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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 Academic publishing is the subfield of publishing which distributes academic research and scholarship. Most academic work is published in academic journal articles, books or thes ...
by
Charles Darwin Charles Robert Darwin (; ; 12 February 1809 – 19 April 1882) was an English naturalist Natural history is a domain of inquiry involving organism In biology, an organism () is any organic, life, living system that fu ...

Charles Darwin
that is considered to be the foundation of
evolutionary biology Evolutionary biology is the subfield of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interacti ...
. Darwin's book introduced the
scientific theory A scientific theory is an explanation of an aspect of the natural science, natural world and universe that has been reproducibility, repeatedly tested and verified in accordance with the scientific method, using accepted protocol (science), protoc ...
that populations
evolve
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 right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
. The book presented a body of evidence that
the diversity of life
the diversity of life
arose by
common descent Common descent is a concept in evolutionary biology Evolutionary biology is the subfield of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chem ...
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 a Christian church Christian Church is a Protestant Protestantism is a form of Christianity that originated with the 16th-century Reformation, a movement against what its followers perceived to be Critic ...
, while science was part of
natural theology Natural theology, once also termed physico-theology, is a type of theology Theology is the systematic study of the nature of the divine Divinity or the divine are things that are either related to, devoted to, or proceeding from a deity.
. Ideas about the
transmutation of species Transmutation of species and transformism are 18th and 19th-century evolutionary ideas about the change of one species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism ...
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
T. H. Huxley
and his fellow members of the
X Club The X Club was a dining club A dining club (UK) or eating club (US) is a Social club, social group, usually requiring membership (which may, or may not be available only to certain people), which meets for dinners and discussion on a regular b ...
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 Sir Julian Sorell Huxley (22 June 1887 – 14 February 1975) was an English evolutionary biologist, eugenicist, and internationalist. He was a proponent of natural selection Natural selection is the differential surv ...
" 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 , adaptation has three related meanings. Firstly, it is the dynamic evolutionary process that fits s to their environment, enhancing their . Secondly, it is a state reached by the population during that process. Thirdly, it is a or adapti ...

adaptation
through natural selection became central to modern evolutionary theory, and it has now become the unifying concept of the
life science Life is a characteristic that distinguishes physical entities that have biological processes, such as signaling In signal processing Signal processing is an electrical engineering subfield that focuses on analysing, modifying, a ...
s.


Summary of Darwin's theory

Darwin's theory of evolution is based on key
fact A fact is something that is true True most commonly refers to truth Truth is the property of being in accord with fact or reality.Merriam-Webster's Online Dictionarytruth 2005 In everyday language, truth is typically ascribed to things ...
s and the
inference Inferences are steps in reasoning Reason is the capacity of consciously applying logic Logic is an interdisciplinary field which studies truth and reasoning Reason is the capacity of consciously making sense of things, applying logic ...

inference
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 File:Francesco Redi.jpg, Francesco Redi, the founder of biology, is recognized to be one of the greatest biologists of all ti ...
summarised as follows: :* Every
species In biology, a species is the basic unit of biological classification, 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 individu ...

species
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
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 Phenotypic trait, traits from parents to their offspring; either through asexual reproduction or sexual reproduction, the offspring cell (biology), cells or orga ...

heritable
(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 right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
(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 A philosopher is someone who practices philosophy Philosophy (from , ) is the study of general and fundamental questio ...

Aristotle
; the text he cites is a summary by Aristotle of the ideas of the earlier Greek philosopher
Empedocles Empedocles (; grc-gre, Ἐμπεδοκλῆς Empedocles (; grc-gre, wikt:Ἐμπεδοκλῆς, Ἐμπεδοκλῆς; , 444–443 BC) was a Ancient Greece, Greek pre-Socratic philosopher and a native citizen of Akragas, a Greek city ...

Empedocles
. Early
Christian Christians () are people who follow or adhere to Christianity, a monotheistic Abrahamic religion based on the life and teachings of Jesus in Christianity, Jesus Christ. The words ''Christ (title), Christ'' and ''Christian'' derive from the Koi ...

Christian
Church Fathers The Church Fathers, Early Church Fathers, Christian Fathers, or Fathers of the Church were ancient and influential Christian theologians Christian theology is the theology Theology is the systematic study of the nature of the Divinity, di ...
and
Medieval Europe In the history of Europe The history of Europe concerns itself with the discovery and collection, the study, organization and presentation and the interpretation of past events and affairs of the people of Europe since the beginning of ...
an scholars interpreted the
Genesis creation narrative The Genesis creation narrative is the creation myth A creation myth (or cosmogonic myth) is a symbol A symbol is a mark, sign, or that indicates, signifies, or is understood as representing an , , or . Symbols allow people to go beyo ...
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 body of thought on the ordinary formation of living organisms without descent from similar organisms. The theory of spontaneous generation held that living creatures could arise from nonliving matter and that such process ...
of life. The
Protestant Reformation The Reformation (alternatively named the Protestant Reformation or the European Reformation) was a major movement within Western Christianity Western Christianity is one of two sub-divisions of Christianity Christianity is an Abra ...
inspired a literal interpretation of the Bible, with concepts of creation that conflicted with the findings of an
emerging science
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 A machine is a man-made device that uses power to apply forces and control movement to perform an action. Mach ...
of
René Descartes René Descartes ( or ; ; Latinized Latinisation or Latinization can refer to: * Latinisation of names, the practice of rendering a non-Latin name in a Latin style * Latinisation in the Soviet Union, the campaign in the USSR during the 1920s ...

René Descartes
and the
empiricism In philosophy Philosophy (from , ) is the study of general and fundamental questions, such as those about Metaphysics, existence, reason, Epistemology, knowledge, Ethics, values, Philosophy of mind, mind, and Philosophy of language, l ...
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 scientific method is an Empirical evidence, empirical method o ...
. After the turmoil of the
English Civil War The English Civil War (1642–1651) was a series of civil wars A civil war, also known as an intrastate war in polemology, is a war War is an intense armed conflict between states State may refer to: Arts, entertainment, ...
, the
Royal Society The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society A learned society (; also known as a learned academy, scholarly society, or academic association) is an organization that exis ...
wanted to show that science did not threaten religious and political stability.
John Ray John Ray Fellow of the Royal Society, FRS (29 November 1627 – 17 January 1705) was a Christian England, English Natural history, naturalist widely regarded as one of the earliest of the English parson-naturalists. Until 1670, he wrote his na ...

John Ray
developed an influential
natural theology Natural theology, once also termed physico-theology, is a type of theology Theology is the systematic study of the nature of the divine Divinity or the divine are things that are either related to, devoted to, or proceeding from a deity.
of rational order; in his
taxonomy Taxonomy (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also refer to a specific classification scheme. Originally used only about biological ...
,
species In biology, a species is the basic unit of biological classification, 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 individu ...

species
were static and fixed, their adaptation and complexity designed by God, and
varieties Variety may refer to: Science and technology Mathematics * Algebraic variety Algebraic varieties are the central objects of study in algebraic geometry Algebraic geometry is a branch of mathematics Mathematics (from Ancient Greek, ...
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 In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, ...

parasitism
was a puzzling problem. The
biological classification In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mecha ...
introduced by
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement Ennoblement is the conferring of nobility—the induction of an individual into the noble social class, class. Currently only a few kingdoms still grant nob ...

Carl Linnaeus
in 1735 also viewed species as fixed according to the divine plan. In 1766, Georges Buffon 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 The Ussher chronology is a 17th-century chronology Chronology (from Latin ''chronologia'', from Ancient Greek , ''chrónos'', "time"; and , ''wikt:-logia, -logia'') is the science of arranging events in their order of occurrence in time. Co ...
of the 1650s had calculated creation at 4004 BC, but by the 1780s geologists assumed a much older world.
Werner
Werner
ians thought
strata In geology and related fields, a stratum (plural: strata) is a layer of sedimentary rock or soil, or igneous rock that was formed at the Earth's surface, with internally consistent characteristics that distinguish it from other layers. The "str ...
were deposits from shrinking seas, but
James Hutton James Hutton (; 3 June 172614 June 1726 New Style Old Style (O.S.) and New Style (N.S.) indicate a dating system from before and after a calendar change, respectively. Usually this is the change from the Julian calendar The Julian c ...

James Hutton
proposed a self-maintaining infinite cycle, anticipating
uniformitarianism at Jedburgh.Above: John Clerk of Eldin's 1787 illustration.Below: 2003 photograph. Uniformitarianism, also known as the Doctrine of Uniformity or the Uniformitarian Principle, is the assumption that the same natural laws and processes that operat ...
. 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 philosophy, natural philosopher, physiology, physiologist, Society for Effecting the ...
outlined a hypothesis of
transmutation of species Transmutation of species and transformism are 18th and 19th-century evolutionary ideas about the change of one species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism ...
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 Natural history is a domain of inquiry involving organisms, including animals, fu ...

Jean-Baptiste Lamarck
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 In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system that embodies ...

Lamarckism
. 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
era An era is a span of time defined for the purposes of chronology 222px, Joseph Scaliger's ''De emendatione temporum'' (1583) began the modern science of chronology Chronology (from Latin Latin (, or , ) is a classical language belonging t ...

era
s 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 Natural history is a domain of inquiry involving organism In biology, an organism () is any orga ...

Georges Cuvier
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 apologetics, Christian apologist, philosopher, and Utilitarianism, utilitarian. He is best known for his natural theology exposition of the teleological argument for the ex ...
'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 In the United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain,Usage is mixed. The Guardian' and Telegraph' u ...
and
Cambridge Cambridge ( ) is a university city and the county town In the United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain,Usage is mixed. The Guardian' and Telegraph' ...
) were
Church of England The Church of England (C of E) is a Christian church Christian Church is a Protestant Protestantism is a form of Christianity that originated with the 16th-century Reformation, a movement against what its followers perceived to be Critic ...
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 is the flood myth found in chapters 6–9 of the Book of Genesis in the Hebrew Bible and the Christian Old Testament. The story tells of God's decision to return the Earth to its pre-creation state of watery Chaos (co ...
. Some anatomists such as
Robert Grant
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 , latin_name = Universitas Academica Edinburgensis , image_name = University of Edinburgh ceremonial roundel.svg , image_size = 150px , established = , type = Public university/ancient university , endowment = Pounds sterling, £487.7 mil ...
in 1825 to study medicine. In his second year he neglected his medical studies for
natural history Natural history is a domain of inquiry involving organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecul ...

natural history
and spent four months assisting Robert Grant's research into
marine invertebrate Marine invertebrates are the invertebrate Invertebrates are animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This includes all animals apart from the subphyl ...
s. Grant revealed his enthusiasm for the transmutation of species, but Darwin rejected it. Starting in 1827, at
Cambridge University The University of Cambridge is a collegiate university, collegiate research university in Cambridge, Cambridge, United Kingdom. Founded in 1209 and granted a royal charter by Henry III of England, Henry III in 1231, Cambridge is the second-oldest ...
, Darwin learnt science as
natural theology Natural theology, once also termed physico-theology, is a type of theology Theology is the systematic study of the nature of the divine Divinity or the divine are things that are either related to, devoted to, or proceeding from a deity.
from botanist
John Stevens Henslow John Stevens Henslow (6 February 1796 – 16 May 1861) was a British people, British priest, botanist and geologist. He is best remembered as friend and mentor to his pupil Charles Darwin. Early life Henslow was born at Rochester, Kent, the s ...

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 wor ...
and
Alexander von Humboldt Friedrich Wilhelm Heinrich Alexander von Humboldt (14 September 17696 May 1859) was a , , , , and proponent of philosophy and . He was the younger brother of the Prussian minister, philosopher, and (1767–1835). Humboldt's quantitative work ...

Alexander von Humboldt
. Filled with zeal for science, he studied catastrophist
geology Geology (from the Ancient Greek Ancient Greek includes the forms of the Greek language Greek ( el, label=Modern Greek Modern Greek (, , or , ''Kiní Neoellinikí Glóssa''), generally referred to by speakers simply as Greek ...

geology
with
Adam Sedgwick Adam Sedgwick (; 22 March 1785 – 27 January 1873) was a British geologist A geologist is a scientist who studies the solid, liquid, and gaseous matter that constitutes Earth and other terrestrial planets, as well as the processes that shap ...

Adam Sedgwick
. 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 Earth is the third planet from the Sun and the only astro ...

Charles Lyell
's ''
Principles of Geology A principle is a proposition or value that is a guide for behavior or evaluation. In law, it is a Legal rule, rule that has to be or usually is to be followed. It can be desirably followed, or it can be an inevitable consequence of something, such ...
'' 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 armadillo
huge armadillo
s, and noted the geographical distribution of modern species in hope of finding their "centre of creation". The three
Fuegian Fuegians are one of the three tribes of indigenous inhabitants of Tierra del Fuego, at the southern tip of South America. In English, the term originally referred to the Yaghan people of Tierra del Fuego. In Spanish, the term ''fueguino'' can refe ...
missionaries the expedition returned to
Tierra del Fuego #REDIRECT Tierra del Fuego #REDIRECT Tierra del Fuego#REDIRECT Tierra del Fuego Tierra del Fuego (, ; Spanish for "Land of Fire", formerly also Fireland in English) is an archipelago An archipelago ( ), sometimes called an island group or i ...

Tierra del Fuego
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. Despite being a controversial figure, Owen is generally considered to have been an outstanding naturalist with a remark ...

Richard Owen
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 FRS FRS may also refer to: Government and politics * Facility Registry System, a centrally managed Environmental Protection Agency database that identifies places of environmental interest in the United States * Family Resources ...

John Gould
announced that Darwin's rhea was a separate species from the previously described rhea (though their territories overlapped), that
mockingbird Mockingbirds are a group of New World The "New World" is a term for the majority of Earth's Western Hemisphere, specifically the Americas."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: Oxf ...

mockingbird
s collected on the
Galápagos Islands The Galápagos Islands (official name: , other Spanish language, Spanish name: , , ), part of the Republic of Ecuador, are an archipelago of High island, volcanic islands. They are distributed on either side of the equator in the Pacific Ocean ...
represented three separate species each unique to a particular island, and that several distinct birds from those islands were all classified as
finches The true finches are small to medium-sized passerine birds in the family (biology), family Fringillidae. Finches have stout conical bills adapted for eating seeds and nuts and often have colourful plumage. They occupy a great range of habitats wh ...
. 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 (from el, γενεαλογία ' "the making of a pedigree") 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 ob ...

genealogical
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 A diagram is a symbolic representation Representation may refer to: Law a ...
, discarding Lamarck's independent lineages progressing to higher forms. Unconventionally, Darwin asked questions of
fancy pigeon Fancy pigeon refers to any breed of domestic pigeon, which is a Domestication, domesticated form of the wild rock dove (''Columba livia''). They are bred by Pigeon keeping, pigeon fanciers for various traits relating to size, shape, color, and behav ...
and animal breeders 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 The Hominidae (), whose members are known as great apes or hominids (), are a taxonomic family In human society A society is a Social group, group of individuals involved in persistent Social relati ...

orangutan
seemed. In late September 1838, he started reading
Thomas Malthus Thomas Robert Malthus (; 13/14 February 1766 – 23 December 1834) was an English cleric Clergy are formal leaders within established religion Religion is a - of designated and practices, , s, s, , , , , or , that relates humanit ...

Thomas Malthus
's ''
An Essay on the Principle of Population The book ''An Essay on the Principle of Population'' was first published anonymously in 1798, but the author was soon identified as Thomas Robert Malthus Thomas Robert Malthus (; 13/14 February 1766 – 23 December 1834) was an English cler ...

An Essay on the Principle of Population
'' with its statistical argument that human populations, if unrestrained, breed beyond their means and
struggle to survive
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 Charl ...
'' 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 United Kingdom of Great Britain and Ireland, British botanist and explorer in the 19th century. He was a founder of geographical botany and Charles Darwin's closest friend ...

Joseph Dalton Hooker
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 called pop-science or popsci) is an interpretation of science Science () is a systematic enterprise that Scientific method, builds and organizes knowledge in the form of Testability, testable explanations and predic ...

popular science
book ''
Vestiges of the Natural History of Creation ''Vestiges of the Natural History of Creation'' is an 1844 work of speculative natural history Natural history is a domain of inquiry involving organisms, including animals, fungus, fungi, and plants, in their natural environment, leaning mor ...
'', 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 , adaptation has three related meanings. Firstly, it is the dynamic evolutionary process that fits s to their environment, enhancing their . Secondly, it is a state reached by the population during that process. Thirdly, it is a or adapti ...

adaptation
. 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, biologist Francesco Redi, the founder of biology, is recognized to be one of the greatest biologists of all time A biologist is a professional who has speciali ...

Herbert Spencer
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 (general) is the practice and science of classification of things or concepts, including the principles that underlie such classification. The term may also refer to a specific classification scheme. Originally used only about biological ...
, Darwin began an eight-year study of
barnacle A barnacle is a type of arthropod An arthropod (, (gen. ποδός)) is an invertebrate animal having an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Euarthropoda,Reference ...

barnacle
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 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 Ecology (from el, οἶκος, "house" and el, -λογία, label=none, "study of") is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms ...

ecological niche
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 Fancy pigeon refers to any breed of domestic pigeon, which is a Domestication, domesticated form of the wild rock dove (''Columba livia''). They are bred by Pigeon keeping, pigeon fanciers for various traits relating to size, shape, color, and behav ...
breeding, and experimented (with the help of his son
Francis Francis may refer to: People *Pope Francis Pope Francis ( la, Franciscus; it, Francesco; es, link=, Francisco; born Jorge Mario Bergoglio, 17 December 1936) is the head of the Catholic Church and sovereign of the Vatican City State since ...
) 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 Darw ...
caused repeated delays. His paper on
Glen Roy Glen Roy ( gd, Gleann Ruaidh, meaning "red glen") in the Lochaber area of the Scottish Highlands, Highlands of Scotland is a glen noted for the geological phenomenon of three loch terrace (geology), terraces known as the Parallel Roads of Gle ...
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 Science (from the Latin word ''scientia'', meaning "knowledge") is a systematic enterprise that Scientific method, builds and Taxonomy (general), organizes knowledge in the form of ...

David Quammen
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 Charles Robert Darwin (; 12 February 1809 – 19 April 1882) was an English naturalist Natural history is a domain of inquiry involving o ...
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 18237 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 throug ...
, 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 Earth is the third planet from the Sun and the only astro ...

Charles Lyell
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 United Kingdom of Great Britain and Ireland, British botanist and explorer in the 19th century. He was a founder of geographical botany and Charles Darwin's closest friend ...

Joseph Dalton Hooker
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 right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
''. His theory including the principle of
divergence In vector calculus Vector calculus, or vector analysis, is concerned with derivative, differentiation and integral, integration of vector fields, primarily in 3-dimensional Euclidean space \mathbb^3. The term "vector calculus" is sometimes ...
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 botany, botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutual ...
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 right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
'', when on 18 June 1858 he received a parcel from Wallace, who stayed on the
Maluku Islands The Maluku Islands or the Moluccas () (''Molukken'') are an archipelago An archipelago ( ), sometimes called an island group or island chain, is a chain, cluster or collection of island An island (or isle) is an isolated piec ...

Maluku Islands
(
Ternate Ternate is a city A city is a large human settlement.Goodall, B. (1987) ''The Penguin Dictionary of Human Geography''. London: Penguin.Kuper, A. and Kuper, J., eds (1996) ''The Social Science Encyclopedia''. 2nd edition. London: Routledge. It ...

Ternate
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 to 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 (biology), taxonomy. It possesses several important biological specimen, manuscript and ...
, 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 "On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection" is the title of a journal article, comprising and resulting from the joint presentation of two scientific papers to the ...
'', 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 This Chihuahua (dog), Chihuahua mixed-breed dog, mix and Great Dane shows the wide range of dog breed sizes created using selective breeding. Selective breeding (also called artificial selection) is the process by which humans use animal breed ...
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 Animal breeding is a branch of animal science Animal science (also bioscience) is described as "studying the biology Biology i ...
, 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 more conventional level of the individual. Early authors such as V. C. Wynne-Edwards and Konrad Lorenz argued that ...
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 (biology), taxonomy. It possesses several important biological specimen, manuscript and ...
, 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 A seaside resort is a resort town, town, village, or hotel that serves as a Resort, vacation resort and is located on a coast. Sometimes the concept includes an aspect of official accreditation based on the satisfact ...
, 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 right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
''.


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 MurrayJohn Murray or John Murry may refer to: Arts and media Literature and music *John Murray (publishing house), a British publishing house, founded by John Murray (1745–1793) *John Murray (publisher, born 1778) (died 1843), second head of the pub ...

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.LTranscribed 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 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 In , family (from la, familia) is a of people related either by (by recognized birth) or (by marriage or other relationship). The purpose of families is to maintain the well-being of its members and o ...

pigeon
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: Science and technology Mathematics * Algebraic variety Algebraic varieties are the central objects of study in algebraic geometry Algebraic geometry is a branch of mathematics Mathematics (from Ancient Greek, ...
"'', meaning varieties within a
species In biology, a species is the basic unit of biological classification, 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 individu ...

species
. 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 A currency, "in circulation", from la, currens, -entis, literally meaning "running" or "traversing" in the most specific sense is money Image:National-D ...
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 The Worshipful Company of Stationers and Newspaper Makers (until 1937 the Worshipful Company of Stationers), usually known as the Stationers' Company, is one of the livery company, livery companies of the City of London. The Stationers' Company ...
copyright, around 1,170 copies were available for sale. Significantly, 500 were taken by , 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 working me ...
, 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'', while the fourth 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 Darwinism is a theory A theory is a reason, rational type of abstraction, abstract thinking about a phenomenon, or the results of such thinking. The process of contemp ...
", which had been coined by the philosopher
Herbert Spencer Herbert Spencer (27 April 1820 – 8 December 1903) was an English philosopher, biologist Francesco Redi, the founder of biology, is recognized to be one of the greatest biologists of all time A biologist is a professional who has speciali ...

Herbert Spencer
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 A biologist is a professional who has specialized knowledge in the field of biology Biology is the natural science that studies life and ...
'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 first principles of being, identity and change, space and time, causality, necessity and possibility. It includes questions about the nature of consciousness and the relationship between ...

metaphysics
. 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 Heredity, also called inheritance or biological inheritance, is the passing on of Phenotypic trait, traits from parents to their offspring; either through asexual reproduction or sexual reproduction, ...

evolution
" which had commonly been associated with embryological development, 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 a non-metropolitan and ceremonial county The counties and areas for the purposes of the lieutenancies, also referred to as the lieutenancy areas of England and informally known as ceremonial co ...

Lancashire
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 A metal (from Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approxim ...

font
. 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 botany, botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutual ...
, 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 Copyright is a type of intellectual property Intellectual property (IP) is a category of property Property is a system of rights that gives people legal control of valuable things, and also refers to the valuable things themselves. ...

copyright
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 Heinrich Georg Bronn (3 March 1800 – 5 July 1862) was a German geologist A geologist is a scientist who studies the solid, liquid, and gaseous matter that constitutes the Earth and other terrestrial planets, as well as t ...

Heinrich Georg Bronn
, 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 philosophy, philosophical tradition of German idealism, as applied to the study of nature in the earlier 19th century ...
''. 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, who published an improved translation in 1867. Darwin's attempts to find a translator in France fell through, and the translation by Clémence Royer published in 1862 added an introduction praising Darwin's ideas as an alternative to religious revelation and promoting ideas anticipating social Darwinism and eugenics, 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 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 and Francis Bacon on the theology of Physical law, natural laws, harmonising science and religion in accordance with Isaac Newton's belief in a rational God who established a law-abiding cosmos. In the second edition, Darwin added an epigraph from Joseph Butler affirming that God could work through scientific laws as much as through miracles, 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 wor ...
'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, and to that of the Galápagos tortoises and
mockingbird Mockingbirds are a group of New World The "New World" is a term for the majority of Earth's Western Hemisphere, specifically the Americas."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: Oxf ...

mockingbird
s. 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 and plant breeding, 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 Animal breeding is a branch of animal science Animal science (also bioscience) is described as "studying the biology Biology i ...
. As an illustration of
artificial selection This Chihuahua (dog), Chihuahua mixed-breed dog, mix and Great Dane shows the wide range of dog breed sizes created using selective breeding. Selective breeding (also called artificial selection) is the process by which humans use animal breed ...
, he describes
fancy pigeon Fancy pigeon refers to any breed of domestic pigeon, which is a Domestication, domesticated form of the wild rock dove (''Columba livia''). They are bred by Pigeon keeping, pigeon fanciers for various traits relating to size, shape, color, and behav ...
breeding, noting that "[t]he diversity of the breeds is something astonishing", yet all were descended from one species of rock pigeon. Darwin saw two distinct kinds of variation: (1) rare abrupt changes he called "sports" or "monstrosities" (example: Ancon sheep 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 biological classification, 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 individu ...

species
and Variety (biology), varieties 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 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 right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
"; 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, biologist Francesco Redi, the founder of biology, is recognized to be one of the greatest biologists of all time A biologist is a professional who has speciali ...

Herbert Spencer
, of the Survival of the Fittest, 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 Augustin Pyramus de Candolle, 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 Earth is the third planet from the Sun and the only astro ...

Charles Lyell
had stated that all organisms are exposed to severe competition. Darwin emphasizes that he used the phrase "The struggle for existence, struggle for existence" 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 ecology, ecological 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 Ecology (from el, οἶκος, "house" and el, -λογία, label=none, "study of") is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms ...

ecological niche
. 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, driven by competition between males for mates, to explain sexual dimorphism, sexually dimorphic 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'' (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 Phylogenetic tree, 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 (science), Tree of life ... with its ever branching and beautiful ramifications".


Variation and heredity

In Darwin's time there was no agreed-upon model of heredity; 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 In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system that embodies ...

Lamarckism
), 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 Evolutionary biology is the subfield of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chem ...
. He recounted how Lord Morton's mare apparently demonstrated telegony (pregnancy), telegony, 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'', which tried to explain heredity through his hypothesis of pangenesis. Although Darwin had privately questioned blending inheritance, 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 Natural history is a domain of inquiry involving organism In biology, an organism () is any organic, life, living system that fu ...

Charles Darwin
's theories of evolution with Gregor Mendel'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. This difficulty can be referred to as the absence or rarity of transitional varieties in habitat space. 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, 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 evolution of the eye, 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 Domestication, domesticated breeds of animals, and Race (human categorization), human races. He suggests that sexual selection 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 ants 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 A biologist is a professional who has specialized knowledge in the field of biology Biology is the natural science that studies life and ...
's criticisms, including his claim that features such as baleen filters in whales, flatfish with both eyes on one side and the camouflage of Phasmatodea, stick insects 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 appears to show forms of life suddenly arising, without the innumerable transitional fossils 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 Earth is the third planet from the Sun and the only astro ...

Charles Lyell
's argument in ''
Principles of Geology A principle is a proposition or value that is a guide for behavior or evaluation. In law, it is a Legal rule, rule that has to be or usually is to be followed. It can be desirably followed, or it can be an inevitable consequence of something, such ...
'' that the record is extremely imperfect as fossilisation 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 Formation (stratigraphy), geological formations, 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 Weald Basin, The Weald as discussed in ''Principles of Geology'' together with other observations from Hugh Miller, James Smith of Jordanhill and Andrew Ramsay (geologist), 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, 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 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 phyletic gradualism, the same rate – some organisms such as ''Lingula (genus), 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 Lamarckism, Lamarckian laws of inevitable progress. It has been argued that this anticipated the punctuated equilibrium 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. Despite being a controversial figure, Owen is generally considered to have been an outstanding naturalist with a remark ...

Richard Owen
'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 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, 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 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 (biology), Taxonomy'', 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 (biology), morphology, including the importance of homology (biology), homologous structures. 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. Despite being a controversial figure, Owen is generally considered to have been an outstanding naturalist with a remark ...

Richard Owen
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 Evolutionary biology is the subfield of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chem ...
. He notes that animals of the same class often have extremely similar embryology, embryos. 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, 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 Human evolution, 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 right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
had been the chief agent of change. He knew that his readers were already familiar with the concept of transmutation of species from ''Vestiges of the Natural History of Creation, 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. 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 wor ...
, 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 Science (from the Latin word ''scientia'', meaning "knowledge") is a systematic enterprise that Scientific method, builds and Taxonomy (general), organizes knowledge in the form of ...

David Quammen
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 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 right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
'' 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 MurrayJohn Murray or John Murry may refer to: Arts and media Literature and music *John Murray (publishing house), a British publishing house, founded by John Murray (1745–1793) *John Murray (publisher, born 1778) (died 1843), second head of the pub ...

John Murray
assurances "That I do not discuss origin of man". In the final chapter of ''On the Origin of Species'', "#Concluding remarks, 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 [Light will be thrown on the origin of man and his history] I 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 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 trap, Malthusian population growth. In his discussions on Morphology (biology), morphology, Darwin compares and comments on bone structures that are Homology (biology), 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 Race (human classification), races of humans differing over ideas of beauty. In his 1856 notes responding to Robert Knox's ''The Races of Man: A Fragment'', he called this effect sexual selection. 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'' 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. Samuel Wilberforce 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'', 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' was used to cover the whole range of evolutionism, 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 became central to the debate and was strongly argued by Thomas Henry Huxley, Huxley who featured it in his popular "working-men's lectures". Darwin did not publish his own views on this until 1871. The naturalism (philosophy), naturalism of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
conflicted with presumptions of teleology, purpose in nature and while this could be reconciled by theistic evolution, other mechanisms implying more progress or purpose were more acceptable.
Herbert Spencer Herbert Spencer (27 April 1820 – 8 December 1903) was an English philosopher, biologist Francesco Redi, the founder of biology, is recognized to be one of the greatest biologists of all time A biologist is a professional who has speciali ...

Herbert Spencer
had already incorporated
Lamarckism Lamarckism, also known as Lamarckian inheritance or neo-Lamarckism, is the notion that an organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system that embodies ...

Lamarckism
into his popular philosophy of progressive free market human society. He popularised the terms 'evolution' and '
survival of the fittest "Survival of the fittest" is a phrase that originated from Darwinian Darwinism is a theory A theory is a reason, rational type of abstraction, abstract thinking about a phenomenon, or the results of such thinking. The process of contemp ...
', 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 Physical law, 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 right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
as a scientifically testable mechanism while accepting that other mechanisms such as inheritance of acquired characters 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 Philosophy (from , ) is the study of general and fundamental questions, such as those about Metaphysics, existence, reason, Epistemology, knowledge, Ethics, values, Philosophy of mind, mind, and Philosophy of language, l ...
of John Stuart Mill's ''A System of Logic'', while opponents held to the idealist school of William Whewell'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 and ecology, including
Joseph Dalton Hooker Sir Joseph Dalton Hooker (30 June 1817 – 10 December 1911) was a United Kingdom of Great Britain and Ireland, British botanist and explorer in the 19th century. He was a founder of geographical botany and Charles Darwin's closest friend ...

Joseph Dalton Hooker
in 1860, and
Asa Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botany, botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutual ...
in 1862. Henry Walter Bates presented research in 1861 that explained Batesian mimicry, insect mimicry using natural selection.
Alfred Russel Wallace Alfred Russel Wallace (8 January 18237 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 throug ...
discussed evidence from his Malay archipelago research, including an 1864 paper with an evolutionary explanation for the Wallace line. Evolution had less obvious applications to anatomy and morphology (biology), morphology, and at first had little impact on the research of the anatomist Thomas Henry Huxley. 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 was sufficient without Saltation (biology), sudden leaps to cause speciation. Huxley wanted science to be secular, without religious interference, and his article in the April 1860 ''Westminster Review'' promoted naturalism (philosophy), 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" 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 A dining club (UK) or eating club (US) is a Social club, social group, usually requiring membership (which may, or may not be available only to certain people), which meets for dinners and discussion on a regular b ...
, which used the journal ''Nature (journal), Nature'' to promote evolution and naturalism, shaping much of late-Victorian science. Later, the German morphologist Ernst Haeckel would convince Huxley that comparative anatomy and paleontology, palaeontology could be used to reconstruct phylogenetics, 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. Despite being a controversial figure, Owen is generally considered to have been an outstanding naturalist with a remark ...

Richard Owen
, 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 teleology, teleological 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 George Campbell, 8th Duke of Argyll, Duke of Argyll 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 meeting featuring the legendary 1860 Oxford evolution debate. 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 working me ...
satirised as the "Great Hippocampus Question" and parodied in ''The Water-Babies, A Fairy Tale for a Land Baby, 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 Earth is the third planet from the Sun and the only astro ...

Charles Lyell
and
Alfred Russel Wallace Alfred Russel Wallace (8 January 18237 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 throug ...
, 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 (biology), homology in morphology (biology), morphology from Johann Wolfgang von Goethe, Goethe's ''Metamorphosis of Plants'' and from their long tradition of comparative anatomy. Heinrich Georg Bronn, Bronn's alterations in his German translation added to the misgivings of conservatives, but enthused political radicals. Ernst Haeckel was particularly ardent, aiming to synthesise Darwin's ideas with those of Lamarckism, Lamarck 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 philosophy, philosophical tradition of German idealism, as applied to the study of nature in the earlier 19th century ...
''. Their ambitious programme to reconstruct the evolutionary history of life was joined by Huxley and supported by discoveries in :paleontology, palaeontology. Haeckel used embryology extensively in his recapitulation theory, which embodied a progressive, almost linear model of evolution. Darwin was cautious about such histories, and had already noted that Karl Ernst von Baer, 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 botany, botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutual ...
promoted and defended ''Origin'' against those American naturalists with an idealist approach, notably Louis Agassiz 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 and Alpheus Hyatt reconciled this view with evolutionism in a form of Lamarckism#Neo-Lamarckism, neo-Lamarckism involving recapitulation theory. French-speaking naturalists in several countries showed appreciation of the much-modified French translation by Clémence Royer, 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 Malthusianism, Malthusian aspects were felt to be relatively unimportant. The political economy of struggle was criticised as a British stereotype by Karl Marx and by Leo Tolstoy, who had the character Levin in his novel ''Anna Karenina'' 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 right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
as the key mechanism of evolution, including Karl Wilhelm von Nägeli, Karl von Nägeli'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 allowed gradual evolution was disputed by William Thomson, 1st Baron Kelvin, William Thomson (later awarded the title Lord Kelvin), who calculated that it had cooled in less than 100 million years. Darwin accepted blending inheritance, 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. St. George Jackson Mivart, Mivart 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 Sir Julian Sorell Huxley (22 June 1887 – 14 February 1975) was an English evolutionary biologist, eugenicist, and internationalist. He was a proponent of natural selection Natural selection is the differential surv ...
" included forms of "Saltation (biology), saltationism" in which new species were thought to arise through "jumps" rather than gradual adaptation, forms of orthogenesis 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, that natural selection was the only mechanism, was called neo-Darwinism. It was thought that the rediscovery of Mendelian inheritance 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 18237 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 throug ...
, 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'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 to provide scientific evidence for the superiority of Caucasian race, Caucasians over non-white races and justify European imperialism. 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, 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. history of geology, Developments in geology meant that there was little opposition based on a literal reading of Book of Genesis, Genesis, but defence of the teleological argument, argument from design and
natural theology Natural theology, once also termed physico-theology, is a type of theology Theology is the systematic study of the nature of the divine Divinity or the divine are things that are either related to, devoted to, or proceeding from a deity.
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 Christianity, 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 (mathematician), Baden Powell praised "Mr Darwin's masterly volume [supporting] the 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 botany, botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutual ...
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, ''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 were favoured over
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
as being more compatible with purpose. Even though the book did not explicitly spell out Darwin's beliefs about human evolution, 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 (spirit), soul, 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 argued in his 1874 critique "What is Darwinism?" that "Darwinism", 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'' were explicitly open to the possibility that God created through evolution, but fundamentalism inspired the American creation–evolution controversy that began in the 1920s. Some conservative Roman Catholic writers and influential Jesuits 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 Christian theology is the theology Theology is the systematic study of the nature of the Divinity, di ...
had not interpreted Genesis literally in this area. The Holy See, Vatican stated its Evolution and the Roman Catholic Church, official position in a Humani generis, 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 Sir Julian Sorell Huxley (22 June 1887 – 14 February 1975) was an English evolutionary biologist, eugenicist, and internationalist. He was a proponent of natural selection Natural selection is the differential surv ...
" became untenable as more was learned about inheritance and mutation. The full significance of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype right , Here the relation between genotype and phenotype is illustrated, using a Punnett square, for the character of peta ...
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, Sewall Wright and J. B. S. Haldane, merged Darwinian selection with a statistical understanding of Mendelian genetics. evolution, Modern evolutionary theory 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 Evolutionary biology is the subfield of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chem ...
, has become the unifying theory of the
life science Life is a characteristic that distinguishes physical entities that have biological processes, such as signaling In signal processing Signal processing is an electrical engineering subfield that focuses on analysing, modifying, a ...
s. The theory explains the diversity of living organisms and their adaptation to the environment. It makes sense of the geological record, biogeography, parallels in embryonic development, Homology (biology), biological homologies, vestigiality, cladistics, phylogenetics and other fields, with unrivalled explanatory power; it has also become essential to applied sciences such as evolutionary medicine, medicine and agriculture. Despite the scientific consensus, a religion-based political creation–evolution controversy, controversy 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, about his life and work. The text of ''Origin'' itself has been subject to much analysis including a variorum, detailing the changes made in every edition, first published in 1959, and a Concordance (publishing), 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

* s:On the Origin of Species (1859), ''On the Origin of Species'' – full text at Wikisource of the first edition, 1859 * s:The Origin of Species (1872), ''The Origin of Species'' – full text at Wikisource of the 6th edition, 1872 * Charles Darwin bibliography * History of biology * History of evolutionary thought * 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'', published in 1871; his second major book on evolutionary theory. * Transmutation of species


References


Works cited

* * * * * * * *. Published anonymously. * * * * * * * * * * * * * * * * * * * * 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 {{DEFAULTSORT:Origin Of Species, The 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 Science textbooks