History of evolutionary thought

This article is about the history of evolutionary thought in biology. For the history of evolutionary thought in
the social sciences, see social evolutionism. For the history of evolutionary thought generally, see
evolutionism.
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Evolutionary thought, the idea that species develop over time, has its roots in antiquity, in the ideas of the Greeks,
Romans, Chinese and Muslims. However, until the 18th century, Western biological thinking was dominated by
essentialism, the idea that living forms are unchanging. During the Enlightenment, evolutionary cosmology and the
mechanical philosophy spread from the physical sciences to natural history. Naturalists such as Maupertuis and
Buffon began to focus on the variability of species, and Erasmus Darwin included even more explicit evolutionary
speculations in his writings. The emergence of paleontology and with it the notion of extinction, combined with the
dramatic expansion of known species undermined the static essentialist view of nature. The first fully developed
theory of evolution was proposed by Jean-Baptiste Lamarck in the early 19th century; Lamarck's theory of
transmutation held that species had an innate drive pushing them up the great chain of being, and that the
mechanism of inheritance of acquired characteristics helped them adapt to local conditions.

The evolutionary theory often referred to as Darwinism was first publicly put forward by Charles Darwin and Alfred
Russel Wallace and discussed in detail in Darwin's On the Origin of Species (1859). Unlike Lamarck's theory,
Darwinism proposed common descent and a branching tree of life. It was based on the idea of natural selection, and
it synthesized evidence from animal husbandry, biogeography, geology, morphology, and embryology. The debate
over Origin raised serious questions about the place of humanity in nature, and was a key step in the process of
methodological naturalism replacing natural theology in the sciences.

Darwin's work led to the rapid acceptance of evolution, but the actual mechanism he proposed, natural selection, was
not widely accepted until the 1930s. Most biologists argued that other factors drove evolution, such as inheritance of
acquired characteristics (neo-Lamarckism), an innate drive for change (orthogenesis), or sudden large mutations
(saltationism). With the rediscovery of Mendelian genetics, and with T.H. Morgan's studies on mutation, scientists
began to better understand the nature of inheritance and variation. This eventually led to the synthesis of natural
selection with Mendelian genetics during the 1920s and 1930s, forming the new discipline of population genetics.
Throughout the 1930s and 1940s, population genetics became integrated with other branches of biology, finally
resulting in a unified theory of evolution - the modern evolutionary synthesis, which restored natural selection to a
central role in evolutionary theory.

Following this establishment of evolutionary biology, the theory of evolution developed rapidly. Studies
of mutation and variation in natural populations, combined with biogeography and systematics, led to sophisticated
mathematical and causal models of evolution. Paleontology and comparative anatomy allowed more detailed
reconstructions of the history of life. After the rise of molecular genetics in the 1950s, the field of molecular
evolution developed, based on DNA, RNA, and protein sequences. The gene-centered view of evolution then rose to
prominence in the 1960s, followed by the neutral theory of molecular evolution, sparking debates over adaptationism,
the units of selection, and the importance of genetic drift. In the late 20th century, genetic sequencing led to a
reorganization of the tree of life into the three-domain system, and the newly-recognized factors
of symbiogenesis and horizontal gene transfer have introduced yet more complexity into evolutionary history.

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Contents

 1 Antiquity
o 1.1 Greek thought
 1.1.1 Plato and the theory of forms
 1.1.2 Aristotle and the ladder of life
o 1.2 Chinese thought
o 1.3 Roman thought
 2 Middle Ages
o 2.1 Christian thought and the great chain of
being
o 2.2 Islamic thought and the struggle for
existence
 3 Modern period
o 3.1 Early modern thought
o 3.2 Early 19th century
 3.2.1 Paleontology and geology
 3.2.2 Lamarckism and
transmutation
 3.2.3 Opposition to transmutation
 3.2.4 Anticipations of natural
selection
 3.2.5 Natural selection
o 3.3 1859–1930s: Darwin and after Darwin
 3.3.1 Application of the theory to
humans
 3.3.2 Alternatives to natural
selection
 3.3.2.1 Theistic evolution
 3.3.2.2 Neo-Lamarckism
 3.3.2.3 Orthogenesis
 3.3.2.4 Saltationism
 3.3.3 Mendelian genetics,
biometrics, and mutation
o 3.4 1920s–1940s:
 3.4.1 Population genetics
 3.4.2 Modern evolutionary
synthesis
o 3.5 1940s–1960s: Molecular biology
o 3.6 Since the 1960s:
 3.6.1 Gene centered view of
evolution
 3.6.2 Punctuated equilibrium
 3.6.3 Sociobiology
 3.6.4 Evolutionary paths and
processes
 3.6.5 Microbiology and horizontal
gene transfer
 3.6.6 Evo-devo
 4 Unconventional evolutionary thought
o 4.1 Gaia hypothesis
 5 Notes
 6 References
 7 See also
Antiquity
Greek thought

Some Greek philosophers discussed ideas that involved forms of organic evolution. Anaximander claimed that life
had originally developed in the sea and only later moved onto land, and Empedocles also discussed a non-
supernatural origin for living things.[1] Empedocles even suggested a form of natural selection, which Aristotle
summarized as, "Wherever then all the parts came about [to be] just what they would have been if they had come to
be for an end, such things survived, being organized spontaneously in a fitting way; whereas those which grew
otherwise perished and continue to perish..."[2]

Plato and the theory of forms

Plato (427/8–347/8 BC) was, in the words of biologist and historian Ernst Mayr, "the great antihero of
evolutionism,"[3] as he established the philosophy of essentialism, which he called the theory of forms. This theory
holds that objects observed in the real world are only reflections of a limited number of essences (eide). Variation is
merely the result of an imperfect reflection of these constant essences. In his Timaeus, Plato set forth the idea that
God had created the cosmos and everything in it because He is good, and hence, "... free from jealousy, He desired
that all things should be as like Himself as they could be." God created all conceivable forms of life, since "... without
them the universe will be incomplete, for it will not contain every kind of animal which it ought to contain, if it is to be
perfect." This idea, that all potential forms of life are essential to a perfect creation, is called the plenitude principle,
and it greatly influenced Christian thought.[4]

Aristotle and the ladder of life

Aristotle, (384–322 BC), one of the most influential of the Greek philosophers, is the earliest natural historian whose
work has come down to us in any real detail. His writings on biology, the result of his research into natural history on
the isle of Lesbos, have survived in the form of four books, usually known by their Latin names, [5]

 De anima (on the essence of life)

 Historia animalium (inquiries about animals)
 De generatione animalium (reproduction)
 De partibus animalium (anatomy)

These works contain some remarkably astute observations and interpretations by Aristotle, along with sundry myths
and mistakes — reflecting the uneven state of knowledge during his time. The most striking passages are about the
sea-life visible in observations on Lesbos, and available from the catches of fishermen. He separated the aquatic
mammals from the fish, and knew that sharks and rays were part of the group he called Selachē (selachians). His
description of the hectocotyl arm (see cephalopod) was about two thousand years ahead of its time, and was in fact
widely disbelieved until the nineteenth century. His observations on catfish, electric fish of the genus Torpedo, and
angler-fish are also exceptional, as is his writing on cephalopod molluscs Octopus, cuttlefish and the paper nautilus.[5]

However, for Charles Singer, "Nothing is more remarkable than [Aristotle's] efforts to [exhibit] the relationships of
living things as a scala naturæ".[5] This scala naturæ, described in History of Animals, classified organisms in relation
to a hierarchical "Ladder of Life" or "Chain of Being", placing them according to complexity of structure and function,
with organisms that showed greater vitality and ability to move described as "higher organisms". [4]

Chinese thought

Ideas on evolution were expressed by ancient Chinese thinkers such as Zhuangzi (Chang Tzu). According to Joseph
Needham, Taoism explicitly denied the fixity of biological species.[6]

Roman thought

Titus Lucretius Carus (d. 50 BC), the Roman Epicurean and atomist, wrote the poem On the Nature of Things (De
rerum natura), describing the development of the living earth in stages: from atoms colliding in the void as swirls of
dust to early plants and animals springing from the early earth's substance, then a succession of animals, including a
series of progressively less brutish humans. Lucretius may be seen as the earliest believer in hard inheritance. He
said "For if each organism had not its own genetic bodies, how could we with certainty assign each to its mother?".
[7]
The essence of Lucretius' ideas was naturalism, and the avoidance of supernatural interventions or explanations.

Middle Ages
Christian thought and the great chain of being

During the so-called Dark Ages, Greek classical learning was all but lost to the West. However, contact with the
Islamic world, where Greek manuscripts were preserved and elaborated on, soon led to a massive spate of Latin
translations in the 12th century. Europeans were thus re-introduced to the works of Plato and Aristotle, as well as
Islamic thought. Christian thinkers combined Aristotlean classification with Plato's ideas of the goodness of God, and
of all potential life forms being present in a perfect creation, to organize all inanimate, animate, and spiritual beings,
into a huge interconnected system: the Scala Naturæ, or great chain of being.

Within this system, everything that existed could be placed in order, from "lowest" to "highest", with Hell at the bottom
and God at the top — below God, an angelic hierarchy marked by the orbits of the planets, mankind in an
intermediate position, and worms the lowest of the animals. As the universe was ultimately perfect, the Great Chain
was also perfect. There were no empty links in the chain, and no link was represented by more than one species. But
this also implied that, since every link is occupied, and none can be occupied twice, then no species can ever move
from one position to another. To do so would leave one level empty and put two species on another. Thus, in this
Christianized version of Plato's perfect universe, species could never change, but must remain forever fixed, in
accordance with the text of Genesis. For humans to forget their position was even seen as sinful, whether they
behaved like lower animals or aspired to a higher station than was given them by their Creator.

Creatures on adjacent steps were expected to closely resemble each other, an idea expressed in a saying
which Charles Darwin often quoted: natura non facit saltum ("nature does not make leaps").[8][4] This basic concept of
the great chain of being greatly influenced the thinking of Western civilization for centuries (and still has an influence
today). It also formed a part of the argument from design presented by natural theology. As a classification system, it
became the major organizing principle and foundation of the emerging science of biology in the 17th and 18th
centuries.[4]

Islamic thought and the struggle for existence

Main article: Early Islamic philosophy – Evolution

Whereas Greek and Roman evolutionary ideas more or less died out in Europe after the fall of the Roman Empire,
they were not lost to Islamic scientists and philosophers. In the Islamic Golden Age, early theories
on evolution and natural selection were widely taught in Madrasahs (Islamic schools). According to al-Khazini, writing
in the 12th century, ideas on evolution were widespread among "common people" in the Islamic world of his time.
John William Draper, the 19th century scientist, philosopher and historian, referred to the early Muslim theories on
evolution as the "Mohammedan theory of evolution", using the then-current term for Muslims. He compared these
early theories to the modern Darwinian theory of evolution of his time, arguing that the former were developed "...
much farther than we are disposed to do, extending them even to inorganic or mineral things." [9]

The first Muslim biologist and philosopher to develop a theory of evolution was the Afro-Arab writer al-Jahiz in the 9th
century. He considered the effects of the environment on the likelihood of an animal to survive and evolve, and first
described the struggle for existence, as well as an early approach to natural selection.[10][11]

Ibn Miskawayh's al-Fawz al-Asghar and the Brethren of Purity's Encyclopedia of the Brethren of Purity (The Epistles
of Ikhwan al-Safa) expressed sophisticated evolutionary ideas regarding how the species evolved: from matter into
vapor and thence to water, then minerals into plants and then animals, leading to apes and, finally, humans. [12][13]

The polymath Ibn al-Haytham wrote a book in which he argued for evolutionism (although not natural selection).
Numerous other Islamic scholars and scientists, such as Abū Rayhān al-Bīrūnī, Nasir al-Din Tusi, and Ibn Khaldun
discussed and developed these ideas. Translated into Latin, these works began to appear in the West after the
Renaissance and may have had an impact on Western science.[11]
Modern period
Early modern thought

The word evolution (from the Latin evolutio, meaning "to unroll like a scroll") appeared in English in the 17th century,
referring to an orderly sequence of events, particularly one in which the outcome was somehow contained within it
from the start. Notably, in 1677 Sir Matthew Hale, attacking the atheistic atomism of Democritus and Epicurus. used
the term evolution to describe his opponent's ideas that vibrations and collisions of atoms in the void — without divine
intervention — had formed "Primordial Seeds" (semina) which were the "immediate, primitive, productive Principles of
Men, Animals, Birds and Fishes."[14] For Hale, this mechanism was "absurd", because "it must have potentially at
least the whole Systeme of Humane Nature, or at least that Ideal Principle or Configuration thereof, in
the evolution whereof the complement and formation of the Humane Nature must consist ... and all this drawn from a
fortuitous coalition of senseless and dead Atoms."[14]

While Hale (ironically) first used the term evolution in arguing against the exact mechanistic view the word would
come to symbolize, he also demonstrates that at least some evolutionist theories explored between 1650 and 1800
postulated that the universe, including life on earth, had developed mechanically, entirely without divine guidance.
Around this time, the mechanical philosophy of Descartes, reinforced by the physics of Galileo and Newton, began to
encourage the machine-like view of the universe which would come to characterise the scientific revolution.
[15]
However, most contemporary theories of evolution, including those developed by the German idealist philosophers
Schelling and Hegel (and mocked by Schopenhauer), held that evolution was a fundamentally spiritual process, with
the entire course of natural and human evolution being "a self-disclosing revelation of the Absolute". [16]

Typical of these theorists, Gottfried Leibniz postulated in 1714 that "monads" inside objects caused motion by internal
forces, and maintained that "the 'germs' of all things have always existed ... [and] contain within themselves an
internal principle of development which drives them on through a vast series of metamorphoses" to become the
geological formations, lifeforms, psychologies, and civilizations of the present. Leibniz clearly felt that evolution
proceeded on divine principles — in his De rerum originatione radicali (1697), he wrote: "A cumulative increase of the
beauty and universal perfection of the works of God, a perpetual and unrestricted progress of the universe as a whole
must be recognized, such that it advances to a higher state of cultivation." [17] Others, such as J. G. von Herder,
expressed similar ideas.[18]

In his Venus Physique in 1745, and System of Nature in 1751, Pierre Louis Maupertuis veered toward more
materialist ground. He wrote of natural modifications occurring during reproduction and accumulating over the course
of many generations, producing races and even new species. He also anticipated in general terms the idea of natural
selection.[19]

Vague and general ideas of evolution continued to proliferate among the mid-eighteenth century Enlightenment
philosophers. G. L. L. Buffon suggested that what most people referred to as species were really just well-marked
varieties. He thought that the members of what was then called a genus (which in terms of modern scientific
classification would be considered a family) are all descended from a single, common ancestor. The ancestor of each
family had arisen through spontaneous generation; environmental effects then caused them to diverge into different
species. Buffon's concept of evolution was strictly limited. He believed there were "internal molds" that shaped the
spontaneous generation of each family and that the families themselves were entirely and eternally distinct. Thus,
lions and tigers and house cats could all share a common ancestor, but dogs and house cats could not. [20][21] Although
Darwin's foreword to his 6th edition of Origin credited Aristotle with foreshadowing the concept of natural selection, he
also wrote that "the first author who in modern times has treated it in a scientific spirit was Buffon". [22]

Between 1767 and 1792 James Burnett, Lord Monboddo included in his writings not only the concept that that man
had descended from primates, but also that, in response to their environment, creatures had found methods of
transforming their characteristics over long time intervals. He also produced research on the evolution of linguistics,
which was cited by Erasmus Darwin in his poem (see below).[23] Jan-Andrew Henderson states that Monboddo was
the first to articulate the theory of natural selection.[24]

In 1792, philosopher Immanuel Kant presented, in his Critique of Judgement, what he referred to as “a daring venture
of reason”, in which “one organic being [is] derived from another organic being, although from one which is
specifically different; e.g., certain water-animals transform themselves gradually into marsh-animals and from these,
after some generations, into land-animals.”[25]
In 1796, Erasmus Darwin published Zoönomia, which suggested "that all warm-blooded animals have arisen from
one living filament ... with the power of acquiring new parts" [26] in response to stimuli, with each round of
improvements being inherited by successive generations. In his 1802 poem Temple of Nature, he described the rise
of life from minute organisms living in the mud to its modern diversity:

First forms minute, unseen by spheric glass,

Move on the mud, or pierce the watery mass;
These, as successive generations bloom,
New powers acquire and larger limbs assume;
Whence countless groups of vegetation spring,
And breathing realms of fin and feet and wing.[27]
Early 19th century

Paleontology and geology

See also: History of paleontology

In 1796, Georges Cuvier published his findings on the differences between living elephants and those found in the
fossil record. His analysis demonstrated that mammoths and mastodons were distinct species different from any
living animal, effectively ending a long-running debate over the possibility of the extinction of a species. [28] William
Smith began the process of ordering rock strata by examining fossils in the layers while he worked on his geologic
map of England. Independently, in 1811, Georges Cuvier and Alexandre Brongniart published an influential study of
the geologic history of the region around Paris, which was based on the stratigraphic succession of layers of rock.
These works helped establish the antiquity of the earth.[29] Cuvier advocated catastrophism to explain the patterns of
extinction and faunal succession revealed by the fossil record.

Knowledge of the fossil record continued to advance rapidly during the first few decades of the 19th century. By the
1840s, the outlines of the geologic timescale were becoming clear, and in 1841 John Phillips named three major
eras, based on their predominant fauna: the Paleozoic, dominated by marine invertebrates and fish, the Mesozoic,
the age of reptiles, and the current Cenozoic age of mammals. This progressive picture of the history of life was
accepted even by conservative English geologists like Adam Sedgwick and William Buckland; however, also like
Cuvier, they attributed the progression to repeated catastrophic episodes of extinction followed by new episodes of
creation.[30] Unlike Cuvier, Buckland and some other advocates natural theology among English geologists made
efforts to explicitly link the last catastrophic episode to the biblical flood. [31]

From 1830 to 1833, Charles Lyell published his multi-volume work Principles of Geology, which advocated a
uniformitarian alternative to the catastrophic theory of geology. Lyell claimed that, rather than being the products of
cataclysmic (and possibly supernatural) events, the geologic features of the earth are better explained as the result of
the same gradual geologic forces observable in the present day — but acting over immensely long periods of time.
Although Lyell opposed evolutionary ideas (even questioning the consensus that the fossil record demonstrates a
true progression), his concept that the earth was shaped by forces working gradually over an extended period, and
the immense age of the earth assumed by his theories, would strongly influence future evolutionary thinkers such
as Charles Darwin.[32]

Lamarckism and transmutation

See also: Lamarckism

Jean-Baptiste Lamarck proposed in his Philosophie Zoologique of 1809 a theory of the transmutation of species.
Lamarck did not believe that all living things shared a common ancestor. Rather he believed that simple forms of life
were created continuously by spontaneous generation. He also believed that an innate life force, which he sometimes
described as a nervous fluid, drove species to become more complex over time, advancing up a linear ladder of
complexity that was related to the great chain of being. Lamarck also recognized that species were adapted to their
environment. He explained this observation by saying that the same nervous fluid driving increasing complexity, also
caused the organs of an animal (or a plant) to change based on the use or disuse of that organ, just as muscles are
affected by exercise. He argued that these changes would be inherited by the next generation and produce slow
adaptation to the environment. It was this secondary mechanism of adaptation through the inheritance of acquired
characteristics that became closely associated with his name and would influence discussions of evolution into the
20th century.[33][34]

A radical British school of comparative anatomy that included the surgeon Robert Knox and the anatomist Robert
Grant was closely in touch with Lamarck's school of French Transformationism, which contained scientists such as
Étienne Geoffroy Saint-Hilaire. Grant developed Lamarck's and Erasmus Darwin's ideas of transmutation and
evolutionism, investigating homology to prove common descent. As a young student Charles Darwin joined Grant in
investigations of the life cycle of marine animals. He also studied geology under professor Robert Jameson who
wrote an anonymous paper in 1826 praising "Mr. Lamarck" for explaining how the higher animals had "evolved" from
the "simplest worms" – this was the first use of the word "evolved" in a modern sense. Jameson's course closed with
lectures on the "Origin of the Species of Animals".[35][36]

The computing pioneer Charles Babbage published his unofficial Ninth Bridgewater Treatise in 1837, putting forward
the thesis that God had the omnipotence and foresight to create as a divine legislator, making laws (or programs)
which then produced species at the appropriate times, rather than continually interfering with ad hoc miracles each
time a new species was required. In 1844 the Scottish publisher Robert Chambers anonymously published an
influential, and extremely controversial book of popular science entitled Vestiges of the Natural History of Creation.
This book proposed an evolutionary scenario for the origins of the solar system and life on earth. It claimed that the
fossil record showed a progressive ascent of animals with current animals being branches off a main line that lead
progressively to humanity. It implied that the transmutations lead to the unfolding of a preordained plan that had been
woven into the laws that governed the universe. In this sense it was less completely materialistic than the ideas of
radicals like Robert Grant, but its implication that humans were just the last step in the ascent of animal life incensed
many conservative thinkers. Both conservatives like Adam Sedgwick, and radical materialists like Thomas Henry
Huxley, who disliked Chambers' implications of preordained progress, were able to find scientific inaccuracies in the
book that they could disparage. However, the high profile of the public debate over Vestiges, with its depiction of
evolution as a progressive process, would greatly influence the perception of Darwin's theory a decade later. [37][38]

Opposition to transmutation

Ideas about the transmutation of species were strongly associated with the radical materialism of the enlightenment
and were greeted with hostility by more conservative thinkers. Cuvier attacked the ideas of Lamarck and Geoffroy
Saint-Hilaire, agreeing with Aristotle that species were immutable. Cuvier believed that the individual parts of an
animal were too closely correlated with one another to allow for one part of the anatomy to change in isolation from
the others, and argued that the fossil record showed patterns of catastrophic extinctions followed by re-population,
rather than gradual change over time. He also noted that drawings of animals and animal mummies from Egypt,
which were thousands of years old, showed no signs of change when compared with modern animals. The strength
of Cuvier's arguments and his reputation as a leading scientist helped keep transmutational ideas out of the scientific
mainstream for decades.[39]

In Britain, where the philosophy of natural theology remained influential, William Paley wrote the book Natural
Theology with its famous watchmaker analogy, at least in part as a response to the transmutational ideas of Erasmus
Darwin.[40] Geologists influenced by natural theology, such as Buckland and Sedgwick, made a regular practice of
attacking the evolutionary ideas of Lamarck and Grant, and Sedgwick wrote a famously harsh review of The Vestiges
of the Natural History of Creation.[41][42] Although the geologist Charles Lyell opposed scriptural geology he also
believed in the immutability of species, and in his Principles of Geology (1830–1833), criticized and dismissed
Lamarck's theories of development. Instead, he advocated a form of progressive creation, in which each species had
its "centre of creation" and was designed for this particular habitat, but would go extinct when this habitat changed. [32]

Another source of opposition to transmutation was a school of naturalists who were influenced by the German
philosophers and naturalists associated with idealism, such as Goethe, Hegel and Lorenz Oken. Idealists such
as Louis Agassiz and Richard Owen believed that each species was fixed and unchangeable because it represented
an idea in the mind of the creator. They believed that relationships between species could be discerned from
developmental patterns in embryology, as well as in the fossil record: but that these relationships represented an
underlying pattern of divine thought, with progressive creation leading to increasing complexity and culminating in
humanity. Owen developed the idea of "archetypes" in the Divine mind that would produce a sequence of species
related by anatomical homologies, such as vertebrate limbs. Owen was concerned by the political implications of the
ideas of transmutationists like Robert Grant, and he led a public campaign by conservatives that successfully
marginalized Grant in the scientific community. In his famous 1841 paper, which coined the term dinosaur for the
giant reptiles discovered by Buckland and Gideon Mantell, Owen argued that these reptiles contradicted the
transmutational ideas of Lamarck because they were more sophisticated than the reptiles of the modern world.
Darwin would make good use of the homologies analyzed by Owen in his own theory, but the harsh treatment of
Grant, along with the controversy surrounding Vestiges, would be major factors in his decision to delay publishing his
ideas.[36][43]

Anticipations of natural selection

Several writers anticipated aspects of Darwin's theory, and in the third edition of On the Origin of Species published
in 1861 he named those he had learnt about in an introductory appendix, An Historical Sketch of the Recent
Progress of Opinion on the Origin of Species, which he added to in later editions.[44]

In 1813, William Charles Wells read before the Royal Society essays assuming that there had been evolution of
humans, and recognising the principle of natural selection. Charles Darwin and Alfred Russel Wallace were unaware
of this work when they jointly published the theory in 1858, but Darwin later acknowledged that Wells had recognised
the principle before them, writing that the paper "An Account of a White Female, part of whose Skin resembles that of
a Negro" was published in 1818, and "he distinctly recognises the principle of natural selection, and this is the first
recognition which has been indicated; but he applies it only to the races of man, and to certain characters
alone."[45] When Darwin was developing his theory, he was influenced by Augustin de Candolle's natural system of
classification, which laid emphasis on the war between competing species. [46][47]

Patrick Matthew wrote in the obscure book Naval Timber & Arboriculture that was published in 1831 of "continual
balancing of life to circumstance. ... [The] progeny of the same parents, under great differences of circumstance,
might, in several generations, even become distinct species, incapable of co-reproduction." [48] Charles Darwin
discovered this work after the initial publication of the Origin. In the brief historical sketch that Darwin included in the
3rd addition he says "Unfortunately the view was given by Mr. Matthew very briefly in an Appendix to a work on a
different subject ... He clearly saw, however, the full force of the principle of natural selection." [49]

It is important to understand that it is possible to look through the history of biology from the ancient Greeks onwards
and discover anticipations of almost all of Darwin's key ideas. However, there are at least three major differences
between Darwin and his predecessors. Perhaps the most important difference is that the vast majority of Darwin's
predecessors seem to have failed to understand the implications of their own ideas. As an example, Matthew chose
to relegate his idea on natural selection to the appendix of a work on an unrelated subject, and William Charles Wells
seems to have made little or no attempts to publicise his ideas beyond reading them to the Royal Society. Secondly,
despite having enunciated the basic idea of natural selection, precursors of Darwin either assumed that it was self-
evidently true, or gave merely logical arguments for its importance and failed to provide any empirical data. In other
words, the anticipations of Darwin were merely formal and verbal. Finally, there is Darwin's naturalism. As numerous
Arabic and Muslim authors have pointed out, earlier Islamic scientists such as al-Jahiz did not only enunciate an early
theory of natural selection, but also provided some empirical evidence for it. But these theories were placed in an
Islamic context, and presupposed a Deity. Darwin's theory (at least in its fully developed form) was a secular theory.
[11][50]

T. H. Huxley pointed out in his essay on the reception of the Origin of Species:

The suggestion that new species may result from the selective action of external conditions upon the variations from
their specific type which individuals present and which we call spontaneous because we are ignorant of their
causation is as wholly unknown to the historian of scientific ideas as it was to biological specialists before 1858. But
that suggestion is the central idea of the Origin of Species, and contains the quintessence of Darwinism.[51]

Natural selection

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

The biogeographical patterns Charles Darwin observed in places such as the Galapagos islands during the voyage of
the Beagle caused him to doubt the fixity of species, and in 1837 Darwin started the first of a series of secret
notebooks on transmutation. Darwin's observations lead him to view transmutation as a process of divergence and
branching, rather than the ladder-like progression envisioned by Lamarck and others. In 1838 he read the new 6th
edition of An Essay on the Principle of Population, written in the late 1700s by Thomas Malthus. Malthus' idea of
population growth leading to a struggle for survival combined with Darwin's knowledge on how breeders selected
traits, led to the inception of Darwin's theory of natural selection. Concerned by the intense controversy raging over
other transmutational ideas, Darwin would develop this idea in private for the next 20 years, sharing it only with a
handful of friendly naturalists through correspondence.[52][53]

Unlike Darwin, Alfred Russel Wallace, influenced by the book Vestiges of the Natural History of Creation, already
believed that transmutation of species occurred when he began his career as a naturalist. By 1855 his
biogeographical observations during his field work in South America and the Malay Archipelago made him confident
enough in a branching pattern of evolution to publish a paper that stated that every species originated in close
proximity to an already existing closely allied species. Once again it was consideration of how the ideas of Malthus
might apply to animal populations that lead Wallace to conclusions very similar to the ones reached by Darwin about
the role of natural selection. In February 1858 Wallace, unaware of Darwin's unpublished ideas, wrote up his
thoughts into an essay and mailed them to Darwin, asking for his opinion. The result was the joint publication of
Darwin's theory of natural selection with Wallace in July. Darwin also began work in earnest on The Origin of
Species, which he would publish in 1859.[54]

1859–1930s: Darwin and after Darwin

See also: reaction to Darwin's theory

While transmutation of species was accepted by a sizable number of scientists before 1859, it was the publication of
Charles Darwin's On the Origin of Species that fundamentally transformed the debate over biological origins. Darwin
argued that his branching version of evolution explained a wealth of facts in biogeography, anatomy, embryology, and
other fields of biology. He also provided the first cogent mechanism by which evolutionary change could persist: his
theory of natural selection.[55]

One of the first and most important naturalists to be convinced by Origin was the British anatomist Thomas Henry
Huxley. Huxley recognized that unlike the earlier transmutational ideas of Lamarck and Vestiges, Darwin's theory
provided a mechanism for evolution without supernatural involvement. Huxley would make advocacy of evolution a
cornerstone of the program of the X-club to reform and professionalize science by displacing natural theology with
methodological naturalism, ending the domination of British natural science by the clergy. By the early 1870s in
English-speaking countries, thanks partly to these efforts, evolution had become the mainstream scientific
explanation for the origin of species.[55] In his campaign for public and scientific acceptance of Darwin's theory, Huxley
made extensive use of new evidence for evolution from paleontology. This included evidence that birds had evolved
from reptiles, including the discovery of Archaeopteryx in Europe, and a number of fossils of primitive birds with teeth
found in North America. Another important line of evidence involved fossils that helped trace the evolution of the
horse from its small five-toed ancestors.[56] However, acceptance of evolution among scientists in non-English
speaking nations such as France, and the countries of southern Europe and Latin America was slower. An exception
to this was Germany, where both August Weismann and Ernst Haeckel championed this idea: with Haeckel using
evolution to challenge the established tradition of metaphysical idealism in German biology, much as Huxley used it
to challenge natural theology in Britain.[57]

Darwin's theory succeeded in profoundly shaking scientific opinion regarding the development of life and resulted in a
small social revolution. However, this theory could not explain several critical components of the evolutionary
process. Namely, Darwin was unable to explain the source of variation in traits within a species, and could not identify
a mechanism that could pass traits faithfully from one generation to the next. Darwin's hypothesis of pangenesis,
while relying in part on the inheritance of acquired characteristics, proved to be useful for statistical models of
evolution that were developed by his cousin Francis Galton and the "biometric" school of evolutionary thought. This
idea was, however, of little use to biologists.

Application of the theory to humans

Charles Darwin was aware of the severe reaction in some parts of the scientific community against the suggestion
made in Vestiges of the Natural History of Creation that humans had arisen from animals by a process of
transmutation. Therefore he almost completely ignored the topic of human evolution in The Origin of Species. Despite
this precaution, the issue featured prominently in the debate that followed the book's publication. For most of the first
half of the 19th century the scientific community believed that, although geology had shown that the earth and life
were very old, human beings had appeared suddenly just a few thousand years before the present. However, a
series of archaeological discoveries in the 1840s and 1850s showed stone tools associated with the remains of
extinct animals. By the early 1860s, as summarized in Charles Lyell's 1863 book Geological Evidences of the
Antiquity of Man, it had become widely accepted that humans had existed during a prehistoric period - which
stretched many thousands of years before the start of written history. This new view of human history was more
compatible than the older one with an evolutionary origin for humanity. On the other hand, there was no fossil
evidence of human evolution known at the time, since the only human fossils discovered prior to the very end of the
19th century were of anatomically modern humans, or of very similar Neanderthals. [58]

Therefore the debate that immediately followed the publication of The Origin of Species centered on the similarities
and differences between humans and modern apes. Richard Owen vigorously defended the traditional classification,
suggested by Carolus Linnaeus and Cuvier, that placed humans in a completely separate order from any of the other
mammals. On the other hand, Huxley sought to demonstrate a close anatomical relationship between humans and
apes. In one very famous incident, Huxley showed that Owen was mistaken in claiming that the brains of gorillas
lacked a structure present in human brains. Huxley summarized his argument in his highly influential 1863
book Evidence as to Man's place in Nature. Another viewpoint was advocated by Charles Lyell and Alfred Russel
Wallace. They agreed that humans shared a common ancestor with apes, but questioned whether any purely
materialistic mechanism could account for some of the differences between humans and apes, especially some
aspects of the human mind.[58]

In 1871 Darwin published The Descent of Man, and Selection in Relation to Sex, which contained his views on
human evolution. Darwin argued that the differences between the human mind and the minds of the higher animals
were a matter of degree rather than of kind. For example, he viewed morality as a natural outgrowth of instincts that
were beneficial to animals living in social groups. He believed that all the differences between humans and apes
could be explained by a combination of the selective pressures resulting from our ancestors moving from the trees to
the plains, and sexual selection. The debate over human origins, and over the degree of human uniqueness would
continue well into the 20th century.[58]

Alternatives to natural selection

Evolution was widely accepted in scientific circles within a few years after the publication of Origin, but the
acceptance of natural selection as its driving mechanism was much less widespread. There were a variety of reasons
for this. Natural selection, with its emphasis on death and competition, did not appeal to many naturalists because
they felt it was immoral and left little room for teleology (purpose) or the concept of "progress" in the development of
life. In addition, some felt that natural selection would be too slow, given the estimates of the age of the earth and sun
(10–100 million years) being made at the time by physicists such as Lord Kelvin. Another objection was that natural
selection could not work because at the time the models for inheritance involved blending of inherited characteristics.
The four major alternatives to natural selection in the late 19th century, were theistic evolution, neo-
Lamarckism, orthogenesis, and saltationism.[59][60]

Theistic evolution

Theistic evolution was the idea that a God intervened in the process of evolution to guide it in such a way that the
living world could still be considered to be designed. Some of its advocates included Asa Gray, George Jackson
Mivart, and the Duke of Argyle. However, this idea rapidly fell out of favor among scientists, as they became more
and more committed to the idea of methodological naturalism and came to believe that direct appeals to supernatural
involvement were scientifically unproductive and a form of special pleading. By 1900 it had completely disappeared
from mainstream scientific discussions, although it continued to be used as a way to reconcile religious belief with
scientific discoveries among non-scientists.[59][60]

Neo-Lamarckism

The term Lamarckism was used for the idea that characteristics acquired during the course of an organism's life,
such as changes caused by the use or disuse of a particular organ, could be inherited by the next generation.
Although Wallace completely rejected the concept in favor of natural selection, Darwin had included it in the first
edition of The Origin of Species as a possible supplemental mechanism of evolution. In the late 19th century the
term neo-Lamarckism came to be associated with the position of naturalists who viewed the inheritance of acquired
characteristics as the most important evolutionary mechanism. Advocates of this position included the British writer
and Darwin critic Samuel Butler, the German biologist Ernst Haeckel, and the American paleontologist Edward
Drinker Cope. They considered Lamarckism to be philosophically superior to Darwin's idea of selection acting on
random variation. Butler and Cope both believed that this allowed organisms to effectively drive their own evolution,
since organisms that developed new behaviors would change the patterns of use of their organs and thus kick-start
the evolutionary process. In addition, Cope and Haeckel both believed that evolution was a progressive process.
Cope looked for, and thought he found, patterns of linear progression in the fossil record. The idea of linear progress
was also an important part of Haeckel's recapitulation theory of evolution, which held that the embryological
development of an organism repeats its evolutionary history.[59][60]

Critics of neo-Lamarckism pointed out that no one had ever produced solid evidence for the inheritance of acquired
characteristics. The experimental work of the German biologist August Weismann resulted in the germ plasm
theory of inheritance. This led him to declare that inheritance of acquired charateristics was flatly impossible, since
the Weismann barrier would prevent any changes that occurred to the body after birth from being inherited by the
next generation. Despite these criticisms, neo-Lamarckism remained the most popular alternative to natural selection
at the end of the 19th century, and would remain the position of some naturalists well into the 20th century. [59][60]

Orthogenesis

Orthogenesis or orthogenetic evolution was the hypothesis that life has an innate tendency to change, in a unilinear
fashion, towards ever-greater perfection. It had a significant following in the 19th century, and its proponents included
the Russian biologist Leo Berg, and the American paleontologist Henry Fairfield Osborn. Orthogenesis was
particularly popular among some paleontologists, who believed that the fossil record showed a gradual and constant
unidirectional change. Those who accepted this idea, however, did not necessarily accept that the mechanism driving
orthogenesis was teleological (goal-directed). The orthogenesis hypothesis began to collapse when it became clear
that it could not explain the patterns found by paleontologists in the fossil record, which were non-linear and
contained many complications. A few hung on to the orthogenesis hypothesis as late as the 1950s by claiming that
the processes of macroevolution, the long term trends in evolution, were distinct from the processes of
microevolution.[59][60]

Saltationism

Saltationism was the idea that new species arise as a result of large mutations. It was seen as a much faster
alternative to the Darwinian concept of a gradual process of small random variations being acted on by natural
selection. It was very popular with early geneticists such as Hugo DeVries, William Bateson, and early in his
career, T. H. Morgan. This later became the basis of the mutation theory of evolution. [59][60]

Mendelian genetics, biometrics, and mutation

The rediscovery of Gregor Mendel's laws of inheritance in 1900 ignited a fierce debate between two camps of
biologists. In one camp were the Mendelians, who were focused on discrete variations and the laws of inheritance.
They were led by William Bateson (who coined the word genetics) and Hugo de Vries (who coined the
word mutation). Their opponents were the biometricians, who were interested in the continuous variation of
characteristics within populations. Their leaders, Karl Pearson and Walter Frank Raphael Weldon, followed in the
tradition of Francis Galton - who had focused on measurement and statistical analysis of variation within a population.
The biometricians rejected Mendelian genetics because they felt that discrete units of heredity, such as genes, could
not explain the continuous range of variation seen in wild populations. Weldon's work with crabs and snails provided
evidence that selection pressure from environmental factors really could shift the range of variation in real world
populations, but the Mendelians maintained that the variations measured by biometricians were too insignificant to
account for the evolution of new species.[61][62]

When T. H. Morgan began experimenting with breeding the fruit fly Drosophila melanogaster he was a saltationist
who hoped to demonstrate that a new species could be created in the lab by mutation alone. Instead, the work at his
lab between 1910 and 1915 reconfirmed Mendelian genetics and provided solid experimental evidence linking it to
chromosomal inheritance. It also demonstrated that most mutations had relatively small affects (such as a change in
eye color), and that rather than creating a new species in a single step, they served to increase variation within the
existing population.[61][62]

1920s–1940s:
See also: Modern evolutionary synthesis
Biston betularia f. typica is the white-bodied form of the peppered moth.
Biston betularia f. carbonaria is the black-bodied form of the peppered moth.

Population genetics

Eventually, the Mendelian and biometrician models were reconciled and merged during the development of the
discipline of population genetics. A key part of this development was the work of the British biologist and
statistician R.A. Fisher. In a series of papers starting in 1918 and culminating in his 1930 book Genetical Theory of
Natural Selection Fisher showed that the continuous variation measured by the biometricians could be produced by
the combined action of many discrete genes, and that natural selection could change gene frequencies in a
population, driving evolution. In a series of papers starting in 1924 another British geneticist, J.B.S. Haldane, applied
statistical analysis to real-world examples of natural selection, such as the evolution of industrial melanism in
peppered moths, and showed that natural selection worked at a faster rate than even Fisher had thought possible.
The American biologist Sewall Wright, who had a background in animal breeding experiments, focused on
combinations of interacting genes, and the effects of inbreeding on small relatively isolated populations that
exhibited genetic drift. In 1932 Wright introduced the concept of an adaptive landscape and argued that genetic drift
and inbreeding could drive a small isolated sub-populations away from an adaptive peak, allowing natural selection to
drive it towards different adaptive peaks. The work of Fisher, Haldane, and Wright founded the discipline
of population genetics, which integrated natural selection with Mendelian genetics. [63][64]

Modern evolutionary synthesis

In the first couple of decades of the 20th century most field naturalists continued to believe that Lamarckian and
orthogenic mechanisms of evolution provided the best explanation for the complexity they observed in the living
world. However, as the field of genetics continued to develop, those views became less tenable. Theodosius
Dobzhansky had been a postdoctoral worker in T. H. Morgan's lab and had been influenced by the work on genetic
diversity done by Russian geneticists such as Sergei Chetverikov. He would help to bridge the divide between the
population geneticists and the field biologists with his 1937 book Genetics and the origin of species. Dobzhansky
examined the genetic diversity of wild populations, and showed that contrary to the assumptions of the population
geneticists, these populations had large amounts of genetic diversity with marked differences between sub-
populations. The book also took the highly mathematical work of the population geneticists and put it into more
accessible form. Ernst Mayr was influenced by the work of the German biologist Bernhard Rensch on how local
environmental factors influenced the geographic distribution of sub-species and closely related species. Mayr
followed up on Dobzhansky's work with the 1942 book Systematics and the Origin of Species, which emphasized the
importance of allopatric speciation in the formation of new species. This form of speciation occurs when geographical
isolation of a sub-population is followed by the development of mechanisms for reproductive isolation. Mayr also
formulated the biological species concept that defined a species as a group of interbreeding or potentially
interbreeding populations that were reproductively isolated from all other populations. In the 1944 book Mode and
Tempo in Evolution George Gaylord Simpson showed that the fossil record was consistent with the irregular non-
directional pattern predicted by the developing evolutionary synthesis, and that the linear trends that earlier
paleontologists had claimed supported orthogenesis and neo-Lamarckism did not hold up to closer examination. In
1950 G. Ledyard Stebbins published Variation and Evolution in Plants, which helped integrate botany into the
synthesis. The emerging cross-discipline consensus on how evolution worked would be known as the modern
evolutionary synthesis. It received its name from the book Evolution: the modern synthesis by Julian Huxley.[63][64][65]

1940s–1960s: Molecular biology

Main article: History of molecular evolution

In the 1940s, following up on Griffith's experiment on bacterial transformation, Avery, MacLeod and McCarty
definitively identified deoxyribonucleic acid (DNA) as the transforming principle responsible for transmitting genetic
information.[66] In 1953, Francis Crick and James D. Watson published their famous paper on the structure of DNA,
based on the research of Rosalind Franklin and Maurice Wilkins.[67] These developments ignited the era of molecular
biology and transformed the understanding of evolution into a molecular process: the mutation of segments of DNA.

During this era of molecular biology, it also became clear that a major mechanism for variation within a population
is mutations of DNA. In the mid-1970s, Motoo Kimura formulated the neutral theory of molecular evolution, firmly
establishing the importance of genetic drift as a major mechanism of evolution.[68] The theory sparked the "neutralist-
selectionist" debate, partially solved by the development of Tomoko Ohta's nearly neutral theory of evolution.[69]

Since the 1960s:

Gene centered view of evolution

In the mid-1960s, George C. Williams strongly critiqued verbal explanations of adaptations couched in terms of
"survival of the species" (essentially group selection arguments). Such explanations were largely replaced by a gene-
centered view of evolution, epitomised by the kin selection arguments of W. D. Hamilton, George R. Price and John
Maynard Smith.[70] This viewpoint would be summarized and popularized in the influential 1976 book The Selfish
Gene by Richard Dawkins.[71] Models of the period showed that group selection was severely limited in its strength,
although these models have since been shown to be too limited and newer models do admit the possibility of
significant multi-level selection.[72]

In 1973 Leigh Van Valen proposed the term Red Queen, which he took from Through the Looking Glass by Lewis
Carroll, to describe a scenario where a species involved in one or more evolutionary arms races would have to
constantly change just to keep pace with the species it was co-evolving with. Hamilton, Williams and others
suggested that this idea might help explain the evolution of sexual reproduction, because the increased genetic
diversity caused by sexual reproduction would help maintain resistance against rapidly evolving parasites. They felt
this might explain why sexual reproduction was so common despite the tremendous cost from the gene-centric point
of view of a system where only half of an organism's genome is passed on during reproduction.[73][74] The gene-centric
view has also led to an increased interest in Darwin's old idea of sexual selection,[75] and more recently in topics such
as sexual conflict and intragenomic conflict.

Punctuated equilibrium

One of the most prominent debates arising during this time period was over the theory of punctuated equilibrium, a
theory propounded by Niles Eldredge and Stephen Jay Gould to account for the pattern of fossil species persisting
phenotypically unchanged for long periods (what they termed stasis), with relatively brief periods of phenotypic
change during speciation.[76][77]

Sociobiology

W. D. Hamilton's work on kin selection also contributed to the emergence of the discipline of
sociobiology. Altruism has been a difficult problem for evolutionary theorists going all the way back to Darwin.
[78]
Significant progress was made in 1964 when Hamilton formulated the inequality known as Hamilton's rule which
showed how eusociality (sterile worker classes) in insects and many other examples of altruistic behavior could have
evolved through kin selection. Other theories, some derived from game theory, such as reciprocal altruism followed.
[79]
In 1975 E.O. Wilson published the influential and highly controversial book Sociobiology: The New
Synthesis which claimed evolutionary theory could help explain many aspects of animal, including human, behavior.
Critics of sociobiology, including Stephen Jay Gould, and Richard Lewontin, claimed that sociobiology greatly
overstated the degree to which complex human behaviors could be determined by genetic factors. They also claimed
that the theories of sociobiologists often reflected their own ideological biases. Despite these criticisms, work in
sociobiology and the related discipline of evolutionary psychology, including work on other aspects of the altruism
problem, has continued.[80][81]

Evolutionary paths and processes

Improvements in sequencing methods have resulted in a large increase of sequenced genomes, allowing the testing
and refining of the theory of evolution using this huge amount of genome data. [82] This research is providing insights
into the molecular mechanisms of speciation and adaptation.[83][84] Such genetic analysis has produced fundamental
changes, such as Carl Woese's Three-domain system, in our understanding of the evolutionary history of life.
[85]
Advances in computational hardware and software have allowed for the testing and extrapolation of increasingly
advanced evolutionary models and the development of the field of systems biology.[86] Discoveries
in biotechnology are now producing methods for the synthesis and modification of entire genomes, driving
evolutionary studies to the level where future experiments may involve the creation of entirely synthetic organisms. [87]

Microbiology and horizontal gene transfer

Main article: Horizontal gene transfer

Microbiology has just recently developed into an evolutionary discipline. It was originally ignored due to the paucity of
morphological traits and the lack of a species concept in microbiology, particularly amongst prokaryotes.[88] Now,
evolutionary researchers are taking advantage their improved understanding of microbial physiology and ecology,
produced by the comparative ease of microbial genomics, to explore the taxonomy and evolution of these organisms.
[89]
These studies are revealing completely unanticipated levels of diversity amongst microbes, demonstrating that
these organisms are the dominant form of life on Earth.[90][91]

One particularly important outcome from studies on microbial evolution was the discovery in Japan of horizontal gene
transfer in 1959.[92] This transfer of genetic material between different species of bacteria has played a major role in
the propagation of antibiotic resistance.[93] More recently, as knowledge of genomes has continued to expand, it has
been suggested that lateral transfer of genetic material has played an important role in the evolution of all organisms.
[94]
Indeed, as part of the endosymbiotic theory for the origin of organelles, horizontal gene transfer has been a critical
step in the evolution of eukaryotes such as fungi, plants, and animals. [95][96]

Evo-devo

Main article: Evolutionary developmental biology

In the 1980s and 1990s the tenets of the modern evolutionary synthesis came under increasing scrutiny. There was a
renewal of structuralist themes in evolutionary biology in the work of biologists such as Brian Goodwin and Stuart
Kauffman, which incorporated ideas from cybernetics and systems theory, and emphasized the self-organizing
processes of development as factors directing the course of evolution. The evolutionary biologist Stephen Jay
Gould revived earlier ideas of heterochrony, alterations in the relative rates of developmental processes over the
course of evolution, to account for the generation of novel forms, and, with the evolutionary biologist Richard
Lewontin, wrote an influential paper in 1979 suggesting that a change in one biological structure could arise
incidentally as an accidental result of selection on another structure, rather than through direct selection for that
particular adaptation.[97]

Molecular data regarding the mechanisms underlying development accumulated rapidly during the 1980s and '90s.
For example, it became clear that the diversity of animal morphology was not the result of different sets of proteins
regulating the development of different animals, but from changes in the deployment of a small set of proteins that
were common to all animals.[98] These proteins became known as the "developmental toolkit".[99] These various
perspectives came to inform the disciplines of phylogenetics, paleontology and comparative developmental biology,
spawning the new discipline of "evo-devo."[100]

More recent work in this field has emphasized phenotypic and developmental plasticity. It has been hypothesized, for
example, that the rapid emergence of basic metazoan body plans in the Cambrian Explosion was due in part to
changes in the environment acting on inherent material properties of cell aggregates, such as differential cell
adhesion and biochemical oscillation. The resulting forms were later “locked in” by means of stabilizing natural
selection.[101] Experimental and theoretical research on these and related ideas has been presented in the multi-
authored volume Origination of Organismal Form.

Unconventional evolutionary thought

Gaia hypothesis
Main article: Gaia hypothesis
Pierre Teilhard de Chardin formulated theories describing the gradual development of the Universe from subatomic
particles to human society, considered by Teilhard as the last stage (see Gaia theory), but his ideas were not
accepted by the scientific community. However, this hypothesis was later developed in a more limited and rigorous
form by James Lovelock, who proposed that the living and nonliving parts of Earth can be viewed as a complex
interacting system with similarity to a single organism.[102] This modified hypothesis postulates that all living things
have a regulatory effect on the Earth's environment that promotes life overall. Although not fully accepted by the
scientific community, this hypothesis has been a useful spur to further research and is a topic of current scientific
debate.[103][104]

Notes
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3. ^ (Mayr 1982 p. 304)
4. ^ a b c d Johnston, Ian (1999). Section Three: The Origins of Evolutionary Theory. . . . And Still We Evolve, A
Handbook on the History of Modern Science. Liberal Studies Department, Malaspina University College.
Retrieved on 2007-08-11.
5. ^ a b c (Singer 1931)
6. ^ (Needham and Ronan 1995 p. 101)
7. ^ (Darlington 1959)
8. ^ Fancher, Lynn. Aristotle and the Great Chain. College of DuPage. Retrieved on 2007-08-10.
9. ^ (Draper 1878 pp. 154–155, 237)
10. ^ Conway Zirkle (1941). Natural Selection before the "Origin of Species", Proceedings of the American
Philosophical Society 84 (1), p. 71–123.
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Islamic Quarterly. London. [1]
12. ^ Muhammad Hamidullah and Afzal Iqbal (1993), The Emergence of Islam: Lectures on the Development of
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21. ^ (Larson 2004 pp. 14–15)
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23. ^ (Darwin, Erasmus 1825 p. 9)
24. ^ (Henderson 2000)
25. ^ (Kant 1792 Section 80)
26. ^ (Darwin, Erasmus 1818 Vol I section XXXIX)
27. ^ (Darwin, Erasmus 1825 p. 15)
28. ^ (Larson 2004 p. 7)
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53. ^ (Larson 2004 pp. 55–71)
54. ^ (Bowler 2003 pp. 173–176)
55. ^ a b (Larson 2004 pp. 79–111)
56. ^ (Larson 2004 pp. 139–40)
57. ^ (Larson 2004 pp. 109–110)
58. ^ a b c (Bowler 2003 pp. 207–216)
59. ^ a b c d e f (Larson 2004 pp. 105–129)
60. ^ a b c d e f (Bowler 2003 pp. 196–253)
61. ^ a b (Bowler 2003 pp. 256–273)
62. ^ a b (Larson 2004 pp. 153–174)
63. ^ a b (Bowler 2003 pp. 325–339)
64. ^ a b (Larson 2004 pp. 221–243)
65. ^ Mayr and Provine (1998) pp. 33–34, 297–298, 416
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transformation of pneumococcal types. Inductions of transformation by a desoxyribonucleic acid fraction
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74. ^ (Bowler 2003 p. 358)
75. ^ (Bowler 2003 pp. 358–359)
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See also
 The Voyage of the Beagle
 Galápagos Islands
 Faith and rationality
 List of paradigm shifts in science

This article incorporates text from the Encyclopædia Britannica Eleventh Edition, a publication now in the public
domain.

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