The same may be said of another conclusion, namely, that the geological succession of animals and plants appears to correspond in a general way with their relative standing or rank in a natural system of classification. It seems clear that, though no one of the grand types of the animal kingdom can be traced back farther than the rest, yet the lower classes long preceded the higher; that there has been on the whole a steady progression within each class and order; and that the highest plants and animals have appeared only in relatively modern times. It is only, however, in a broad sense that this generalization is now thought to hold good. It encounters many apparent exceptions, and sundry real ones. So far as the rule holds, all is as it should be upon an hypothesis of derivation.

The rule has its exceptions. But, curiously enough, the most striking class of exceptions, if such they be, seems to us even more favorable to the doctrine of derivation than is the general rule of a pure and simple ascending gradation. We refer to what Agassiz calls prophetic and synthetic types; for which the former name may suffice, as the difference between the two is evanescent.

‘It has been noticed,’ writes our great zoölogist, ‘that certain types, which are frequently prominent among the representatives of past ages, combine in their structure peculiarities which at later periods are only observed separately in different, distinct types. Sauroid fishes before reptiles, Pterodactyles before birds, Ichthyosauri before dolphins, etc. There are entire families, of nearly every class of animals, which in the state of their perfect development exemplify such prophetic relations. . . . The sauroid fishes of the past geological ages are an example of this kind These fishes which preceded the appearance of reptiles present a combination of ichthyic and reptilian characters not to be found in the true members of this class, which form its bulk at present. The Pterodactyles, which preceded the class of birds, and the Ichthyosauri, which preceded the Cetacea, are other examples of such prophetic types.’—(Agassiz, ‘Contributions, Essay on Classification,’ p. 117.)

Now, these reptile-like fishes, of which gar-pikes are the living representatives, though of earlier appearance, are admittedly of higher rank than common fishes. They dominated until reptiles appeared, when they mostly gave place to (or, as the derivationists will insist, were resolved by divergent variation and natural selection into) common fishes, destitute of reptilian characters, and saurian reptiles—the intermediate grades, which, according to a familiar piscine saying, are ‘neither fish, flesh, nor good red-herring,’ being eliminated and extinguished by natural consequence of the struggle for existence which Darwin so aptly portrays. And so, perhaps, of the other prophetic types. Here type and antitype correspond. If these are true prophecies, we need not wonder that some who read them in Agassiz’s book will read their fulfillment in Darwin’s.

Note also, in this connection, that along with a wonderful persistence of type, with change of species, genera, orders, etc., from formation to formation, no species and no higher group which has once unequivocally died out ever afterward reappears. Why is this, but that the link of generation has been sundered? Why, on the hypothesis of independent originations, were not failing species recreated, either identically or with a difference, in regions eminently adapted to their well-being? To take a striking case. That no part of the world now offers more suitable conditions for wild horses and cattle than the pampas and other plains of South America, is shown by the facility with which they have there run wild and enormously multiplied, since introduced from the Old World not long ago. There was no wild American stock. Yet in the times of the mastodon and megatherium, at the dawn of the present period, wild-horses—certainly very much like the existing horse—roamed over those plains in abundance. On the principle of original and direct created adaptation of species to climate and other conditions, why were they not reproduced, when, after the colder intervening era, those regions became again eminently adapted to such animals? Why, but because, by their complete extinction in South America, the line of descent was there utterly broken? Upon the ordinary hypothesis, there is no scientific explanation possible of this series of facts, and of many others like them. Upon the new hypothesis, ‘the succession of the same types of structure within the same areas during the later geological periods ceases to be mysterious, and is simply explained by inheritance.’ Their cessation is failure of issue.

Along with these considerations the fact (alluded to on page 98) should be remembered that, as a general thing, related species of the present age are geographically associated. The larger part of the plants, and still more of the animals, of each separate country are peculiar to it; and, as most species now flourish over the graves of their by-gone relatives of former ages, so they now dwell among or accessibly near their kindred species.

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