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''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''. In the 1872 sixth edition, "On" was omitted, so the full title is ''The origin of species by means of natural selection, or the preservation of favoured races in the struggle for life.'' This edition is usually known as ''The Origin of Species.'' The 6th is Darwin's final edition; there were minor modifications in the text of certain subsequent issues. See Freeman, R. B.

In Van Wyhe, John, ed. ''Darwin Online: On the Origin of Species'', 2002.
published on 24 November 1859, is a work of
scientific literature : ''For a broader class of literature, see Academic publishing.'' Scientific literature comprises scholarly publications that report original empirical and theoretical work in the natural and social sciences. Within an academic field, scie ...
by
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
that is considered to be the foundation of
evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life ...
. Darwin's book introduced the
scientific theory A scientific theory is an explanation of an aspect of the natural world and universe that has been repeatedly tested and corroborated in accordance with the scientific method, using accepted protocols of observation, measurement, and evaluati ...
that populations evolve over the course of generations through a process of
natural selection 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 heritable traits characteristic of a population over generations. Cha ...
. The book presented a body of evidence that the diversity of life arose by
common descent Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal com ...
through a branching pattern of evolution. Darwin included evidence that he had collected on the ''Beagle'' expedition in the 1830s and his subsequent findings from research, correspondence, and experimentation. Various evolutionary ideas had already been proposed to explain new findings in biology. There was growing support for such ideas among dissident anatomists and the general public, but during the first half of the 19th century the English scientific establishment was closely tied to the
Church of England The Church of England (C of E) is the established Christian church in England and the mother church of the international Anglican Communion. It traces its history to the Christian church recorded as existing in the Roman province of Brit ...
, while science was part of
natural theology Natural theology, once also termed physico-theology, is a type of theology that seeks to provide arguments for theological topics (such as the existence of a deity) based on reason and the discoveries of science. This distinguishes it from ...
. Ideas about the
transmutation of species Transmutation of species and transformism are unproven 18th and 19th-century evolutionary ideas about the change of one species into another that preceded Charles Darwin's theory of natural selection. The French ''Transformisme'' was a term used ...
were controversial as they conflicted with the beliefs that species were unchanging parts of a designed hierarchy and that humans were unique, unrelated to other animals. The political and theological implications were intensely debated, but transmutation was not accepted by the scientific mainstream. The book was written for non-specialist readers and attracted widespread interest upon its publication. Darwin was already highly regarded as a scientist, so his findings were taken seriously and the evidence he presented generated scientific, philosophical, and religious discussion. The debate over the book contributed to the campaign by T. H. Huxley and his fellow members of the
X Club The X Club was a dining club of nine men who supported the theories of natural selection and academic liberalism in late 19th-century England. Thomas Henry Huxley was the initiator; he called the first meeting for 3 November 1864. The club m ...
to secularise science by promoting scientific naturalism. Within two decades, there was widespread scientific agreement that evolution, with a branching pattern of common descent, had occurred, but scientists were slow to give natural selection the significance that Darwin thought appropriate. During "
the eclipse of Darwinism Julian Huxley used the phrase "the eclipse of Darwinism" to describe the state of affairs prior to what he called the "modern synthesis". During the "eclipse", evolution was widely accepted in scientific circles but relatively few biologists be ...
" from the 1880s to the 1930s, various other mechanisms of evolution were given more credit. With the development of the modern evolutionary synthesis in the 1930s and 1940s, Darwin's concept of evolutionary
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
through natural selection became central to modern evolutionary theory, and it has now become the unifying concept of the
life science Life is a quality that distinguishes matter that has biological processes, such as signaling and self-sustaining processes, from that which does not, and is defined by the capacity for growth, reaction to stimuli, metabolism, energy ...
s.


Summary of Darwin's theory

Darwin's theory of evolution is based on key
fact A fact is a datum about one or more aspects of a circumstance, which, if accepted as true and proven true, allows a logical conclusion to be reached on a true–false evaluation. Standard reference works are often used to check facts. Scie ...
s and the
inference Inferences are steps in reasoning, moving from premises to logical consequences; etymologically, the word ''wikt:infer, infer'' means to "carry forward". Inference is theoretically traditionally divided into deductive reasoning, deduction and in ...
s drawn from them, which biologist
Ernst Mayr Ernst Walter Mayr (; 5 July 1904 – 3 February 2005) was one of the 20th century's leading evolutionary biologists. He was also a renowned taxonomist, tropical explorer, ornithologist, philosopher of biology, and historian of science. His ...
summarised as follows: :* Every
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
is fertile enough that if all offspring survived to reproduce, the population would grow (fact). :* Despite periodic fluctuations, populations remain roughly the same size (fact). :* Resources such as food are limited and are relatively stable over time (fact). :* A struggle for survival ensues (inference). :* Individuals in a population vary significantly from one another (fact). :* Much of this variation is
heritable Heredity, also called inheritance or biological inheritance, is the passing on of traits from parents to their offspring; either through asexual reproduction or sexual reproduction, the offspring cells or organisms acquire the genetic informa ...
(fact). :* Individuals less suited to the environment are less likely to survive and less likely to reproduce; individuals more suited to the environment are more likely to survive and more likely to reproduce and leave their heritable traits to future generations, which produces the process of
natural selection 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 heritable traits characteristic of a population over generations. Cha ...
(fact). :* This slowly effected process results in populations changing to adapt to their environments, and ultimately, these variations accumulate over time to form new species (inference).


Background


Developments before Darwin's theory

In later editions of the book, Darwin traced evolutionary ideas as far back as
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ...
; the text he cites is a summary by Aristotle of the ideas of the earlier Greek philosopher
Empedocles Empedocles (; grc-gre, Ἐμπεδοκλῆς; , 444–443 BC) was a Greek pre-Socratic philosopher and a native citizen of Akragas, a Greek city in Sicily. Empedocles' philosophy is best known for originating the cosmogonic theory of the ...
. Early
Christian Christians () are people who follow or adhere to Christianity, a monotheistic Abrahamic religion based on the life and teachings of Jesus Christ. The words ''Christ'' and ''Christian'' derive from the Koine Greek title ''Christós'' (Χρι� ...
Church Fathers The Church Fathers, Early Church Fathers, Christian Fathers, or Fathers of the Church were ancient and influential Christian theologians and writers who established the intellectual and doctrinal foundations of Christianity. The historical per ...
and
Medieval Europe In the history of Europe, the Middle Ages or medieval period lasted approximately from the late 5th to the late 15th centuries, similar to the post-classical period of global history. It began with the fall of the Western Roman Empire a ...
an scholars interpreted the
Genesis creation narrative The Genesis creation narrative is the creation myth of both Judaism and Christianity. The narrative is made up of two stories, roughly equivalent to the first two chapters of the Book of Genesis. In the first, Elohim (the Hebrew generic word ...
allegorically rather than as a literal historical account; organisms were described by their mythological and heraldic significance as well as by their physical form. Nature was widely believed to be unstable and capricious, with monstrous births from union between species, and
spontaneous generation Spontaneous generation is a superseded scientific theory that held that living creatures could arise from nonliving matter and that such processes were commonplace and regular. It was hypothesized that certain forms, such as fleas, could arise f ...
of life. The
Protestant Reformation The Reformation (alternatively named the Protestant Reformation or the European Reformation) was a major movement within Western Christianity in 16th-century Europe that posed a religious and political challenge to the Catholic Church and i ...
inspired a literal interpretation of the Bible, with concepts of creation that conflicted with the findings of an emerging science seeking explanations congruent with the
mechanical philosophy The mechanical philosophy is a form of natural philosophy which compares the universe to a large-scale mechanism (i.e. a machine). The mechanical philosophy is associated with the scientific revolution of early modern Europe. One of the first expo ...
of
René Descartes René Descartes ( or ; ; Latinized: Renatus Cartesius; 31 March 1596 – 11 February 1650) was a French philosopher, scientist, and mathematician, widely considered a seminal figure in the emergence of modern philosophy and science. Ma ...
and the
empiricism In philosophy, empiricism is an epistemological theory that holds that knowledge or justification comes only or primarily from sensory experience. It is one of several views within epistemology, along with rationalism and skepticism. Empir ...
of the
Baconian method The Baconian method is the investigative method developed by Sir Francis Bacon, one of the founders of modern science, and thus a first formulation of a modern scientific method. The method was put forward in Bacon's book ''Novum Organum'' (1620) ...
. After the turmoil of the
English Civil War The English Civil War (1642–1651) was a series of civil wars and political machinations between Parliamentarians (" Roundheads") and Royalists led by Charles I (" Cavaliers"), mainly over the manner of England's governance and issues of r ...
, the
Royal Society The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society and the United Kingdom's national academy of sciences. The society fulfils a number of roles: promoting science and its benefits, re ...
wanted to show that science did not threaten religious and political stability.
John Ray John Ray FRS (29 November 1627 – 17 January 1705) was a Christian English naturalist widely regarded as one of the earliest of the English parson-naturalists. Until 1670, he wrote his name as John Wray. From then on, he used 'Ray', after ...
developed an influential
natural theology Natural theology, once also termed physico-theology, is a type of theology that seeks to provide arguments for theological topics (such as the existence of a deity) based on reason and the discoveries of science. This distinguishes it from ...
of rational order; in his
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
,
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
were static and fixed, their adaptation and complexity designed by God, and
varieties Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
showed minor differences caused by local conditions. In God's benevolent design, carnivores caused mercifully swift death, but the suffering caused by
parasitism Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
was a puzzling problem. The
biological classification In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are give ...
introduced by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...
in 1735 also viewed species as fixed according to the divine plan. In 1766,
Georges Buffon Georges-Louis Leclerc, Comte de Buffon (; 7 September 1707 – 16 April 1788) was a French naturalist, mathematician, cosmologist, and encyclopédiste. His works influenced the next two generations of naturalists, including two prominent F ...
suggested that some similar species, such as horses and asses, or lions, tigers, and leopards, might be varieties descended from a common ancestor. The Ussher chronology of the 1650s had calculated creation at 4004 BC, but by the 1780s geologists assumed a much older world. Wernerians thought
strata In geology and related fields, a stratum ( : strata) is a layer of rock or sediment characterized by certain lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by visible surfaces known as e ...
were deposits from shrinking seas, but
James Hutton James Hutton (; 3 June O.S.172614 June 1726 New Style. – 26 March 1797) was a Scottish geologist, agriculturalist, chemical manufacturer, naturalist and physician. Often referred to as the father of modern geology, he played a key role ...
proposed a self-maintaining infinite cycle, anticipating
uniformitarianism Uniformitarianism, also known as the Doctrine of Uniformity or the Uniformitarian Principle, is the assumption that the same natural laws and processes that operate in our present-day scientific observations have always operated in the universe in ...
. Charles Darwin's grandfather
Erasmus Darwin Erasmus Robert Darwin (12 December 173118 April 1802) was an English physician. One of the key thinkers of the Midlands Enlightenment, he was also a natural philosopher, physiologist, slave-trade abolitionist, inventor, and poet. His poems ...
outlined a hypothesis of
transmutation of species Transmutation of species and transformism are unproven 18th and 19th-century evolutionary ideas about the change of one species into another that preceded Charles Darwin's theory of natural selection. The French ''Transformisme'' was a term used ...
in the 1790s, and French naturalist
Jean-Baptiste Lamarck Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biolo ...
published a more developed theory in 1809. Both envisaged that spontaneous generation produced simple forms of life that progressively developed greater complexity, adapting to the environment by inheriting changes in adults caused by use or disuse. This process was later called
Lamarckism Lamarckism, also known as Lamarckian inheritance or neo-Lamarckism, is the notion that an organism can pass on to its offspring physical characteristics that the parent organism acquired through use or disuse during its lifetime. It is also calle ...
. Lamarck thought there was an inherent progressive tendency driving organisms continuously towards greater complexity, in parallel but separate lineages with no extinction. Geoffroy contended that embryonic development recapitulated transformations of organisms in past eras when the environment acted on embryos, and that animal structures were determined by a constant plan as demonstrated by homologies.
Georges Cuvier Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French naturalist and zoologist, sometimes referred to as the "founding father of paleontology". Cuvier was a major figure in na ...
strongly disputed such ideas, holding that unrelated, fixed species showed similarities that reflected a design for functional needs. His palæontological work in the 1790s had established the reality of extinction, which he explained by local catastrophes, followed by repopulation of the affected areas by other species. In Britain,
William Paley William Paley (July 174325 May 1805) was an English clergyman, Christian apologist, philosopher, and utilitarian. He is best known for his natural theology exposition of the teleological argument for the existence of God in his work ''Natu ...
's ''Natural Theology'' saw adaptation as evidence of beneficial "design" by the Creator acting through natural laws. All naturalists in the two English universities (
Oxford Oxford () is a city in England. It is the county town and only city of Oxfordshire. In 2020, its population was estimated at 151,584. It is north-west of London, south-east of Birmingham and north-east of Bristol. The city is home to the ...
and
Cambridge Cambridge ( ) is a university city and the county town in Cambridgeshire, England. It is located on the River Cam approximately north of London. As of the 2021 United Kingdom census, the population of Cambridge was 145,700. Cambridge bec ...
) were
Church of England The Church of England (C of E) is the established Christian church in England and the mother church of the international Anglican Communion. It traces its history to the Christian church recorded as existing in the Roman province of Brit ...
clergymen, and science became a search for these laws. Geologists adapted catastrophism to show repeated worldwide annihilation and creation of new fixed species adapted to a changed environment, initially identifying the most recent catastrophe as the
biblical flood The Genesis flood narrative (chapters 6–9 of the Book of Genesis) is the Hebrew version of the universal flood myth. It tells of God's decision to return the universe to its pre- creation state of watery chaos and remake it through the microc ...
. Some anatomists such as Robert Grant were influenced by Lamarck and Geoffroy, but most naturalists regarded their ideas of transmutation as a threat to divinely appointed social order.


Inception of Darwin's theory

Darwin went to
Edinburgh University The University of Edinburgh ( sco, University o Edinburgh, gd, Oilthigh Dhùn Èideann; abbreviated as ''Edin.'' in Post-nominal letters, post-nominals) is a Public university, public research university based in Edinburgh, Scotland. Granted ...
in 1825 to study medicine. In his second year he neglected his medical studies for natural history and spent four months assisting Robert Grant's research into marine invertebrates. Grant revealed his enthusiasm for the transmutation of species, but Darwin rejected it. Starting in 1827, at
Cambridge University , mottoeng = Literal: From here, light and sacred draughts. Non literal: From this place, we gain enlightenment and precious knowledge. , established = , other_name = The Chancellor, Masters and Schola ...
, Darwin learnt science as
natural theology Natural theology, once also termed physico-theology, is a type of theology that seeks to provide arguments for theological topics (such as the existence of a deity) based on reason and the discoveries of science. This distinguishes it from ...
from botanist John Stevens Henslow, and read Paley,
John Herschel Sir John Frederick William Herschel, 1st Baronet (; 7 March 1792 – 11 May 1871) was an English polymath active as a mathematician, astronomer, chemist, inventor, experimental photographer who invented the blueprint and did botanical wo ...
and
Alexander von Humboldt Friedrich Wilhelm Heinrich Alexander von Humboldt (14 September 17696 May 1859) was a German polymath, geographer, naturalist, explorer, and proponent of Romantic philosophy and science. He was the younger brother of the Prussian minister ...
. Filled with zeal for science, he studied catastrophist
geology Geology () is a branch of natural science concerned with Earth and other Astronomical object, astronomical objects, the features or rock (geology), rocks of which it is composed, and the processes by which they change over time. Modern geology ...
with
Adam Sedgwick Adam Sedgwick (; 22 March 1785 – 27 January 1873) was a British geologist and Anglican priest, one of the founders of modern geology. He proposed the Cambrian and Devonian period of the geological timescale. Based on work which he did on ...
. In December 1831, he joined the ''Beagle'' expedition as a gentleman naturalist and geologist. He read
Charles Lyell Sir Charles Lyell, 1st Baronet, (14 November 1797 – 22 February 1875) was a Scottish geologist who demonstrated the power of known natural causes in explaining the earth's history. He is best known as the author of ''Principles of Geolo ...
's ''
Principles of Geology ''Principles of Geology: Being an Attempt to Explain the Former Changes of the Earth's Surface, by Reference to Causes Now in Operation'' is a book by the Scottish geologist Charles Lyell that was first published in 3 volumes from 1830–1833. Ly ...
'' and from the first stop ashore, at St. Jago, found Lyell's uniformitarianism a key to the geological history of landscapes. Darwin discovered fossils resembling huge armadillos, and noted the geographical distribution of modern species in hope of finding their "centre of creation". The three Fuegian missionaries the expedition returned to
Tierra del Fuego Tierra del Fuego (, ; Spanish for "Land of the Fire", rarely also Fireland in English) is an archipelago off the southernmost tip of the South American mainland, across the Strait of Magellan. The archipelago consists of the main island, Isla ...
were friendly and civilised, yet to Darwin their relatives on the island seemed "miserable, degraded savages", and he no longer saw an unbridgeable gap between humans and animals. As the ''Beagle'' neared England in 1836, he noted that species might not be fixed.
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Ow ...
showed that fossils of extinct species Darwin found in South America were allied to living species on the same continent. In March 1837, ornithologist
John Gould John Gould (; 14 September 1804 – 3 February 1881) was an English ornithologist. He published a number of monographs on birds, illustrated by plates produced by his wife, Elizabeth Gould, and several other artists, including Edward Lear, ...
announced that
Darwin's rhea Darwin's rhea or the lesser rhea (''Rhea pennata'') is a large flightless bird, the smaller of the two extant species of rheas. It is found in the Altiplano and Patagonia in South America. Description The lesser rhea stands at tall. Leng ...
was a separate species from the previously described rhea (though their territories overlapped), that mockingbirds collected on the
Galápagos Islands The Galápagos Islands ( Spanish: , , ) are an archipelago of volcanic islands. They are distributed on each side of the equator in the Pacific Ocean, surrounding the centre of the Western Hemisphere, and are part of the Republic of Ecuad ...
represented three separate species each unique to a particular island, and that several distinct birds from those islands were all classified as finches. Darwin began speculating, in a series of notebooks, on the possibility that "one species does change into another" to explain these findings, and around July sketched a
genealogical Genealogy () is the study of families, family history, and the tracing of their lineages. Genealogists use oral interviews, historical records, genetic analysis, and other records to obtain information about a family and to demonstrate kin ...
branching of a single
evolutionary tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
, discarding Lamarck's independent lineages progressing to higher forms. Unconventionally, Darwin asked questions of fancy pigeon and
animal breeders Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
as well as established scientists. At the zoo he had his first sight of an ape, and was profoundly impressed by how human the
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genu ...
seemed. In late September 1838, he started reading
Thomas Malthus Thomas Robert Malthus (; 13/14 February 1766 – 29 December 1834) was an English cleric, scholar and influential economist in the fields of political economy and demography. In his 1798 book ''An Essay on the Principle of Population'', Mal ...
's ''
An Essay on the Principle of Population An, AN, aN, or an may refer to: Businesses and organizations * Airlinair (IATA airline code AN) * Alleanza Nazionale, a former political party in Italy * AnimeNEXT, an annual anime convention located in New Jersey * Anime North, a Canadian ...
'' with its statistical argument that human populations, if unrestrained, breed beyond their means and struggle to survive. Darwin related this to the struggle for existence among wildlife and botanist de Candolle's "warring of the species" in plants; he immediately envisioned "a force like a hundred thousand wedges" pushing well-adapted variations into "gaps in the economy of nature", so that the survivors would pass on their form and abilities, and unfavourable variations would be destroyed. By December 1838, he had noted a similarity between the act of breeders selecting traits and a Malthusian Nature selecting among variants thrown up by "chance" so that "every part of newly acquired structure is fully practical and perfected". Darwin now had the basic framework of his theory of natural selection, but he was fully occupied with his career as a geologist and held back from compiling it until his book on ''
The Structure and Distribution of Coral Reefs ''The Structure and Distribution of Coral Reefs, Being the first part of the geology of the voyage of the Beagle, under the command of Capt. Fitzroy, R.N. during the years 1832 to 1836'', was published in 1842 as Charles Darwin's first monogr ...
'' was completed. As he recalled in his autobiography, he had "at last got a theory by which to work", but it was only in June 1842 that he allowed himself "the satisfaction of writing a very brief abstract of my theory in pencil".


Further development

Darwin continued to research and extensively revise his theory while focusing on his main work of publishing the scientific results of the ''Beagle'' voyage. He tentatively wrote of his ideas to Lyell in January 1842; then in June he roughed out a 35-page "Pencil Sketch" of his theory. Darwin began correspondence about his theorising with the botanist
Joseph Dalton Hooker Sir Joseph Dalton Hooker (30 June 1817 – 10 December 1911) was a British botanist and explorer in the 19th century. He was a founder of geographical botany and Charles Darwin's closest friend. For twenty years he served as director of ...
in January 1844, and by July had rounded out his "sketch" into a 230-page "Essay", to be expanded with his research results and published if he died prematurely. In November 1844, the anonymously published
popular science ''Popular Science'' (also known as ''PopSci'') is an American digital magazine carrying popular science content, which refers to articles for the general reader on science and technology subjects. ''Popular Science'' has won over 58 awards, incl ...
book ''
Vestiges of the Natural History of Creation ''Vestiges of the Natural History of Creation'' is an 1844 work of speculative natural history and philosophy by Robert Chambers. Published anonymously in England, it brought together various ideas of stellar evolution with the progressive tra ...
'', written by Scottish journalist Robert Chambers, widened public interest in the concept of transmutation of species. ''Vestiges'' used evidence from the fossil record and embryology to support the claim that living things had progressed from the simple to the more complex over time. But it proposed a linear progression rather than the branching common descent theory behind Darwin's work in progress, and it ignored
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
. Darwin read it soon after publication, and scorned its amateurish geology and zoology, but he carefully reviewed his own arguments after leading scientists, including Adam Sedgwick, attacked its morality and scientific errors. ''Vestiges'' had significant influence on public opinion, and the intense debate helped to pave the way for the acceptance of the more scientifically sophisticated ''Origin'' by moving evolutionary speculation into the mainstream. While few naturalists were willing to consider transmutation,
Herbert Spencer Herbert Spencer (27 April 1820 – 8 December 1903) was an English philosopher, psychologist, biologist, anthropologist, and sociologist famous for his hypothesis of social Darwinism. Spencer originated the expression " survival of the f ...
became an active proponent of Lamarckism and progressive development in the 1850s. Hooker was persuaded to take away a copy of the "Essay" in January 1847, and eventually sent a page of notes giving Darwin much-needed feedback. Reminded of his lack of expertise in
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
, Darwin began an eight-year study of
barnacle A barnacle is a type of arthropod constituting the subclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in erosiv ...
s, becoming the leading expert on their classification. Using his theory, he discovered homologies showing that slightly changed body parts served different functions to meet new conditions, and he found an intermediate stage in the evolution of distinct sexes. Darwin's barnacle studies convinced him that variation arose constantly and not just in response to changed circumstances. In 1854, he completed the last part of his ''Beagle''-related writing and began working full-time on evolution. He now realised that the branching pattern of
evolutionary divergence Divergent evolution or divergent selection is the accumulation of differences between closely related populations within a species, leading to speciation. Divergent evolution is typically exhibited when two populations become separated by a geog ...
was explained by natural selection working constantly to improve adaptation. His thinking changed from the view that species formed in isolated populations only, as on islands, to an emphasis on speciation without isolation; that is, he saw increasing specialisation within large stable populations as continuously exploiting new
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s. He conducted empirical research focusing on difficulties with his theory. He studied the developmental and anatomical differences between different breeds of many domestic animals, became actively involved in fancy pigeon breeding, and experimented (with the help of his son
Francis Francis may refer to: People *Pope Francis, the head of the Catholic Church and sovereign of the Vatican City State and Bishop of Rome * Francis (given name), including a list of people and fictional characters * Francis (surname) Places *Rural ...
) on ways that plant seeds and animals might disperse across oceans to colonise distant islands. By 1856, his theory was much more sophisticated, with a mass of supporting evidence.


Publication


Time taken to publish

In his autobiography, Darwin said he had "gained much by my delay in publishing from about 1839, when the theory was clearly conceived, to 1859; and I lost nothing by it". On the first page of his 1859 book he noted that, having begun work on the topic in 1837, he had drawn up "some short notes" after five years, had enlarged these into a sketch in 1844, and "from that period to the present day I have steadily pursued the same object." Various biographers have proposed that Darwin avoided or delayed making his ideas public for personal reasons. Reasons suggested have included fear of religious persecution or social disgrace if his views were revealed, and concern about upsetting his clergymen naturalist friends or his pious wife Emma.
Charles Darwin's illness For much of his adult life, Charles Darwin's health was repeatedly compromised by an uncommon combination of symptoms, leaving him severely debilitated for long periods of time. However, in some ways this may have helped his work, and Charles Dar ...
caused repeated delays. His paper on Glen Roy had proved embarrassingly wrong, and he may have wanted to be sure he was correct.
David Quammen David Quammen (born February 24, 1948) is an American science, nature, and travel writer and the author of fifteen books. His articles have appeared in ''Outside Magazine'', '' National Geographic'', '' Harper's'', '' Rolling Stone'', ''The New ...
has suggested all these factors may have contributed, and notes Darwin's large output of books and busy family life during that time. A more recent study by science historian
John van Wyhe John van Wyhe (born 1971) is a British historian of science, with a focus on Charles Darwin and Alfred Russel Wallace, at the National University of Singapore. He holds various academic and research positions, ranging from founder and director of ...
has determined that the idea that Darwin delayed publication only dates back to the 1940s, and Darwin's contemporaries thought the time he took was reasonable. Darwin always finished one book before starting another. While he was researching, he told many people about his interest in transmutation without causing outrage. He firmly intended to publish, but it was not until September 1854 that he could work on it full-time. His 1846 estimate that writing his "big book" would take five years proved optimistic.


Events leading to publication: "big book" manuscript

An 1855 paper on the "introduction" of species, written by
Alfred Russel Wallace Alfred Russel Wallace (8 January 1823 – 7 November 1913) was a British natural history, naturalist, explorer, geographer, anthropologist, biologist and illustrator. He is best known for independently conceiving the theory of evolution thro ...
, claimed that patterns in the geographical distribution of living and fossil species could be explained if every new species always came into existence near an already existing, closely related species.
Charles Lyell Sir Charles Lyell, 1st Baronet, (14 November 1797 – 22 February 1875) was a Scottish geologist who demonstrated the power of known natural causes in explaining the earth's history. He is best known as the author of ''Principles of Geolo ...
recognised the implications of Wallace's paper and its possible connection to Darwin's work, although Darwin did not, and in a letter written on 1–2 May 1856 Lyell urged Darwin to publish his theory to establish priority. Darwin was torn between the desire to set out a full and convincing account and the pressure to quickly produce a short paper. He met Lyell, and in correspondence with
Joseph Dalton Hooker Sir Joseph Dalton Hooker (30 June 1817 – 10 December 1911) was a British botanist and explorer in the 19th century. He was a founder of geographical botany and Charles Darwin's closest friend. For twenty years he served as director of ...
affirmed that he did not want to expose his ideas to review by an editor as would have been required to publish in an academic journal. He began a "sketch" account on 14 May 1856, and by July had decided to produce a full technical treatise on species as his "big book" on ''
Natural Selection 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 heritable traits characteristic of a population over generations. Cha ...
''. His theory including the principle of
divergence In vector calculus, divergence is a vector operator that operates on a vector field, producing a scalar field giving the quantity of the vector field's source at each point. More technically, the divergence represents the volume density of ...
was complete by 5 September 1857 when he sent
Asa Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutually excl ...
a brief but detailed abstract of his ideas.


Joint publication of papers by Wallace and Darwin

Darwin was hard at work on the manuscript for his "big book" on ''
Natural Selection 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 heritable traits characteristic of a population over generations. Cha ...
'', when on 18 June 1858 he received a parcel from Wallace, who stayed on the
Maluku Islands The Maluku Islands (; Indonesian: ''Kepulauan Maluku'') or the Moluccas () are an archipelago in the east of Indonesia. Tectonically they are located on the Halmahera Plate within the Molucca Sea Collision Zone. Geographically they are located ...
(
Ternate Ternate is a city in the Indonesian province of North Maluku and an island in the Maluku Islands. It was the ''de facto'' provincial capital of North Maluku before Sofifi on the nearby coast of Halmahera became the capital in 2010. It is off the ...
and Gilolo). It enclosed twenty pages describing an evolutionary mechanism, a response to Darwin's recent encouragement, with a request to send it on to Lyell if Darwin thought it worthwhile. The mechanism was similar to Darwin's own theory. Darwin wrote to Lyell that "your words have come true with a vengeance, ... forestalled" and he would "of course, at once write and offer to send tto any journal" that Wallace chose, adding that "all my originality, whatever it may amount to, will be smashed". Lyell and Hooker agreed that a joint publication putting together Wallace's pages with extracts from Darwin's 1844 Essay and his 1857 letter to Gray should be presented at the
Linnean Society The Linnean Society of London is a learned society dedicated to the study and dissemination of information concerning natural history, evolution, and taxonomy. It possesses several important biological specimen, manuscript and literature colle ...
, and on 1 July 1858, the papers entitled '' On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection'', by Wallace and Darwin respectively, were read out but drew little reaction. While Darwin considered Wallace's idea to be identical to his concept of natural selection, historians have pointed out differences. Darwin described natural selection as being analogous to the
artificial selection Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ma ...
practised by animal breeders, and emphasised competition between individuals; Wallace drew no comparison to
selective breeding Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ...
, and focused on ecological pressures that kept different varieties adapted to local conditions. Some historians have suggested that Wallace was actually discussing
group selection Group selection is a proposed mechanism of evolution in which natural selection acts at the level of the group, instead of at the level of the individual or gene. Early authors such as V. C. Wynne-Edwards and Konrad Lorenz argued that the behavi ...
rather than selection acting on individual variation.


Abstract of Species book

Soon after the meeting, Darwin decided to write "an abstract of my whole work" in the form of one or more papers to be published by the
Linnean Society The Linnean Society of London is a learned society dedicated to the study and dissemination of information concerning natural history, evolution, and taxonomy. It possesses several important biological specimen, manuscript and literature colle ...
, but was concerned about "how it can be made scientific for a Journal, without giving facts, which would be impossible." He asked Hooker how many pages would be available, but "If the Referees were to reject it as not strictly scientific I would, perhaps publish it as pamphlet." He began his "abstract of Species book" on 20 July 1858, while on holiday at
Sandown Sandown is a seaside resort and civil parish on the south-east coast of the Isle of Wight, United Kingdom with the resort of Shanklin to the south and the settlement of Lake in between. Together with Shanklin, Sandown forms a built-up area of ...
, and wrote parts of it from memory, while sending the manuscripts to his friends for checking. By early October, he began to "expect my abstract will run into a small volume, which will have to be published separately." Over the same period, he continued to collect information and write large fully detailed sections of the manuscript for his "big book" on Species, ''
Natural Selection 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 heritable traits characteristic of a population over generations. Cha ...
''.


Murray as publisher; choice of title

By mid-March 1859 Darwin's abstract had reached the stage where he was thinking of early publication; Lyell suggested the publisher John Murray, and met with him to find if he would be willing to publish. On 28 March Darwin wrote to Lyell asking about progress, and offering to give Murray assurances "that my Book is not more ''un''-orthodox, than the subject makes inevitable." He enclosed a draft title sheet proposing ''An abstract of an Essay on the Origin of Species and Varieties Through natural selection'', with the year shown as "1859".
Darwin, C. R
proposed title page for ''Origin of species'' draft
(1859) APS-B-D25.L 38Transcribed by Kees Rookmaaker, edited by John van Wyhe
Murray's response was favourable, and a very pleased Darwin told Lyell on 30 March that he would "send shortly a large bundle of M.S. but unfortunately I cannot for a week, as the three first chapters are in three copyists' hands". He bowed to Murray's objection to "abstract" in the title, though he felt it excused the lack of references, but wanted to keep "natural selection" which was "constantly used in all works on Breeding", and hoped "to retain it with Explanation, somewhat as thus",— ''Through Natural Selection or the preservation of favoured races''. On 31 March Darwin wrote to Murray in confirmation, and listed headings of the 12 chapters in progress: he had drafted all except "XII. Recapitulation & Conclusion". Murray responded immediately with an agreement to publish the book on the same terms as he published Lyell, without even seeing the manuscript: he offered Darwin ⅔ of the profits. Darwin promptly accepted with pleasure, insisting that Murray would be free to withdraw the offer if, having read the chapter manuscripts, he felt the book would not sell well (eventually Murray paid £180 to Darwin for the first edition and by Darwin's death in 1882 the book was in its sixth edition, earning Darwin nearly £3000). On 5 April, Darwin sent Murray the first three chapters, and a proposal for the book's title. An early draft title page suggests ''On the Mutability of Species''. Murray cautiously asked
Whitwell Elwin Whitwell Elwin (26 February 1816 – 1 January 1900) was an English clergyman, critic and editor of the '' Quarterly Review''. Life He was the son of Marsham Elwin, a country gentleman of Thurning, Norfolk, and a descendant of John Rolfe and ...
to review the chapters. At Lyell's suggestion, Elwin recommended that, rather than "put forth the theory without the evidence", the book should focus on observations upon
pigeon Columbidae () is a bird family consisting of doves and pigeons. It is the only family in the order Columbiformes. These are stout-bodied birds with short necks and short slender bills that in some species feature fleshy ceres. They primarily ...
s, briefly stating how these illustrated Darwin's general principles and preparing the way for the larger work expected shortly: "Every body is interested in pigeons." Darwin responded that this was impractical: he had only the last chapter still to write. In September the main title still included "''An essay on the origin of species and varieties''", but Darwin now proposed dropping "varieties". With Murray's persuasion, the title was eventually agreed as ''On the Origin of Species'', with the title page adding ''by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''. In this extended title (and elsewhere in the book) Darwin used the biological term ''" races"'' interchangeably with ''"
varieties Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
"'', meaning varieties within a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
. He used the term broadly,, Quote: "There nonetheless are a few cases in which Darwin does discuss selection processes in which groups are the units, and these will be the focus of the present chapter. But even here it does not matter whether the groups are from different 'races' or from the same race. It is nests of honeybees that compete with each other, and human tribes that compete with other human tribes. For Darwin, the question of group selection had nothing ''special'' to do with 'race.' Still, writing in the heyday of empire, Darwin saw European nations outcompeting the nations, kingdoms, and tribes that occupy the rest of the globe. In this one very salient example, Darwin did see races struggling with each other. In any event, the word ''race'' in Darwin’s subtitle needs to be understood very broadly; it encompasses competition among individuals, competition among groups in the same 'race,' and competition from groups from different 'races.' This is a much broader meaning than the word 'race' tends to have today." and as well as discussions of "the several races, for instance, of the cabbage" and "the hereditary varieties or races of our domestic animals and plants", there are three instances in the book where the phrase "races of man" is used, referring to races of humans.


Publication and subsequent editions

''On the Origin of Species'' was first published on Thursday 24 November 1859, priced at fifteen
shilling The shilling is a historical coin, and the name of a unit of modern currencies formerly used in the United Kingdom, Australia, New Zealand, other British Commonwealth countries and Ireland, where they were generally equivalent to 12 pence ...
s with a first printing of 1250 copies. The book had been offered to booksellers at Murray's autumn sale on Tuesday 22 November, and all available copies had been taken up immediately. In total, 1,250 copies were printed but after deducting presentation and review copies, and five for Stationers' Hall copyright, around 1,170 copies were available for sale. Significantly, 500 were taken by Mudie's Library, ensuring that the book promptly reached a large number of subscribers to the library. The second edition of 3,000 copies was quickly brought out on 7 January 1860, and incorporated numerous corrections as well as a response to religious objections by the addition of a new epigraph on page ii, a quotation from
Charles Kingsley Charles Kingsley (12 June 1819 – 23 January 1875) was a broad church priest of the Church of England, a university professor, social reformer, historian, novelist and poet. He is particularly associated with Christian socialism, the worki ...
, and the phrase "by the Creator" added to the closing sentence. During Darwin's lifetime the book went through six editions, with cumulative changes and revisions to deal with counter-arguments raised. The third edition came out in 1861, with a number of sentences rewritten or added and an introductory appendix, ''An Historical Sketch of the Recent Progress of Opinion on the Origin of Species''. In response to objections that the
origin of life In biology, abiogenesis (from a- 'not' + Greek bios 'life' + genesis 'origin') or the origin of life is the natural process by which life has arisen from non-living matter, such as simple organic compounds. The prevailing scientific hypothes ...
was unexplained, Darwin pointed to acceptance of Newton's law even though the cause of gravity was unknown, and
Leibnitz Leibnitz (Slovenian: ''Lipnica'') is a city in the Austrian state of Styria and on 1 Jan. 2017 had a population of 12,176. It is located to the south of the city of Graz, between the Mur and Sulm rivers. The town is the capital of the Leibni ...
had accused Newton of introducing "occult qualities & miracles".
The fourth edition in 1866 had further revisions. The fifth edition, published on 10 February 1869, incorporated more changes and for the first time included the phrase "
survival of the fittest "Survival of the fittest" is a phrase that originated from Darwinian evolutionary theory as a way of describing the mechanism of natural selection. The biological concept of fitness is defined as reproductive success. In Darwinian terms, ...
", which had been coined by the philosopher
Herbert Spencer Herbert Spencer (27 April 1820 – 8 December 1903) was an English philosopher, psychologist, biologist, anthropologist, and sociologist famous for his hypothesis of social Darwinism. Spencer originated the expression " survival of the f ...
in his ''Principles of Biology'' (1864)."This survival of the fittest, which I have here sought to express in mechanical terms, is that which Mr. Darwin has called 'natural selection', or the preservation of favoured races in the struggle for life." In January 1871,
George Jackson Mivart St. George Jackson Mivart (30 November 1827 – 1 April 1900) was an English biologist. He is famous for starting as an ardent believer in natural selection who later became one of its fiercest critics. Mivart attempted to reconcile D ...
's ''On the Genesis of Species'' listed detailed arguments against natural selection, and claimed it included false
metaphysics Metaphysics is the branch of philosophy that studies the fundamental nature of reality, the first principles of being, identity and change, space and time, causality, necessity, and possibility. It includes questions about the nature of conscio ...
. Darwin made extensive revisions to the sixth edition of the ''Origin'' (this was the first edition in which he used the word "
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
" which had commonly been associated with
embryological development Prenatal development () includes the development of the embryo and of the fetus during a viviparous animal's gestation. Prenatal development starts with fertilization, in the germinal stage of embryonic development, and continues in fetal devel ...
, though all editions concluded with the word "evolved"), and added a new chapter VII, ''Miscellaneous objections'', to address Mivart's arguments. The sixth edition was published by Murray on 19 February 1872 as ''The Origin of Species'', with "On" dropped from the title. Darwin had told Murray of working men in
Lancashire Lancashire ( , ; abbreviated Lancs) is the name of a historic county, ceremonial county, and non-metropolitan county in North West England. The boundaries of these three areas differ significantly. The non-metropolitan county of Lancas ...
clubbing together to buy the fifth edition at 15 shillings and wanted it made more widely available; the price was halved to 7 s 6 d by printing in a smaller
font In metal typesetting, a font is a particular size, weight and style of a typeface. Each font is a matched set of type, with a piece (a " sort") for each glyph. A typeface consists of a range of such fonts that shared an overall design. In mo ...
. It includes a glossary compiled by W.S. Dallas. Book sales increased from 60 to 250 per month.


Publication outside Great Britain

In the United States, botanist
Asa Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutually excl ...
, an American colleague of Darwin, negotiated with a Boston publisher for publication of an authorised American version, but learnt that two New York publishing firms were already planning to exploit the absence of international
copyright A copyright is a type of intellectual property that gives its owner the exclusive right to copy, distribute, adapt, display, and perform a creative work, usually for a limited time. The creative work may be in a literary, artistic, educatio ...
to print ''Origin''. Darwin was delighted by the popularity of the book, and asked Gray to keep any profits. Gray managed to negotiate a 5% royalty with Appleton's of New York, who got their edition out in mid-January 1860, and the other two withdrew. In a May letter, Darwin mentioned a print run of 2,500 copies, but it is not clear if this referred to the first printing only, as there were four that year. The book was widely translated in Darwin's lifetime, but problems arose with translating concepts and metaphors, and some translations were biased by the translator's own agenda. Darwin distributed presentation copies in France and Germany, hoping that suitable applicants would come forward, as translators were expected to make their own arrangements with a local publisher. He welcomed the distinguished elderly naturalist and geologist
Heinrich Georg Bronn Heinrich Georg Bronn (3 March 1800 – 5 July 1862) was a German geologist and paleontologist. He was the first to translate Charles Darwin's ''On the Origin of Species'' into German in 1860, although not without introducing his own interpretation ...
, but the German translation published in 1860 imposed Bronn's own ideas, adding controversial themes that Darwin had deliberately omitted. Bronn translated "favoured races" as "perfected races", and added essays on issues including the origin of life, as well as a final chapter on religious implications partly inspired by Bronn's adherence to ''
Naturphilosophie ''Naturphilosophie'' (German for "nature-philosophy") is a term used in English-language philosophy to identify a current in the philosophical tradition of German idealism, as applied to the study of nature in the earlier 19th century. German s ...
''. In 1862, Bronn produced a second edition based on the third English edition and Darwin's suggested additions, but then died of a heart attack. Darwin corresponded closely with
Julius Victor Carus Julius Victor Carus (25 July 1823 – 10 March 1903) was a German zoologist, comparative anatomist and entomologist. Career Carus was born in Leipzig. He served as curator of the Museum of Comparative Anatomy at Oxford University from 1849 to 1 ...
, who published an improved translation in 1867. Darwin's attempts to find a translator in France fell through, and the translation by
Clémence Royer Clémence Royer (21 April 1830 – 6 February 1902) was a self-taught French scholar who lectured and wrote on economics, philosophy, science and feminism. She is best known for her controversial 1862 French translation of Charles Darwin's ' ...
published in 1862 added an introduction praising Darwin's ideas as an alternative to religious revelation and promoting ideas anticipating
social Darwinism Social Darwinism refers to various theories and societal practices that purport to apply biological concepts of natural selection and survival of the fittest to sociology, economics and politics, and which were largely defined by scholars in We ...
and
eugenics Eugenics ( ; ) is a fringe set of beliefs and practices that aim to improve the genetic quality of a human population. Historically, eugenicists have attempted to alter human gene pools by excluding people and groups judged to be inferior o ...
, as well as numerous explanatory notes giving her own answers to doubts that Darwin expressed. Darwin corresponded with Royer about a second edition published in 1866 and a third in 1870, but he had difficulty getting her to remove her notes and was troubled by these editions. He remained unsatisfied until a translation by Edmond Barbier was published in 1876. A Dutch translation by
Tiberius Cornelis Winkler Tiberius Cornelis Winkler (May 28, 1822 – April 4, 1897) was a Dutch anatomist, zoologist and natural historian, and the second curator of geology, paleontology and mineralogy at Teylers Museum in Haarlem. Besides translating the first edit ...
was published in 1860. By 1864, additional translations had appeared in Italian and Russian. In Darwin's lifetime, ''Origin'' was published in Swedish in 1871, Danish in 1872, Polish in 1873, Hungarian in 1873–1874, Spanish in 1877 and Serbian in 1878. By 1977, ''Origin'' had appeared in an additional 18 languages, including Chinese by Ma Chün-wu who added non-Darwinian ideas; he published the preliminaries and chapters 1–5 in 1902–1904, and his complete translation in 1920.


Content


Title pages and introduction

Page ii contains quotations by
William Whewell William Whewell ( ; 24 May 17946 March 1866) was an English polymath, scientist, Anglican priest, philosopher, theologian, and historian of science. He was Master of Trinity College, Cambridge. In his time as a student there, he achieved ...
and
Francis Bacon Francis Bacon, 1st Viscount St Alban (; 22 January 1561 – 9 April 1626), also known as Lord Verulam, was an English philosopher and statesman who served as Attorney General and Lord Chancellor of England. Bacon led the advancement of both ...
on the theology of natural laws, harmonising science and religion in accordance with
Isaac Newton Sir Isaac Newton (25 December 1642 – 20 March 1726/27) was an English mathematician, physicist, astronomer, alchemist, Theology, theologian, and author (described in his time as a "natural philosophy, natural philosopher"), widely ...
's belief in a rational God who established a law-abiding cosmos. In the second edition, Darwin added an epigraph from
Joseph Butler Joseph Butler (18 May O.S. 1692 – 16 June O.S. 1752) was an English Anglican bishop, theologian, apologist, and philosopher, born in Wantage in the English county of Berkshire (now in Oxfordshire). He is known for critiques of Deism, Thom ...
affirming that God could work through scientific laws as much as through
miracles A miracle is an event that is inexplicable by natural or scientific lawsOne dictionary define"Miracle"as: "A surprising and welcome event that is not explicable by natural or scientific laws and is therefore considered to be the work of a divin ...
, in a nod to the religious concerns of his oldest friends. The ''Introduction'' establishes Darwin's credentials as a naturalist and author, then refers to
John Herschel Sir John Frederick William Herschel, 1st Baronet (; 7 March 1792 – 11 May 1871) was an English polymath active as a mathematician, astronomer, chemist, inventor, experimental photographer who invented the blueprint and did botanical wo ...
's letter suggesting that the origin of species "would be found to be a natural in contradistinction to a miraculous process":
WHEN on board HMS ''Beagle'', as naturalist, I was much struck with certain facts in the distribution of the inhabitants of South America, and in the geological relations of the present to the past inhabitants of that continent. These facts seemed to me to throw some light on the origin of species—that mystery of mysteries, as it has been called by one of our greatest philosophers.
Darwin refers specifically to the distribution of the species
rheas The rheas ( ), also known as ñandus ( ) or South American ostriches, are large ratites (flightless birds without a keel on their sternum bone) in the order Rheiformes, native to South America, distantly related to the ostrich and emu. Most tax ...
, and to that of the
Galápagos tortoise The Galápagos tortoise or Galápagos giant tortoise (''Chelonoidis niger'') is a species of very large tortoise in the genus ''Chelonoidis'' (which also contains three smaller species from mainland South America). It comprises 15 subspecies ...
s and mockingbirds. He mentions his years of work on his theory, and the arrival of Wallace at the same conclusion, which led him to "publish this Abstract" of his incomplete work. He outlines his ideas, and sets out the essence of his theory:
As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be naturally selected. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form.
Starting with the third edition, Darwin prefaced the introduction with a sketch of the historical development of evolutionary ideas. In that sketch he acknowledged that Patrick Matthew had, unknown to Wallace or himself, anticipated the concept of natural selection in an appendix to a book published in 1831; in the fourth edition he mentioned that William Charles Wells had done so as early as 1813.


Variation under domestication and under nature

Chapter I covers
animal husbandry Animal husbandry is the branch of agriculture concerned with animals that are raised for meat, fibre, milk, or other products. It includes day-to-day care, selective breeding, and the raising of livestock. Husbandry has a long history, starti ...
and
plant breeding Plant breeding is the science of changing the traits of plants in order to produce desired characteristics. It has been used to improve the quality of nutrition in products for humans and animals. The goals of plant breeding are to produce cr ...
, going back to ancient Egypt. Darwin discusses contemporary opinions on the origins of different breeds under cultivation to argue that many have been produced from common ancestors by
selective breeding Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ...
. As an illustration of
artificial selection Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ma ...
, he describes fancy pigeon breeding, noting that " e diversity of the breeds is something astonishing", yet all were descended from one species of
rock pigeon The rock dove, rock pigeon, or common pigeon ( also ; ''Columba livia'') is a member of the bird family Columbidae (doves and pigeons). In common usage, it is often simply referred to as the "pigeon". The domestic pigeon (''Columba livia domes ...
. Darwin saw two distinct kinds of variation: (1) rare abrupt changes he called "sports" or "monstrosities" (example:
Ancon sheep Ancon sheep (also called "Otter" sheep) were a grouping of domestic sheep with long bodies and very short legs, with the fore-legs being crooked. The term is generally applied to a line of sheep bred from a single affected lamb born in 1791 in Ma ...
with short legs), and (2) ubiquitous small differences (example: slightly shorter or longer bill of pigeons).David Reznick (2009) ''The Origin Then and Now'', Princeton University Press, p.49. Both types of hereditary changes can be used by breeders. However, for Darwin the small changes were most important in evolution. In this chapter Darwin expresses his erroneous belief that environmental change is necessary to generate variation. In Chapter II, Darwin specifies that the distinction between
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
and
varieties Variety may refer to: Arts and entertainment Entertainment formats * Variety (radio) * Variety show, in theater and television Films * ''Variety'' (1925 film), a German silent film directed by Ewald Andre Dupont * ''Variety'' (1935 film), ...
is arbitrary, with experts disagreeing and changing their decisions when new forms were found. He concludes that "a well-marked variety may be justly called an incipient species" and that "species are only strongly marked and permanent varieties". He argues for the ubiquity of variation in nature. Historians have noted that naturalists had long been aware that the individuals of a species differed from one another, but had generally considered such variations to be limited and unimportant deviations from the
archetype The concept of an archetype (; ) appears in areas relating to behavior, historical psychology, and literary analysis. An archetype can be any of the following: # a statement, pattern of behavior, prototype, "first" form, or a main model that ...
of each species, that archetype being a fixed ideal in the mind of God. Darwin and Wallace made variation among individuals of the same species central to understanding the natural world.


Struggle for existence, natural selection, and divergence

In Chapter III, Darwin asks how varieties "which I have called incipient species" become distinct species, and in answer introduces the key concept he calls "
natural selection 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 heritable traits characteristic of a population over generations. Cha ...
"; in the fifth edition he adds, "But the expression often used by Mr.
Herbert Spencer Herbert Spencer (27 April 1820 – 8 December 1903) was an English philosopher, psychologist, biologist, anthropologist, and sociologist famous for his hypothesis of social Darwinism. Spencer originated the expression " survival of the f ...
, of the
Survival of the Fittest "Survival of the fittest" is a phrase that originated from Darwinian evolutionary theory as a way of describing the mechanism of natural selection. The biological concept of fitness is defined as reproductive success. In Darwinian terms, ...
, is more accurate, and is sometimes equally convenient."
Owing to this struggle for life, any variation, however slight and from whatever cause proceeding, if it be in any degree profitable to an individual of any species, in its infinitely complex relations to other organic beings and to external nature, will tend to the preservation of that individual, and will generally be inherited by its offspring ... I have called this principle, by which each slight variation, if useful, is preserved, by the term of Natural Selection, in order to mark its relation to man's power of selection.
He notes that both A. P. de Candolle and
Charles Lyell Sir Charles Lyell, 1st Baronet, (14 November 1797 – 22 February 1875) was a Scottish geologist who demonstrated the power of known natural causes in explaining the earth's history. He is best known as the author of ''Principles of Geolo ...
had stated that all organisms are exposed to severe competition. Darwin emphasizes that he used the phrase "
struggle for existence The concept of the struggle for existence concerns the competition or battle for resources needed to live. It can refer to human society, or to organisms in nature. The concept is ancient, and the term ''struggle for existence'' was in use by the ...
" in "a large and metaphorical sense, including dependence of one being on another"; he gives examples ranging from plants struggling against drought to plants competing for birds to eat their fruit and disseminate their seeds. He describes the struggle resulting from population growth: "It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms." He discusses checks to such increase including complex
ecological Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...
interdependencies, and notes that competition is most severe between closely related forms "which fill nearly the same place in the economy of nature". Chapter IV details natural selection under the "infinitely complex and close-fitting ... mutual relations of all organic beings to each other and to their physical conditions of life". Darwin takes as an example a country where a change in conditions led to extinction of some species, immigration of others and, where suitable variations occurred, descendants of some species became adapted to new conditions. He remarks that the artificial selection practised by animal breeders frequently produced sharp divergence in character between breeds, and suggests that natural selection might do the same, saying:
But how, it may be asked, can any analogous principle apply in nature? I believe it can and does apply most efficiently, from the simple circumstance that the more diversified the descendants from any one species become in structure, constitution, and habits, by so much will they be better enabled to seize on many and widely diversified places in the polity of nature, and so be enabled to increase in numbers.
Historians have remarked that here Darwin anticipated the modern concept of an
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
. He did not suggest that every favourable variation must be selected, nor that the favoured animals were better or higher, but merely more adapted to their surroundings. Darwin proposes
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (in ...
, driven by competition between males for mates, to explain
sexually dimorphic Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
features such as lion manes, deer antlers, peacock tails, bird songs, and the bright plumage of some male birds. He analysed sexual selection more fully in ''
The Descent of Man, and Selection in Relation to Sex ''The Descent of Man, and Selection in Relation to Sex'' is a book by English natural history, naturalist Charles Darwin, first published in 1871, which applies evolutionary theory to human evolution, and details his theory of sexual selection, ...
'' (1871). Natural selection was expected to work very slowly in forming new species, but given the effectiveness of artificial selection, he could "see no limit to the amount of change, to the beauty and infinite complexity of the coadaptations between all organic beings, one with another and with their physical conditions of life, which may be effected in the long course of time by nature's power of selection". Using a tree diagram and calculations, he indicates the "divergence of character" from original species into new species and genera. He describes branches falling off as extinction occurred, while new branches formed in "the great
Tree of life The tree of life is a fundamental archetype in many of the world's mythological, religious, and philosophical traditions. It is closely related to the concept of the sacred tree.Giovino, Mariana (2007). ''The Assyrian Sacred Tree: A Hist ...
 ... with its ever branching and beautiful ramifications".


Variation and heredity

In Darwin's time there was no agreed-upon model of
heredity Heredity, also called inheritance or biological inheritance, is the passing on of traits from parents to their offspring; either through asexual reproduction or sexual reproduction, the offspring cells or organisms acquire the genetic informa ...
; in Chapter I Darwin admitted, "The laws governing inheritance are quite unknown." He accepted a version of the inheritance of acquired characteristics (which after Darwin's death came to be called
Lamarckism Lamarckism, also known as Lamarckian inheritance or neo-Lamarckism, is the notion that an organism can pass on to its offspring physical characteristics that the parent organism acquired through use or disuse during its lifetime. It is also calle ...
), and Chapter V discusses what he called the effects of use and disuse; he wrote that he thought "there can be little doubt that use in our domestic animals strengthens and enlarges certain parts, and disuse diminishes them; and that such modifications are inherited", and that this also applied in nature. Darwin stated that some changes that were commonly attributed to use and disuse, such as the loss of functional wings in some island-dwelling insects, might be produced by natural selection. In later editions of ''Origin'', Darwin expanded the role attributed to the inheritance of acquired characteristics. Darwin also admitted ignorance of the source of inheritable variations, but speculated they might be produced by environmental factors. However, one thing was clear: whatever the exact nature and causes of new variations, Darwin knew from observation and experiment that breeders were able to select such variations and produce huge differences in many generations of selection. The ''observation'' that selection works in domestic animals is not destroyed by lack of ''understanding'' of the underlying hereditary mechanism. Breeding of animals and plants showed related varieties varying in similar ways, or tending to revert to an ancestral form, and similar patterns of variation in distinct species were explained by Darwin as demonstrating
common descent Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal com ...
. He recounted how
Lord Morton's mare Lord Morton’s mare was an equid hybrid and once an often-noticed example in the history of evolutionary theory. In 1820, George Douglas, 16th Earl of Morton, F.R.S., reported to the President of the Royal Society that, being desirous of dome ...
apparently demonstrated
telegony The ''Telegony'' (Greek: , ''Tēlegoneia''; la, Telegonia) is a lost ancient Greek epic poem about Telegonus, son of Odysseus by Circe. His name ("born far away") is indicative of his birth on Aeaea, far from Odysseus' home of Ithaca. It was p ...
, offspring inheriting characteristics of a previous mate of the female parent, and accepted this process as increasing the variation available for natural selection. More detail was given in Darwin's 1868 book on ''
The Variation of Animals and Plants Under Domestication ''The Variation of Animals and Plants under Domestication'' is a book by Charles Darwin that was first published in January 1868. A large proportion of the book contains detailed information on the domestication of animals and plants but it al ...
'', which tried to explain heredity through his hypothesis of
pangenesis Pangenesis was Charles Darwin's hypothetical mechanism for heredity, in which he proposed that each part of the body continually emitted its own type of small organic particles called gemmules that aggregated in the gonads, contributing herita ...
. Although Darwin had privately questioned
blending inheritance Blending may refer to: * The process of mixing in process engineering * Mixing paints to achieve a greater range of colors * Blending (alcohol production), a technique to produce alcoholic beverages by mixing different brews * Blending (linguisti ...
, he struggled with the theoretical difficulty that novel individual variations would tend to blend into a population. However, inherited variation could be seen, and Darwin's concept of selection working on a population with a range of small variations was workable. It was not until the modern evolutionary synthesis in the 1930s and 1940s that a model of heredity became completely integrated with a model of variation. This modern evolutionary synthesis had been dubbed Neo Darwinian Evolution because it encompasses
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
's theories of evolution with
Gregor Mendel Gregor Johann Mendel, OSA (; cs, Řehoř Jan Mendel; 20 July 1822 – 6 January 1884) was a biologist, meteorologist, mathematician, Augustinian friar and abbot of St. Thomas' Abbey in Brünn (''Brno''), Margraviate of Moravia. Mendel was ...
's theories of genetic inheritance.


Difficulties for the theory

Chapter VI begins by saying the next three chapters will address possible objections to the theory, the first being that often no intermediate forms between closely related species are found, though the theory implies such forms must have existed. As Darwin noted, "Firstly, why, if species have descended from other species by insensibly fine gradations, do we not everywhere see innumerable transitional forms? Why is not all nature in confusion, instead of the species being, as we see them, well defined?" Darwin attributed this to the competition between different forms, combined with the small number of individuals of intermediate forms, often leading to extinction of such forms. Another difficulty, related to the first one, is the absence or rarity of transitional varieties in time. Darwin commented that by the theory of natural selection "innumerable transitional forms must have existed," and wondered "why do we not find them embedded in countless numbers in the crust of the earth?" (For further discussion of these difficulties, see Speciation#Darwin's dilemma: Why do species exist? and Bernstein et al. and Michod.) The chapter then deals with whether natural selection could produce complex specialised structures, and the behaviours to use them, when it would be difficult to imagine how intermediate forms could be functional. Darwin said:
Secondly, is it possible that an animal having, for instance, the structure and habits of a bat, could have been formed by the modification of some animal with wholly different habits? Can we believe that natural selection could produce, on the one hand, organs of trifling importance, such as the tail of a giraffe, which serves as a fly-flapper, and, on the other hand, organs of such wonderful structure, as the eye, of which we hardly as yet fully understand the inimitable perfection?
His answer was that in many cases animals exist with intermediate structures that are functional. He presented
flying squirrels Flying squirrels (scientifically known as Pteromyini or Petauristini) are a tribe of 50 species of squirrels in the family Sciuridae. Despite their name, they are not in fact capable of full flight in the same way as birds or bats, but they ar ...
, and flying lemurs as examples of how bats might have evolved from non-flying ancestors. He discussed various simple eyes found in invertebrates, starting with nothing more than an optic nerve coated with pigment, as examples of how the vertebrate eye could have evolved. Darwin concludes: "If it could be demonstrated that any complex organ existed, which could not possibly have been formed by numerous, successive, slight modifications, my theory would absolutely break down. But I can find out no such case." In a section on "organs of little apparent importance", Darwin discusses the difficulty of explaining various seemingly trivial traits with no evident adaptive function, and outlines some possibilities such as correlation with useful features. He accepts that we "are profoundly ignorant of the causes producing slight and unimportant variations" which distinguish
domesticated Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...
breeds of animals, and
human races A race is a categorization of humans based on shared physical or social qualities into groups generally viewed as distinct within a given society. The term came into common usage during the 1500s, when it was used to refer to groups of variou ...
. He suggests that
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (in ...
might explain these variations:
I might have adduced for this same purpose the differences between the races of man, which are so strongly marked; I may add that some little light can apparently be thrown on the origin of these differences, chiefly through sexual selection of a particular kind, but without here entering on copious details my reasoning would appear frivolous.
Chapter VII (of the first edition) addresses the evolution of instincts. His examples included two he had investigated experimentally:
slave-making ant Slave-making ants are brood parasites that capture broods of other ant species to increase the worker force of their colony. After emerging in the slave-maker nest, slave workers work as if they were in their own colony, while parasite workers o ...
s and the construction of hexagonal cells by honey bees. Darwin noted that some species of slave-making ants were more dependent on slaves than others, and he observed that many ant species will collect and store the pupae of other species as food. He thought it reasonable that species with an extreme dependency on slave workers had evolved in incremental steps. He suggested that bees that make hexagonal cells evolved in steps from bees that made round cells, under pressure from natural selection to economise wax. Darwin concluded:
Finally, it may not be a logical deduction, but to my imagination it is far more satisfactory to look at such instincts as the young cuckoo ejecting its foster-brothers, —ants making slaves, —the larvæ of ichneumonidæ feeding within the live bodies of caterpillars, —not as specially endowed or created instincts, but as small consequences of one general law, leading to the advancement of all organic beings, namely, multiply, vary, let the strongest live and the weakest die.
Chapter VIII addresses the idea that species had special characteristics that prevented hybrids from being fertile in order to preserve separately created species. Darwin said that, far from being constant, the difficulty in producing hybrids of related species, and the viability and fertility of the hybrids, varied greatly, especially among plants. Sometimes what were widely considered to be separate species produced fertile hybrid offspring freely, and in other cases what were considered to be mere varieties of the same species could only be crossed with difficulty. Darwin concluded: "Finally, then, the facts briefly given in this chapter do not seem to me opposed to, but even rather to support the view, that there is no fundamental distinction between species and varieties." In the sixth edition Darwin inserted a new chapter VII (renumbering the subsequent chapters) to respond to criticisms of earlier editions, including the objection that many features of organisms were not adaptive and could not have been produced by natural selection. He said some such features could have been by-products of adaptive changes to other features, and that often features seemed non-adaptive because their function was unknown, as shown by his book on '' Fertilisation of Orchids'' that explained how their elaborate structures facilitated pollination by insects. Much of the chapter responds to
George Jackson Mivart St. George Jackson Mivart (30 November 1827 – 1 April 1900) was an English biologist. He is famous for starting as an ardent believer in natural selection who later became one of its fiercest critics. Mivart attempted to reconcile D ...
's criticisms, including his claim that features such as
baleen Baleen is a filter-feeding system inside the mouths of baleen whales. To use baleen, the whale first opens its mouth underwater to take in water. The whale then pushes the water out, and animals such as krill are filtered by the baleen and ...
filters in whales,
flatfish A flatfish is a member of the ray-finned demersal fish order Pleuronectiformes, also called the Heterosomata, sometimes classified as a suborder of Perciformes. In many species, both eyes lie on one side of the head, one or the other migrating ...
with both eyes on one side and the camouflage of
stick insects The Phasmatodea (also known as Phasmida, Phasmatoptera or Spectra) are an order of insects whose members are variously known as stick insects, stick-bugs, walking sticks, stick animals, or bug sticks. They are also occasionally referred to as ...
could not have evolved through natural selection because intermediate stages would not have been adaptive. Darwin proposed scenarios for the incremental evolution of each feature.


Geological record

Chapter IX deals with the fact that the
geological record The geologic record in stratigraphy, paleontology and other natural sciences refers to the entirety of the layers of rock stratum, strata. That is, deposits laid down by volcanism or by Deposition (sediment), deposition of sediment derived from w ...
appears to show forms of life suddenly arising, without the innumerable
transitional fossil A transitional fossil is any fossilized remains of a life form that exhibits traits common to both an ancestral group and its derived descendant group. This is especially important where the descendant group is sharply differentiated by gross a ...
s expected from gradual changes. Darwin borrowed
Charles Lyell Sir Charles Lyell, 1st Baronet, (14 November 1797 – 22 February 1875) was a Scottish geologist who demonstrated the power of known natural causes in explaining the earth's history. He is best known as the author of ''Principles of Geolo ...
's argument in ''
Principles of Geology ''Principles of Geology: Being an Attempt to Explain the Former Changes of the Earth's Surface, by Reference to Causes Now in Operation'' is a book by the Scottish geologist Charles Lyell that was first published in 3 volumes from 1830–1833. Ly ...
'' that the record is extremely imperfect as
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
isation is a very rare occurrence, spread over vast periods of time; since few areas had been geologically explored, there could only be fragmentary knowledge of
geological formations A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics (lithology) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock expo ...
, and fossil collections were very poor. Evolved local varieties which migrated into a wider area would seem to be the sudden appearance of a new species. Darwin did not expect to be able to reconstruct evolutionary history, but continuing discoveries gave him well-founded hope that new finds would occasionally reveal transitional forms. To show that there had been enough time for natural selection to work slowly, he cited the example of The Weald as discussed in ''Principles of Geology'' together with other observations from
Hugh Miller Hugh Miller (10 October 1802 – 23/24 December 1856) was a self-taught Scottish geologist and writer, folklorist and an evangelical Christian. Life and work Miller was born in Cromarty, the first of three children of Harriet Wright (''b ...
,
James Smith of Jordanhill James Smith of Jordanhill FRSE FRS MWS (1782–1867) was a Scottish merchant, antiquarian, architect, geologist, biblical critic and man of letters. An authority on ancient shipbuilding and navigation, his works included "Newer Pliocene" ...
and Andrew Ramsay. Combining this with an estimate of recent rates of sedimentation and erosion, Darwin calculated that erosion of The Weald had taken around 300 million years. The initial appearance of entire groups of well-developed organisms in the oldest fossil-bearing layers, now known as the
Cambrian explosion The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil record. ...
, posed a problem. Darwin had no doubt that earlier seas had swarmed with living creatures, but stated that he had no satisfactory explanation for the lack of fossils. Fossil evidence of
pre-Cambrian The Precambrian (or Pre-Cambrian, sometimes abbreviated pꞒ, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of the ...
life has since been found, extending the history of life back for billions of years. Chapter X examines whether patterns in the fossil record are better explained by common descent and branching evolution through natural selection, than by the individual creation of fixed species. Darwin expected species to change slowly, but not at the same rate – some organisms such as '' Lingula'' were unchanged since the earliest fossils. The pace of natural selection would depend on variability and change in the environment. This distanced his theory from
Lamarckian Lamarckism, also known as Lamarckian inheritance or neo-Lamarckism, is the notion that an organism can pass on to its offspring physical characteristics that the parent organism acquired through use or disuse during its lifetime. It is also calle ...
laws of inevitable progress. It has been argued that this anticipated the
punctuated equilibrium In evolutionary biology, punctuated equilibrium (also called punctuated equilibria) is a theory that proposes that once a species appears in the fossil record, the population will become stable, showing little evolutionary change for most of ...
hypothesis, but other scholars have preferred to emphasise Darwin's commitment to gradualism. He cited
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Ow ...
's findings that the earliest members of a class were a few simple and generalised species with characteristics intermediate between modern forms, and were followed by increasingly diverse and specialised forms, matching the branching of common descent from an ancestor. Patterns of extinction matched his theory, with related groups of species having a continued existence until extinction, then not reappearing. Recently extinct species were more similar to living species than those from earlier eras, and as he had seen in South America, and
William Clift William Clift FRS (14 February 1775 – 20 June 1849) was a British illustrator and conservator. Early life Clift was born in Burcombe near Bodmin in Cornwall. He was the youngest of seven children and grew up in poverty following his fat ...
had shown in Australia, fossils from recent geological periods resembled species still living in the same area.


Geographic distribution

Chapter XI deals with evidence from
biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
, starting with the observation that differences in flora and fauna from separate regions cannot be explained by environmental differences alone; South America, Africa, and Australia all have regions with similar climates at similar latitudes, but those regions have very different plants and animals. The species found in one area of a continent are more closely allied with species found in other regions of that same continent than to species found on other continents. Darwin noted that barriers to migration played an important role in the differences between the species of different regions. The coastal sea life of the Atlantic and Pacific sides of Central America had almost no species in common even though the
Isthmus of Panama The Isthmus of Panama ( es, Istmo de Panamá), also historically known as the Isthmus of Darien (), is the narrow strip of land that lies between the Caribbean Sea and the Pacific Ocean, linking North and South America. It contains the country ...
was only a few miles wide. His explanation was a combination of migration and descent with modification. He went on to say: "On this principle of inheritance with modification, we can understand how it is that sections of genera, whole genera, and even families are confined to the same areas, as is so commonly and notoriously the case." Darwin explained how a volcanic island formed a few hundred miles from a continent might be colonised by a few species from that continent. These species would become modified over time, but would still be related to species found on the continent, and Darwin observed that this was a common pattern. Darwin discussed ways that species could be dispersed across oceans to colonise islands, many of which he had investigated experimentally. Chapter XII continues the discussion of biogeography. After a brief discussion of freshwater species, it returns to oceanic islands and their peculiarities; for example on some islands roles played by mammals on continents were played by other animals such as flightless birds or reptiles. The summary of both chapters says:
... I think all the grand leading facts of geographical distribution are explicable on the theory of migration (generally of the more dominant forms of life), together with subsequent modification and the multiplication of new forms. We can thus understand the high importance of barriers, whether of land or water, which separate our several zoological and botanical provinces. We can thus understand the localisation of sub-genera, genera, and families; and how it is that under different latitudes, for instance in South America, the inhabitants of the plains and mountains, of the forests, marshes, and deserts, are in so mysterious a manner linked together by affinity, and are likewise linked to the extinct beings which formerly inhabited the same continent ... On these same principles, we can understand, as I have endeavoured to show, why oceanic islands should have few inhabitants, but of these a great number should be endemic or peculiar; ...


Classification, morphology, embryology, rudimentary organs

Chapter XIII starts by observing that classification depends on species being grouped together in a ''
Taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
'', a multilevel system of groups and sub-groups based on varying degrees of resemblance. After discussing classification issues, Darwin concludes:
All the foregoing rules and aids and difficulties in classification are explained, if I do not greatly deceive myself, on the view that the natural system is founded on descent with modification; that the characters which naturalists consider as showing true affinity between any two or more species, are those which have been inherited from a common parent, and, in so far, all true classification is genealogical; that community of descent is the hidden bond which naturalists have been unconsciously seeking, ...
Darwin discusses morphology, including the importance of
homologous structures In biology, homology is similarity due to shared ancestry between a pair of structures or genes in different taxa. A common example of homologous structures is the forelimbs of vertebrates, where the wings of bats and birds, the arms of pri ...
. He says, "What can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include the same bones, in the same relative positions?" This made no sense under doctrines of independent creation of species, as even
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Ow ...
had admitted, but the "explanation is manifest on the theory of the natural selection of successive slight modifications" showing
common descent Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal com ...
. He notes that animals of the same class often have extremely similar
embryos An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
. Darwin discusses rudimentary organs, such as the wings of flightless birds and the rudiments of pelvis and leg bones found in some snakes. He remarks that some rudimentary organs, such as teeth in
baleen whales Baleen whales ( systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea ( whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in t ...
, are found only in embryonic stages. These factors also supported his theory of descent with modification.


Concluding remarks

The final chapter, "Recapitulation and Conclusion", reviews points from earlier chapters, and Darwin concludes by hoping that his theory might produce revolutionary changes in many fields of natural history. He suggests that psychology will be put on a new foundation and implies the relevance of his theory to the first appearance of humanity with the sentence that "Light will be thrown on the origin of man and his history." Darwin ends with a passage that became well known and much quoted:
It is interesting to contemplate an entangled bank, clothed with many plants of many kinds, with birds singing on the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner, have all been produced by laws acting around us ... Thus, from the war of nature, from famine and death, the most exalted object which we are capable of conceiving, namely, the production of the higher animals, directly follows. There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.
Darwin added the phrase "by the Creator" from the 1860 second edition onwards, so that the ultimate sentence begins "There is grandeur in this view of life, with its several powers, having been originally breathed by the Creator into a few forms or into one".


Structure, style, and themes


Nature and structure of Darwin's argument

Darwin's aims were twofold: to show that species had not been separately created, and to show that
natural selection 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 heritable traits characteristic of a population over generations. Cha ...
had been the chief agent of change. He knew that his readers were already familiar with the concept of transmutation of species from '' Vestiges'', and his introduction ridicules that work as failing to provide a viable mechanism. Therefore, the first four chapters lay out his case that selection in nature, caused by the struggle for existence, is analogous to the selection of variations under domestication, and that the accumulation of adaptive variations provides a scientifically testable mechanism for evolutionary
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution withi ...
. Later chapters provide evidence that evolution has occurred, supporting the idea of branching, adaptive evolution without directly proving that selection is the mechanism. Darwin presents supporting facts drawn from many disciplines, showing that his theory could explain a myriad of observations from many fields of natural history that were inexplicable under the alternative concept that species had been individually created. The structure of Darwin's argument showed the influence of
John Herschel Sir John Frederick William Herschel, 1st Baronet (; 7 March 1792 – 11 May 1871) was an English polymath active as a mathematician, astronomer, chemist, inventor, experimental photographer who invented the blueprint and did botanical wo ...
, whose philosophy of science maintained that a mechanism could be called a ''vera causa'' (true cause) if three things could be demonstrated: its existence in nature, its ability to produce the effects of interest, and its ability to explain a wide range of observations.


Literary style

The ''Examiner'' review of 3 December 1859 commented, "Much of Mr. Darwin's volume is what ordinary readers would call 'tough reading;' that is, writing which to comprehend requires concentrated attention and some preparation for the task. All, however, is by no means of this description, and many parts of the book abound in information, easy to comprehend and both instructive and entertaining." While the book was readable enough to sell, its dryness ensured that it was seen as aimed at specialist scientists and could not be dismissed as mere journalism or imaginative fiction. Unlike the still-popular ''Vestiges'', it avoided the narrative style of the historical novel and cosmological speculation, though the closing sentence clearly hinted at cosmic progression. Darwin had long been immersed in the literary forms and practices of specialist science, and made effective use of his skills in structuring arguments.
David Quammen David Quammen (born February 24, 1948) is an American science, nature, and travel writer and the author of fifteen books. His articles have appeared in ''Outside Magazine'', '' National Geographic'', '' Harper's'', '' Rolling Stone'', ''The New ...
has described the book as written in everyday language for a wide audience, but noted that Darwin's literary style was uneven: in some places he used convoluted sentences that are difficult to read, while in other places his writing was beautiful. Quammen advised that later editions were weakened by Darwin making concessions and adding details to address his critics, and recommended the first edition. James T. Costa said that because the book was an abstract produced in haste in response to Wallace's essay, it was more approachable than the big book on natural selection Darwin had been working on, which would have been encumbered by scholarly footnotes and much more technical detail. He added that some parts of ''Origin'' are dense, but other parts are almost lyrical, and the case studies and observations are presented in a narrative style unusual in serious scientific books, which broadened its audience.


Human evolution

From his early transmutation notebooks in the late 1830s onwards, Darwin considered
human evolution Human evolution is the evolutionary process within the history of primates that led to the emergence of '' Homo sapiens'' as a distinct species of the hominid family, which includes the great apes. This process involved the gradual developmen ...
as part of the natural processes he was investigating, and rejected divine intervention. In 1856, his "big book on species" titled ''
Natural Selection 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 heritable traits characteristic of a population over generations. Cha ...
'' was to include a "note on Man", but when Wallace enquired in December 1857, Darwin replied; "You ask whether I shall discuss 'man';—I think I shall avoid whole subject, as so surrounded with prejudices, though I fully admit that it is the highest & most interesting problem for the naturalist." On 28 March 1859, with his manuscript for the book well under way, Darwin wrote to Lyell offering the suggested publisher John Murray assurances "That I do not discuss origin of man". In the final chapter of ''On the Origin of Species'', " Recapitulation and Conclusion", Darwin briefly highlights the human implications of his theory:
"In the distant future I see open fields for far more important researches. Psychology will be based on a new foundation, that of the necessary acquirement of each mental power and capacity by gradation. Light will be thrown on the origin of man and his history."
Discussing this in January 1860, Darwin assured Lyell that "by the sentence ight will be thrown on the origin of man and his historyI show that I believe man is in same predicament with other animals. Many modern writers have seen this sentence as Darwin’s only reference to humans in the book;
Janet Browne Elizabeth Janet Browne (née Bell, born 30 March 1950) is a British historian of science, known especially for her work on the history of 19th-century biology. She taught at the Wellcome Trust Centre for the History of Medicine, University Col ...
describes it as his only discussion there of human origins, while noting that the book makes other references to humanity. Some other statements in the book are quietly effective at pointing out the implication that humans are simply another species, evolving through the same processes and principles affecting other organisms. For example, in Chapter III: "Struggle for Existence" Darwin includes "slow-breeding man" among other examples of Malthusian population growth. In his discussions on morphology, Darwin compares and comments on bone structures that are homologous between humans and other mammals. Darwin's early notebooks discussed how non-adaptive characteristics could be selected when animals or humans chose mates, with races of humans differing over ideas of beauty. In his 1856 notes responding to
Robert Knox Robert Knox (4 September 1791 – 20 December 1862) was a Scottish anatomist and ethnologist best known for his involvement in the Burke and Hare murders. Born in Edinburgh, Scotland, Knox eventually partnered with anatomist and former teach ...
's ''The Races of Man: A Fragment'', he called this effect
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (in ...
. He added notes on sexual selection to his "big book on species", and in mid-1857 he added a section heading "Theory applied to Races of Man", but did not add text on this topic. In ''On the Origin of Species'', Chapter VI: "Difficulties on Theory", Darwin mentions this in the context of "slight and unimportant variations":
I might have adduced for this same purpose the differences between the races of man, which are so strongly marked; I may add that some little light can apparently be thrown on the origin of these differences, chiefly through sexual selection of a particular kind, but without here entering on copious details my reasoning would appear frivolous."
When Darwin published ''
The Descent of Man, and Selection in Relation to Sex ''The Descent of Man, and Selection in Relation to Sex'' is a book by English natural history, naturalist Charles Darwin, first published in 1871, which applies evolutionary theory to human evolution, and details his theory of sexual selection, ...
'' twelve years later, he said that he had not gone into detail on human evolution in the ''Origin'' as he thought that would "only add to the prejudices against my views". He had not completely avoided the topic:, Quote: "During many years I collected notes on the origin or descent of man, without any intention of publishing on the subject, but rather with the determination not to publish, as I thought that I should thus only add to the prejudices against my views."
It seemed to me sufficient to indicate, in the first edition of my 'Origin of Species,' that by this work 'light would be thrown on the origin of man and his history;' and this implies that man must be included with other organic beings in any general conclusion respecting his manner of appearance on this earth.
He also said that he had "merely alluded" in that book to sexual selection differentiating human races.


Reception

The book aroused international interest and a widespread debate, with no sharp line between scientific issues and ideological, social and religious implications. Much of the initial reaction was hostile, in a large part because very few reviewers actually understood his theory, but Darwin had to be taken seriously as a prominent and respected name in science. Bishop
Samuel Wilberforce Samuel Wilberforce, FRS (7 September 1805 – 19 July 1873) was an English bishop in the Church of England, and the third son of William Wilberforce. Known as "Soapy Sam", Wilberforce was one of the greatest public speakers of his day. Natural ...
wrote a review in ''Quarterly Review'' in 1860 where he disagreed with Darwin's 'argument'. There was much less controversy than had greeted the 1844 publication ''
Vestiges of Creation ''Vestiges of the Natural History of Creation'' is an 1844 work of speculative natural history and philosophy by Robert Chambers. Published anonymously in England, it brought together various ideas of stellar evolution with the progressive tr ...
'', which had been rejected by scientists, but had influenced a wide public readership into believing that nature and human society were governed by natural laws. The ''Origin of Species'' as a book of wide general interest became associated with ideas of social reform. Its proponents made full use of a surge in the publication of review journals, and it was given more popular attention than almost any other scientific work, though it failed to match the continuing sales of ''Vestiges''. Darwin's book legitimised scientific discussion of evolutionary mechanisms, and the newly coined term '
Darwinism Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations tha ...
' was used to cover the whole range of
evolutionism Evolutionism is a term used (often derogatorily) to denote the theory of evolution. Its exact meaning has changed over time as the study of evolution has progressed. In the 19th century, it was used to describe the belief that organisms deliberat ...
, not just his own ideas. By the mid-1870s, evolutionism was triumphant. While Darwin had been somewhat coy about human origins, not identifying any explicit conclusion on the matter in his book, he had dropped enough hints about human's animal ancestry for the inference to be made, and the first review claimed it made a creed of the "men from monkeys" idea from ''Vestiges''.
Human evolution Human evolution is the evolutionary process within the history of primates that led to the emergence of '' Homo sapiens'' as a distinct species of the hominid family, which includes the great apes. This process involved the gradual developmen ...
became central to the debate and was strongly argued by Huxley who featured it in his popular "working-men's lectures". Darwin did not publish his own views on this until 1871. The naturalism of
natural selection 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 heritable traits characteristic of a population over generations. Cha ...
conflicted with presumptions of purpose in nature and while this could be reconciled by
theistic evolution Theistic evolution (also known as theistic evolutionism or God-guided evolution) is a theological view that God creates through laws of nature. Its religious teachings are fully compatible with the findings of modern science, including biologica ...
, other mechanisms implying more progress or purpose were more acceptable.
Herbert Spencer Herbert Spencer (27 April 1820 – 8 December 1903) was an English philosopher, psychologist, biologist, anthropologist, and sociologist famous for his hypothesis of social Darwinism. Spencer originated the expression " survival of the f ...
had already incorporated
Lamarckism Lamarckism, also known as Lamarckian inheritance or neo-Lamarckism, is the notion that an organism can pass on to its offspring physical characteristics that the parent organism acquired through use or disuse during its lifetime. It is also calle ...
into his popular philosophy of progressive
free market In economics, a free market is an economic system in which the prices of goods and services are determined by supply and demand expressed by sellers and buyers. Such markets, as modeled, operate without the intervention of government or any ot ...
human society. He popularised the terms 'evolution' and '
survival of the fittest "Survival of the fittest" is a phrase that originated from Darwinian evolutionary theory as a way of describing the mechanism of natural selection. The biological concept of fitness is defined as reproductive success. In Darwinian terms, ...
', and many thought Spencer was central to evolutionary thinking.


Impact on the scientific community

Scientific readers were already aware of arguments that species changed through processes that were subject to laws of nature, but the transmutational ideas of Lamarck and the vague "law of development" of ''Vestiges'' had not found scientific favour. Darwin presented
natural selection 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 heritable traits characteristic of a population over generations. Cha ...
as a scientifically testable mechanism while accepting that other mechanisms such as
inheritance of acquired characters Lamarckism, also known as Lamarckian inheritance or neo-Lamarckism, is the notion that an organism can pass on to its offspring physical characteristics that the parent organism acquired through use or disuse during its lifetime. It is also calle ...
were possible. His strategy established that evolution through natural laws was worthy of scientific study, and by 1875, most scientists accepted that evolution occurred but few thought natural selection was significant. Darwin's scientific method was also disputed, with his proponents favouring the
empiricism In philosophy, empiricism is an epistemological theory that holds that knowledge or justification comes only or primarily from sensory experience. It is one of several views within epistemology, along with rationalism and skepticism. Empir ...
of
John Stuart Mill John Stuart Mill (20 May 1806 – 7 May 1873) was an English philosopher, political economist, Member of Parliament (MP) and civil servant. One of the most influential thinkers in the history of classical liberalism, he contributed widely to ...
's ''
A System of Logic ''A System of Logic, Ratiocinative and Inductive'' is an 1843 book by English philosopher John Stuart Mill. Overview In this work, he formulated the five principles of inductive reasoning that are known as Mill's Methods. This work is important ...
'', while opponents held to the idealist school of
William Whewell William Whewell ( ; 24 May 17946 March 1866) was an English polymath, scientist, Anglican priest, philosopher, theologian, and historian of science. He was Master of Trinity College, Cambridge. In his time as a student there, he achieved ...
's ''Philosophy of the Inductive Sciences'', in which investigation could begin with the intuitive idea that species were fixed objects created by design. Early support for Darwin's ideas came from the findings of field naturalists studying
biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
and ecology, including
Joseph Dalton Hooker Sir Joseph Dalton Hooker (30 June 1817 – 10 December 1911) was a British botanist and explorer in the 19th century. He was a founder of geographical botany and Charles Darwin's closest friend. For twenty years he served as director of ...
in 1860, and
Asa Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutually excl ...
in 1862.
Henry Walter Bates Henry Walter Bates (8 February 1825, in Leicester – 16 February 1892, in London) was an English naturalist and explorer who gave the first scientific account of mimicry in animals. He was most famous for his expedition to the rainforests of ...
presented research in 1861 that explained insect mimicry using natural selection.
Alfred Russel Wallace Alfred Russel Wallace (8 January 1823 – 7 November 1913) was a British natural history, naturalist, explorer, geographer, anthropologist, biologist and illustrator. He is best known for independently conceiving the theory of evolution thro ...
discussed evidence from his
Malay archipelago The Malay Archipelago ( Indonesian/ Malay: , tgl, Kapuluang Malay) is the archipelago between mainland Indochina and Australia. It has also been called the " Malay world," " Nusantara", "East Indies", Indo-Australian Archipelago, Spices Arc ...
research, including an 1864 paper with an evolutionary explanation for the
Wallace line The Wallace Line or Wallace's Line is a faunal boundary line drawn in 1859 by the British naturalist Alfred Russel Wallace and named by English biologist Thomas Henry Huxley that separates the biogeographical realms of Asia and Wallacea, a trans ...
. Evolution had less obvious applications to
anatomy Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having i ...
and morphology, and at first had little impact on the research of the anatomist
Thomas Henry Huxley Thomas Henry Huxley (4 May 1825 – 29 June 1895) was an English biologist and anthropologist specialising in comparative anatomy. He has become known as "Darwin's Bulldog" for his advocacy of Charles Darwin's theory of evolution. The stori ...
. Despite this, Huxley strongly supported Darwin on evolution; though he called for experiments to show whether natural selection could form new species, and questioned if Darwin's
gradualism Gradualism, from the Latin ''gradus'' ("step"), is a hypothesis, a theory or a tenet assuming that change comes about gradually or that variation is gradual in nature and happens over time as opposed to in large steps. Uniformitarianism, increment ...
was sufficient without sudden leaps to cause
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution withi ...
. Huxley wanted science to be secular, without religious interference, and his article in the April 1860 ''
Westminster Review The ''Westminster Review'' was a quarterly British publication. Established in 1823 as the official organ of the Philosophical Radicals, it was published from 1824 to 1914. James Mill was one of the driving forces behind the liberal journal unt ...
'' promoted scientific naturalism over natural theology, praising Darwin for "extending the domination of Science over regions of thought into which she has, as yet, hardly penetrated" and coining the term "
Darwinism Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations tha ...
" as part of his efforts to secularise and professionalise science. Huxley gained influence, and initiated the
X Club The X Club was a dining club of nine men who supported the theories of natural selection and academic liberalism in late 19th-century England. Thomas Henry Huxley was the initiator; he called the first meeting for 3 November 1864. The club m ...
, which used the journal ''
Nature Nature, in the broadest sense, is the physical world or universe. "Nature" can refer to the phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. Although humans are ...
'' to promote evolution and naturalism, shaping much of late-Victorian science. Later, the German morphologist
Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new s ...
would convince Huxley that comparative anatomy and
palaeontology Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
could be used to reconstruct evolutionary genealogies. The leading naturalist in Britain was the anatomist
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Ow ...
, an idealist who had shifted to the view in the 1850s that the history of life was the gradual unfolding of a divine plan. Owen's review of the ''Origin'' in the April 1860 ''Edinburgh Review'' bitterly attacked Huxley, Hooker and Darwin, but also signalled acceptance of a kind of evolution as a
teleological Teleology (from and )Partridge, Eric. 1977''Origins: A Short Etymological Dictionary of Modern English'' London: Routledge, p. 4187. or finalityDubray, Charles. 2020 912Teleology" In ''The Catholic Encyclopedia'' 14. New York: Robert Appleton ...
plan in a continuous "ordained becoming", with new species appearing by natural birth. Others that rejected natural selection, but supported "creation by birth", included the
Duke of Argyll Duke of Argyll ( gd, Diùc Earraghàidheil) is a title created in the peerage of Scotland in 1701 and in the peerage of the United Kingdom in 1892. The earls, marquesses, and dukes of Argyll were for several centuries among the most powerfu ...
who explained beauty in plumage by design. Since 1858, Huxley had emphasised anatomical similarities between apes and humans, contesting Owen's view that humans were a separate sub-class. Their disagreement over human origins came to the fore at the
British Association for the Advancement of Science The British Science Association (BSA) is a charity and learned society founded in 1831 to aid in the promotion and development of science. Until 2009 it was known as the British Association for the Advancement of Science (BA). The current Chi ...
meeting featuring the legendary
1860 Oxford evolution debate The 1860 Oxford evolution debate took place at the Oxford University Museum in Oxford, England, on 30 June 1860, seven months after the publication of Charles Darwin's ''On the Origin of Species''. Several prominent British scientists and philo ...
. In two years of acrimonious public dispute that
Charles Kingsley Charles Kingsley (12 June 1819 – 23 January 1875) was a broad church priest of the Church of England, a university professor, social reformer, historian, novelist and poet. He is particularly associated with Christian socialism, the worki ...
satirised as the "
Great Hippocampus Question The Great Hippocampus Question was a 19th-century scientific controversy about the anatomy of ape and human uniqueness. The dispute between Thomas Henry Huxley and Richard Owen became central to the scientific debate on human evolution that follow ...
" and parodied in '' The Water-Babies'' as the "great hippopotamus test", Huxley showed that Owen was incorrect in asserting that ape brains lacked a structure present in human brains. Others, including
Charles Lyell Sir Charles Lyell, 1st Baronet, (14 November 1797 – 22 February 1875) was a Scottish geologist who demonstrated the power of known natural causes in explaining the earth's history. He is best known as the author of ''Principles of Geolo ...
and
Alfred Russel Wallace Alfred Russel Wallace (8 January 1823 – 7 November 1913) was a British natural history, naturalist, explorer, geographer, anthropologist, biologist and illustrator. He is best known for independently conceiving the theory of evolution thro ...
, thought that humans shared a common ancestor with apes, but higher mental faculties could not have evolved through a purely material process. Darwin published his own explanation in the '' Descent of Man'' (1871).


Impact outside Great Britain

Evolutionary ideas, although not natural selection, were accepted by German biologists accustomed to ideas of homology in morphology from
Goethe Johann Wolfgang von Goethe (28 August 1749 – 22 March 1832) was a German poet, playwright, novelist, scientist, statesman, theatre director, and critic. His works include plays, poetry, literature, and aesthetic criticism, as well as tr ...
's '' Metamorphosis of Plants'' and from their long tradition of comparative anatomy. Bronn's alterations in his German translation added to the misgivings of conservatives, but enthused political radicals.
Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new s ...
was particularly ardent, aiming to synthesise Darwin's ideas with those of
Lamarck Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biolo ...
and Goethe while still reflecting the spirit of ''
Naturphilosophie ''Naturphilosophie'' (German for "nature-philosophy") is a term used in English-language philosophy to identify a current in the philosophical tradition of German idealism, as applied to the study of nature in the earlier 19th century. German s ...
''. Their ambitious programme to reconstruct the
evolutionary history of life The history of life on Earth traces the processes by which living and fossil organisms evolved, from the earliest emergence of life to present day. Earth formed about 4.5 billion years ago (abbreviated as ''Ga'', for ''gigaannum'') and evide ...
was joined by Huxley and supported by discoveries in
palaeontology Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
. Haeckel used
embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, '' -logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embr ...
extensively in his
recapitulation theory The theory of recapitulation, also called the biogenetic law or embryological parallelism—often expressed using Ernst Haeckel's phrase "ontogeny recapitulates phylogeny"—is a historical hypothesis that the development of the embryo of an a ...
, which embodied a progressive, almost linear model of evolution. Darwin was cautious about such histories, and had already noted that von Baer's laws of embryology supported his idea of complex branching.
Asa Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutually excl ...
promoted and defended ''Origin'' against those American naturalists with an idealist approach, notably
Louis Agassiz Jean Louis Rodolphe Agassiz ( ; ) FRS (For) FRSE (May 28, 1807 – December 14, 1873) was a Swiss-born American biologist and geologist who is recognized as a scholar of Earth's natural history. Spending his early life in Switzerland, he rec ...
who viewed every species as a distinct fixed unit in the mind of the Creator, classifying as species what others considered merely varieties.
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontologist, comparative anatomist, herpetologist, and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy interes ...
and
Alpheus Hyatt Alpheus Hyatt (April 5, 1838 – January 15, 1902) was an American zoologist and palaeontologist. Biography Alpheus Hyatt II was born in Washington, D.C. to Alpheus Hyatt and Harriet Randolph (King) Hyatt. He briefly attended the Mary ...
reconciled this view with evolutionism in a form of
neo-Lamarckism Lamarckism, also known as Lamarckian inheritance or neo-Lamarckism, is the notion that an organism can pass on to its offspring physical characteristics that the parent organism acquired through use or disuse during its lifetime. It is also calle ...
involving recapitulation theory. French-speaking naturalists in several countries showed appreciation of the much-modified French translation by
Clémence Royer Clémence Royer (21 April 1830 – 6 February 1902) was a self-taught French scholar who lectured and wrote on economics, philosophy, science and feminism. She is best known for her controversial 1862 French translation of Charles Darwin's ' ...
, but Darwin's ideas had little impact in France, where any scientists supporting evolutionary ideas opted for a form of Lamarckism. The intelligentsia in Russia had accepted the general phenomenon of evolution for several years before Darwin had published his theory, and scientists were quick to take it into account, although the
Malthusian Malthusianism is the idea that population growth is potentially exponential while the growth of the food supply or other resources is linear, which eventually reduces living standards to the point of triggering a population die off. This event, ...
aspects were felt to be relatively unimportant. The political economy of struggle was criticised as a British stereotype by
Karl Marx Karl Heinrich Marx (; 5 May 1818 – 14 March 1883) was a German philosopher, economist, historian, sociologist, political theorist, journalist, critic of political economy, and socialist revolutionary. His best-known titles are the 1848 ...
and by
Leo Tolstoy Count Lev Nikolayevich TolstoyTolstoy pronounced his first name as , which corresponds to the romanization ''Lyov''. () (; russian: link=no, Лев Николаевич Толстой,In Tolstoy's day, his name was written as in pre-refor ...
, who had the character Levin in his novel ''
Anna Karenina ''Anna Karenina'' ( rus, «Анна Каренина», p=ˈanːə kɐˈrʲenʲɪnə) is a novel by the Russian author Leo Tolstoy, first published in book form in 1878. Widely considered to be one of the greatest works of literature ever writt ...
'' voice sharp criticism of the morality of Darwin's views.


Challenges to natural selection

There were serious scientific objections to the process of
natural selection 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 heritable traits characteristic of a population over generations. Cha ...
as the key mechanism of evolution, including
Carl Nägeli Carl Wilhelm von Nägeli (26 or 27 March 1817 – 10 May 1891) was a Swiss botanist. He studied cell division and pollination but became known as the man who discouraged Gregor Mendel from further work on genetics. He rejected natural selection ...
's insistence that a trivial characteristic with no adaptive advantage could not be developed by selection. Darwin conceded that these could be linked to adaptive characteristics. His estimate that the
age of the Earth The age of Earth is estimated to be 4.54 ± 0.05 billion years This age may represent the age of Earth's accretion, or core formation, or of the material from which Earth formed. This dating is based on evidence from radiometric age-dating of ...
allowed gradual evolution was disputed by William Thomson (later awarded the title Lord Kelvin), who calculated that it had cooled in less than 100 million years. Darwin accepted
blending inheritance Blending may refer to: * The process of mixing in process engineering * Mixing paints to achieve a greater range of colors * Blending (alcohol production), a technique to produce alcoholic beverages by mixing different brews * Blending (linguisti ...
, but Fleeming Jenkin calculated that as it mixed traits, natural selection could not accumulate useful traits. Darwin tried to meet these objections in the fifth edition.
Mivart St. George Jackson Mivart (30 November 1827 – 1 April 1900) was an English biologist. He is famous for starting as an ardent believer in natural selection who later became one of its fiercest critics. Mivart attempted to reconcile ...
supported directed evolution, and compiled scientific and religious objections to natural selection. In response, Darwin made considerable changes to the sixth edition. The problems of the age of the Earth and heredity were only resolved in the 20th century. By the mid-1870s, most scientists accepted evolution, but relegated natural selection to a minor role as they believed evolution was purposeful and progressive. The range of evolutionary theories during "
the eclipse of Darwinism Julian Huxley used the phrase "the eclipse of Darwinism" to describe the state of affairs prior to what he called the "modern synthesis". During the "eclipse", evolution was widely accepted in scientific circles but relatively few biologists be ...
" included forms of " saltationism" in which new species were thought to arise through "jumps" rather than gradual adaptation, forms of
orthogenesis Orthogenesis, also known as orthogenetic evolution, progressive evolution, evolutionary progress, or progressionism, is an obsolete biological hypothesis that organisms have an innate tendency to evolve in a definite direction towards some g ...
claiming that species had an inherent tendency to change in a particular direction, and forms of neo-Lamarckism in which inheritance of acquired characteristics led to progress. The minority view of
August Weismann August Friedrich Leopold Weismann FRS (For), HonFRSE, LLD (17 January 18345 November 1914) was a German evolutionary biologist. Fellow German Ernst Mayr ranked him as the second most notable evolutionary theorist of the 19th century, after Cha ...
, that natural selection was the only mechanism, was called
neo-Darwinism Neo-Darwinism is generally used to describe any integration of Charles Darwin's theory of evolution by natural selection with Gregor Mendel's theory of genetics. It mostly refers to evolutionary theory from either 1895 (for the combinations of Dar ...
. It was thought that the rediscovery of
Mendelian inheritance Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later popularize ...
invalidated Darwin's views.


Impact on economic and political debates

While some, like Spencer, used analogy from natural selection as an argument against government intervention in the economy to benefit the poor, others, including
Alfred Russel Wallace Alfred Russel Wallace (8 January 1823 – 7 November 1913) was a British natural history, naturalist, explorer, geographer, anthropologist, biologist and illustrator. He is best known for independently conceiving the theory of evolution thro ...
, argued that action was needed to correct social and economic inequities to level the playing field before natural selection could improve humanity further. Some political commentaries, including
Walter Bagehot Walter Bagehot ( ; 3 February 1826 – 24 March 1877) was an English journalist, businessman, and essayist, who wrote extensively about government, economics, literature and race. He is known for co-founding the '' National Review'' in 185 ...
's ''Physics and Politics'' (1872), attempted to extend the idea of natural selection to competition between nations and between human races. Such ideas were incorporated into what was already an ongoing effort by some working in
anthropology Anthropology is the scientific study of humanity, concerned with human behavior, human biology, cultures, societies, and linguistics, in both the present and past, including past human species. Social anthropology studies patterns of be ...
to provide scientific evidence for the superiority of Caucasians over non-white races and justify European
imperialism Imperialism is the state policy, practice, or advocacy of extending power and dominion, especially by direct territorial acquisition or by gaining political and economic control of other areas, often through employing hard power (economic powe ...
. Historians write that most such political and economic commentators had only a superficial understanding of Darwin's scientific theory, and were as strongly influenced by other concepts about social progress and evolution, such as the Lamarckian ideas of Spencer and Haeckel, as they were by Darwin's work. Darwin objected to his ideas being used to justify military aggression and unethical business practices as he believed morality was part of fitness in humans, and he opposed
polygenism Polygenism is a theory of human origins which posits the view that the human races are of different origins (''polygenesis''). This view is opposite to the idea of monogenism, which posits a single origin of humanity. Modern scientific views no ...
, the idea that human races were fundamentally distinct and did not share a recent common ancestry.


Religious attitudes

The book produced a wide range of religious responses at a time of changing ideas and increasing secularisation. The issues raised were complex and there was a large middle ground. Developments in geology meant that there was little opposition based on a literal reading of
Genesis Genesis may refer to: Bible * Book of Genesis, the first book of the biblical scriptures of both Judaism and Christianity, describing the creation of the Earth and of mankind * Genesis creation narrative, the first several chapters of the Book of ...
, but defence of the
argument from design The teleological argument (from ; also known as physico-theological argument, argument from design, or intelligent design argument) is an argument for the existence of God or, more generally, that complex functionality in the natural world wh ...
and
natural theology Natural theology, once also termed physico-theology, is a type of theology that seeks to provide arguments for theological topics (such as the existence of a deity) based on reason and the discoveries of science. This distinguishes it from ...
was central to debates over the book in the English-speaking world. Natural theology was not a unified doctrine, and while some such as Louis Agassiz were strongly opposed to the ideas in the book, others sought a reconciliation in which evolution was seen as purposeful. In the Church of England, some liberal clergymen interpreted natural selection as an instrument of God's design, with the cleric Charles Kingsley seeing it as "just as noble a conception of Deity". In the second edition of January 1860, Darwin quoted Kingsley as "a celebrated cleric", and added the phrase "by the Creator" to the closing sentence, which from then on read "life, with its several powers, having been originally breathed by the Creator into a few forms or into one". While some commentators have taken this as a concession to religion that Darwin later regretted, Darwin's view at the time was of God creating life through the laws of nature, and even in the first edition there are several references to "creation". Baden Powell praised "Mr Darwin's masterly volume upportingthe grand principle of the self-evolving powers of nature". In America,
Asa Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutually excl ...
argued that evolution is the secondary effect, or ''modus operandi'', of the first cause, design, and published a pamphlet defending the book in terms of
theistic evolution Theistic evolution (also known as theistic evolutionism or God-guided evolution) is a theological view that God creates through laws of nature. Its religious teachings are fully compatible with the findings of modern science, including biologica ...
, ''Natural Selection is not inconsistent with Natural Theology''. Theistic evolution became a popular compromise, and St. George Jackson Mivart was among those accepting evolution but attacking Darwin's naturalistic mechanism. Eventually it was realised that supernatural intervention could not be a scientific explanation, and naturalistic mechanisms such as
neo-Lamarckism Lamarckism, also known as Lamarckian inheritance or neo-Lamarckism, is the notion that an organism can pass on to its offspring physical characteristics that the parent organism acquired through use or disuse during its lifetime. It is also calle ...
were favoured over
natural selection 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 heritable traits characteristic of a population over generations. Cha ...
as being more compatible with purpose. Even though the book did not explicitly spell out Darwin's beliefs about human origins, it had dropped a number of hints about human's animal ancestry
and quickly became central to the debate, as mental and moral qualities were seen as spiritual aspects of the immaterial
soul In many religious and philosophical traditions, there is a belief that a soul is "the immaterial aspect or essence of a human being". Etymology The Modern English noun '' soul'' is derived from Old English ''sāwol, sāwel''. The earliest att ...
, and it was believed that animals did not have spiritual qualities. This conflict could be reconciled by supposing there was some supernatural intervention on the path leading to humans, or viewing evolution as a purposeful and progressive ascent to mankind's position at the head of nature. While many conservative theologians accepted evolution,
Charles Hodge Charles Hodge (December 27, 1797 – June 19, 1878) was a Reformed Presbyterian theologian and principal of Princeton Theological Seminary between 1851 and 1878. He was a leading exponent of the Princeton Theology, an orthodox Calvinist theo ...
argued in his 1874 critique "What is Darwinism?" that "
Darwinism Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations tha ...
", defined narrowly as including rejection of design, was atheism though he accepted that Asa Gray did not reject design. Asa Gray responded that this charge misrepresented Darwin's text. By the early 20th century, four noted authors of ''
The Fundamentals ''The Fundamentals: A Testimony To The Truth'' (generally referred to simply as ''The Fundamentals'') is a set of ninety essays published between 1910 and 1915 by the Testimony Publishing Company of Chicago. It was initially published quarterly in ...
'' were explicitly open to the possibility that God created through evolution, but
fundamentalism Fundamentalism is a tendency among certain groups and individuals that is characterized by the application of a strict literal interpretation to scriptures, dogmas, or ideologies, along with a strong belief in the importance of distinguish ...
inspired the American
creation–evolution controversy Recurring cultural, political, and theological rejection of evolution by religious groups (sometimes termed the creation–evolution controversy, the creation vs. evolution debate or the origins debate) exists regarding the origins of the Ea ...
that began in the 1920s. Some conservative
Roman Catholic Roman or Romans most often refers to: * Rome, the capital city of Italy *Ancient Rome, Roman civilization from 8th century BC to 5th century AD * Roman people, the people of ancient Rome *'' Epistle to the Romans'', shortened to ''Romans'', a let ...
writers and influential
Jesuit , image = Ihs-logo.svg , image_size = 175px , caption = ChristogramOfficial seal of the Jesuits , abbreviation = SJ , nickname = Jesuits , formation = , founders ...
s opposed evolution in the late 19th and early 20th century, but other Catholic writers, starting with Mivart, pointed out that early
Church Fathers The Church Fathers, Early Church Fathers, Christian Fathers, or Fathers of the Church were ancient and influential Christian theologians and writers who established the intellectual and doctrinal foundations of Christianity. The historical per ...
had not interpreted Genesis literally in this area. The
Vatican Vatican may refer to: Vatican City, the city-state ruled by the pope in Rome, including St. Peter's Basilica, Sistine Chapel, Vatican Museum The Holy See * The Holy See, the governing body of the Catholic Church and sovereign entity recognized ...
stated its official position in a 1950 papal encyclical, which held that evolution was not inconsistent with Catholic teaching.


Modern influence

Various alternative evolutionary mechanisms favoured during "
the eclipse of Darwinism Julian Huxley used the phrase "the eclipse of Darwinism" to describe the state of affairs prior to what he called the "modern synthesis". During the "eclipse", evolution was widely accepted in scientific circles but relatively few biologists be ...
" became untenable as more was learned about inheritance and
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, m ...
. The full significance of
natural selection 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 heritable traits characteristic of a population over generations. Cha ...
was at last accepted in the 1930s and 1940s as part of the modern evolutionary synthesis. During that synthesis biologists and statisticians, including
R. A. Fisher Sir Ronald Aylmer Fisher (17 February 1890 – 29 July 1962) was a British polymath who was active as a mathematician, statistician, biologist, geneticist, and academic. For his work in statistics, he has been described as "a genius who a ...
,
Sewall Wright Sewall Green Wright FRS(For) Honorary FRSE (December 21, 1889March 3, 1988) was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. He was a founder of population genetics alongsi ...
and
J. B. S. Haldane John Burdon Sanderson Haldane (; 5 November 18921 December 1964), nicknamed "Jack" or "JBS", was a British-Indian scientist who worked in physiology, genetics, evolutionary biology, and mathematics. With innovative use of statistics in biolo ...
, merged Darwinian selection with a statistical understanding of
Mendelian genetics Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later populari ...
.
Modern evolutionary theory Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variati ...
continues to develop. Darwin's theory of evolution by natural selection, with its tree-like model of branching
common descent Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal com ...
, has become the unifying theory of the
life science Life is a quality that distinguishes matter that has biological processes, such as signaling and self-sustaining processes, from that which does not, and is defined by the capacity for growth, reaction to stimuli, metabolism, energy ...
s. The theory explains the diversity of living organisms and their adaptation to the environment. It makes sense of the
geological record The geologic record in stratigraphy, paleontology and other natural sciences refers to the entirety of the layers of rock stratum, strata. That is, deposits laid down by volcanism or by Deposition (sediment), deposition of sediment derived from w ...
, biogeography, parallels in
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
nic development, biological homologies,
vestigiality Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...
,
cladistics Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived cha ...
,
phylogenetics In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
and other fields, with unrivalled explanatory power; it has also become essential to applied sciences such as
medicine Medicine is the science and practice of caring for a patient, managing the diagnosis, prognosis, prevention, treatment, palliation of their injury or disease, and promoting their health. Medicine encompasses a variety of health care pr ...
and agriculture. Despite the scientific consensus, a religion-based political
controversy Controversy is a state of prolonged public dispute or debate, usually concerning a matter of conflicting opinion or point of view. The word was coined from the Latin ''controversia'', as a composite of ''controversus'' – "turned in an opposite d ...
has developed over how evolution is taught in schools, especially in the United States. Interest in Darwin's writings continues, and scholars have generated an extensive literature, the
Darwin Industry The Darwin Industry refers to historical scholarship about, and the large community of historians of science working on, Charles Darwin's life, work, and influence. The term "has a slightly derogatory connotation, as if the scale of the research h ...
, about his life and work. The text of ''Origin'' itself has been subject to much analysis including a
variorum A variorum, short for ''(editio) cum notis variorum'', is a work that collates all known variants of a text. It is a work of textual criticism, whereby all variations and emendations are set side by side so that a reader can track how textual deci ...
, detailing the changes made in every edition, first published in 1959, and a concordance, an exhaustive external index published in 1981. Worldwide commemorations of the 150th anniversary of the publication of ''On the Origin of Species'' and the bicentenary of Darwin's birth were scheduled for 2009. They celebrated the ideas which "over the last 150 years have revolutionised our understanding of nature and our place within it". In a survey conducted by a group of academic booksellers, publishers and librarians in advance of Academic Book Week in the United Kingdom, ''On the Origin of Species'' was voted the most influential academic book ever written. It was hailed as "the supreme demonstration of why academic books matter" and "a book which has changed the way we think about everything".


See also

* ''On the Origin of Species'' – full text at
Wikisource Wikisource is an online digital library of free-content textual sources on a wiki, operated by the Wikimedia Foundation. Wikisource is the name of the project as a whole and the name for each instance of that project (each instance usually re ...
of the first edition, 1859 * ''The Origin of Species'' – full text at Wikisource of the 6th edition, 1872 *
Charles Darwin bibliography This is a list of the writings of Charles Darwin. Published works * 1829–1832. ecords of captured insects, inJames Francis Stephens, Stephens, J. F., ''Illustrations of British entomology'' * 1835: ''Extracts from Letters to Henslow'' (Rea ...
*
History of biology The history of biology traces the study of the living world from ancient to modern times. Although the concept of ''biology'' as a single coherent field arose in the 19th century, the biological sciences emerged from traditions of medicine an ...
*
History of evolutionary thought Evolutionary thought, the recognition that species change over time and the perceived understanding of how such processes work, has roots in antiquity—in the ideas of the ancient Greeks, Romans, Chinese, Church Fathers as well as in medie ...
* History of speciation * Modern synthesis (20th century), Modern evolutionary synthesis * The Complete Works of Charles Darwin Online * ''
The Descent of Man, and Selection in Relation to Sex ''The Descent of Man, and Selection in Relation to Sex'' is a book by English natural history, naturalist Charles Darwin, first published in 1871, which applies evolutionary theory to human evolution, and details his theory of sexual selection, ...
'', published in 1871; his second major book on evolutionary theory. * Transmutation of species


References


Works cited

* * * * * * * *. Published anonymously. *
Full image view
* * * * * * * * * * * * * * * * * * Also available her

* * *. Published anonymously. * * * * * * * * * * * * * * * * * * * * * * *


Further reading

* *
Vol. 2
* * *


Contemporary reviews

*. Published anonymously. *. Extract from ''Proceedings of the American Academy of Arts and Sciences'' 4 (1860): 411–415. *. *. Published anonymously. *. Published anonymously. *. *. Published anonymously. *. Published anonymously. *For further reviews, see


External links

*The Complete Works of Charles Darwin Online:
Table of contents
bibliography o

– links to text and images of all six British editions of ''The Origin of Species'', the 6th edition with additions and corrections (final text), the first American edition, and translations into Danish, Dutch, French, German, Polish, Russian and Spanish

showing every change between the six British editions *
On the Origin of Species
eBook provided by Project Gutenberg * *
''On the Origin of Species''
full text with embedded audio
Darwin Correspondence Project Home Page
University Library, Cambridge *View online at the Biodiversity Heritage Librar
''On the Origin of Species''
1860 American edition, D Appleton and Company, New York, with front insert by H. E. Barker, ''Lincolniana''
Darwin's notes on the creation of ''On the Origin of Species''
digitised in Cambridge Digital Library {{Authority control 1859 non-fiction books 1859 in science Books about evolution Books by Charles Darwin English-language books English non-fiction books John Murray (publishing house) books Biology textbooks