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			{{Short description|English naturalist and biologist (1809–1882)}}
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	{{Infobox scientist
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	|name = Charles Darwin
			{{Use British English|date=October 2024}}
	|image = Charles_Darwin_seated.jpg
			{{Use dmy dates|date=October 2024}}
	|alt=Three quarter length studio photo showing Darwin's characteristic large forehead and bushy eyebrows with deep set eyes, pug nose and mouth set in a determined look. He is bald on top, with dark hair and long side whiskers but no beard or moustache. His jacket is dark, with very wide lapels, and his trousers are a light check pattern. His shirt has an upright wing collar, and his cravat is tucked into his waistcoat which is a light fine checked pattern.
			{{Infobox person
	|caption = Charles Robert Darwin, aged 45 in 1854, by then working towards publication of '']''.
			| name = Charles Darwin
	|birth_date = {{birth date|1809|2|12|df=y}}
			| honorific_suffix = {{post-nominals|country=GBR|size=100%|JP|FRS|FRGS|FLS|FZS}}
	|birth_place = ], ], ], ]
			| image = Charles Darwin seated crop.jpg
	|death_date = {{death date and age|1882|4|19|1809|2|12|df=yes}}
			| alt = Three quarter length studio photo showing Darwin's characteristic large forehead and bushy eyebrows with deep set eyes, pug nose and mouth set in a determined look. He is bald on top, with dark hair and long side whiskers but no beard or moustache. His jacket is dark, with very wide lapels, and his trousers are a light check pattern. His shirt has an upright wing collar, and his cravat is tucked into his waistcoat which is a light fine checked pattern.
	|death_place = ], ], ], England
			| caption = Darwin, {{circa|1854}}, when he was preparing '']''{{sfn|Freeman|2007|p=76}}
	|residence = ]
			| birth_name = Charles Robert Darwin
	|citizenship = ]
			| birth_date = {{birth date|df=y|1809|2|12}}
	|nationality = ]
			| birth_place = ], ], England
	|ethnicity = ]
			| death_date = {{death date and age|1882|df=y|4|19|1809|2|12}}
	|fields = ]
			| death_place = ], ], ], England<!--Letter no. 3368 footnote; "spelling of the village name was variable. 'Down' was the commonest form up to the 1870s, and 'Downe' thereafter", and it was in Kent until 1965, (now in ])-->
	|workplaces = ]
			| resting_place = ]
	|alma_mater = ]<br />]
			| education = {{ubli|
	|doctoral_advisor = <!--there were no PhDs in Cambridge at the time-->
			]|
	|academic_advisors = ]<br />]
			] (], ])<ref>{{cite news|url=https://www.telegraph.co.uk/news/uknews/5032354/Charles-Darwins-personal-finances-revealed-in-new-find.html|archive-url=https://web.archive.org/web/20171019230458/http://www.telegraph.co.uk/news/uknews/5032354/Charles-Darwins-personal-finances-revealed-in-new-find.html|url-status=dead|title=Charles Darwin's personal finances revealed in new find|date=22 March 2009|archive-date=19 October 2017|work=The Telegraph
	|doctoral_students = <!--there were no PhDs in Cambridge at the time-->
			}}</ref>}}
	|notable_students =
	|known_for = '']''<br />'']''<br />]		| known_for = ]
			| spouse = {{marriage|]|1839}}
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			| children = 10, including ], ], ], ], ] and ]
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			| parents = {{ubl|]|]}}
	|influences = ]<br />]
			| family = ]
	|influenced = ]<br />]<br />]
			| awards = {{ubl|] (1839)<ref name="catalogues.royalsociety.org-2015">{{cite web | title=Search Results: Record – Darwin; Charles Robert | website=The Royal Society Collections Catalogues | date=20 June 2015 | url=https://catalogues.royalsociety.org/CalmView/Record.aspx?src=CalmView.Persons&id=NA8196&pos=1 | access-date=2 December 2021 | archive-date=2 December 2021 | archive-url=https://web.archive.org/web/20211202200044/https://catalogues.royalsociety.org/CalmView/Record.aspx?src=CalmView.Persons&id=NA8196&pos=1 | url-status=live }}</ref>|] (1853)<ref name="Freeman 2007">{{harvnb|Freeman|2007|p=}}.</ref>|] (1859)<ref name="Freeman 2007" />|] (1864)<ref name="Freeman 2007" />|{{lang|fr|]}} (1867)<ref name="Freeman 2007" />|] (1879)<ref name="Freeman 2007" />}}
	|awards = ] (1853)<br />] (1859)<br />] (1864)
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	|signature_alt = "Charles Darwin", with the last name underlined by a downward curve that mimics the curve of the initial "C"
			| notableworks = {{plainlist|
	|footnotes =
			* '']''
			* '']''
			* '']''}}
			| module = {{Infobox scientist
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			| author_abbrev_bot = '''Darwin'''
			| author_abbrev_zoo = '''Darwin'''
			| signature = Charles Darwin Signature.svg
			| signature_alt = "Charles Darwin", with the surname underlined by a downward curve that mimics the curve of the initial "C"
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	'''Charles Robert Darwin''' ] (12 February 1809 – 19 April 1882) was an ] ]{{Ref_label|A|I|none}} who proposed that all ] of life have ] over time from ], and published compelling supporting evidence of this in his 1859 book '']'' in which he presented his ] that this branching pattern of evolution resulted from a process that he called ].<ref>{{cite book |title=Why Evolution is True |last = Coyne |first=Jerry A. |authorlink=Jerry Coyne |year=2009 |publisher=] |location= Oxford |isbn=0-199-23084-6 |page=17 |quote=In ''The Origin'', Darwin provided an alternative hypothesis for the development , diversification, and design of life. Much of that book presents evidence that not only supports evolution but at the same time refutes creationism. In Darwin's day, the evidence for his theories was compelling but not completely decisive.}}</ref><ref>{{cite book |title=Forerunners of Darwin |last=Glass |first=Bentley |authorlink=Bentley Glass |coauthors= |year=1959 |publisher=Johns Hopkins University Press |location=Baltimore, MD |isbn= 0801802229|pages= |page=iv |quote=Darwin's solution is a magnificent synthesis of evidence...a synthesis...compelling in honesty and comprehensiveness}}</ref>
	The ] became accepted by the ] and much of the general public in his lifetime,<ref name=JvW>{{Harvnb|van Wyhe|2008}}</ref> but it was not until the emergence of the ] from the 1930s to the 1950s that a broad consensus developed that natural selection was the basic mechanism of evolution.<ref name=b3847>{{harvnb|Bowler|2003|pp=338, 347}}</ref> In modified form, Darwin's scientific discovery is the unifying theory of the ], explaining the ].<ref> ''darwin-online.org.uk''. Retrieved on 2006-12-15<br />{{Harvnb|Dobzhansky|1973}}</ref><ref>As Darwinian scholar Joseph Carroll of the University of Missouri–St. Louis puts it in his introduction to a modern reprint of Darwin's work: "''The Origin of Species'' has special claims on our attention. It is one of the two or three most significant works of all time—one of those works that fundamentally and permanently alter our vision of the world....It is argued with a singularly rigorous consistency but it is also eloquent, imaginatively evocative, and rhetorically compelling." {{cite book |title=On the origin of species by means of natural selection |last= |first= |authorlink= |editor=Carroll, Joseph |year=2003 |publisher=Broadview |location= Peterborough, Ontario|isbn= 1551113376|page=15 |pages= |url= |accessdate= }}</ref>
	<!--Please consider discussing changes on the talk page, as this opening is the result of a very long consensus-building process.-->		<!--Please consider discussing changes on the talk page, as this opening is the result of a very long consensus-building process.-->
			'''Charles Robert Darwin''' ({{IPAc-en|ˈ|d|ɑr|w|ɪ|n}}<ref> {{webarchive|url=https://web.archive.org/web/20140718234042/http://www.collinsdictionary.com/dictionary/english/darwin |date=18 July 2014 }} entry in '']''.</ref> {{Respell|DAR|win}}; 12 February 1809&nbsp;– 19 April 1882) was an English ], ], and ],<ref>{{harvnb|Desmond|Moore|Browne|2004}}.</ref> widely known for his contributions to ]. His proposition that all species of life have descended from a ] is now generally accepted and considered a fundamental scientific concept.<ref>{{cite book |last=Coyne |first=Jerry A. |author-link=Jerry Coyne |title=Why Evolution is True |publisher=Viking |year=2009 |pages= |isbn=978-0-670-02053-9 |url=https://archive.org/details/whyevolutionistr00coyn/page/8}}</ref> In a joint presentation with ], he introduced his scientific theory that this ] of ] resulted from a process he called ], in which the ] has a similar effect to the artificial selection involved in ].<ref name="Larson 2004">{{Harvnb|Larson|2004|pp=79–111}}.</ref> Darwin has been described as one of the most influential figures in ] and was honoured by ].<ref>{{cite news |url=https://www.newscientist.com/special/darwin-200 |title=Special feature: Darwin 200 |access-date=2 April 2011 |work=New Scientist |url-status=live |archive-url=https://web.archive.org/web/20110211051412/http://www.newscientist.com/special/darwin-200 |archive-date=11 February 2011}}</ref><ref name="Westminster Abbey-2016" />
	Darwin's early interest in nature led him to neglect his course in ] at ] and instead help to investigate ], then the ] encouraged a passion for ].<ref name=whowas>{{Harvnb|Leff|2000|loc=}}</ref> His ] on {{HMS|Beagle}} established him as an eminent ] whose observations and theories supported ]'s ] ideas, and publication of his ] made him famous as a popular author.<ref>{{Harvnb|Desmond|Moore|1991|pp= 210, 284–285}}</ref>
			Darwin's early interest in nature led him to neglect his medical education at the ]; instead, he helped to investigate ]. His studies at the ]'s ] from 1828 to 1831 encouraged his passion for ].<ref name="Leff 2000">{{Harvnb|Leff|2000|loc=}}.</ref> However, it was his ] on {{HMS|Beagle}} from 1831 to 1836 that truly established Darwin as an eminent geologist. The observations and theories he developed during his voyage supported ]'s ]. Publication of his ] made Darwin famous as a popular author.<ref>{{Harvnb|Desmond|Moore|1991|pp=210, 284–285}}.</ref>
			Puzzled by the geographical distribution of wildlife and fossils he collected on the voyage, Darwin began detailed investigations and, in 1838, devised his theory of natural selection.<ref>{{Harvnb|Desmond|Moore|1991|pp=263–274}}.</ref> Although he discussed his ideas with several naturalists, he needed time for extensive research, and his geological work had priority.<ref>{{harvnb|van Wyhe|2007|pp=184, 187}}</ref> He was writing up his theory in 1858 when Alfred Russel Wallace sent him an essay that described the same idea, prompting the immediate joint submission of ] to the ].<ref>{{Cite journal | last1=Beddall | first1=B. G. | title=Wallace, Darwin, and the Theory of Natural Selection | journal=Journal of the History of Biology | volume=1 | issue=2 | pages=261–323 | year=1968 | doi=10.1007/BF00351923 | s2cid=81107747 | df=dmy-all |issn=0022-5010}}</ref> Darwin's work established evolutionary descent with modification as the dominant scientific explanation of natural diversification.<ref name="van Wyhe 2008" /> In 1871, he examined ] and ] in '']'', followed by '']'' (1872). His research on plants was published in a series of books, and in his final book, ] (1881), he examined ]s and their effect on soil.
			Darwin published his theory of evolution with compelling evidence in his 1859 book '']''.<ref>{{cite book |title=Why Evolution is True |last=Coyne |first=Jerry A. |author-link=Jerry Coyne |year=2009 |publisher=Oxford University Press |location=Oxford |isbn=978-0-19-923084-6 |page= |quote=In ''The Origin'', Darwin provided an alternative hypothesis for the development, diversification, and design of life. Much of that book presents evidence that not only supports evolution but at the same time refutes creationism. In Darwin's day, the evidence for his theories was compelling but not completely decisive. |url=https://archive.org/details/isbn_9780199230846/page/17 }}</ref><ref>{{cite book |title=Forerunners of Darwin |last=Glass |first=Bentley |author-link=Bentley Glass |year=1959 |publisher=Johns Hopkins University Press |location=Baltimore, MD |isbn=978-0-8018-0222-5 |page=iv |quote=Darwin's solution is a magnificent synthesis of evidence&nbsp;... a synthesis&nbsp;... compelling in honesty and comprehensiveness}}.</ref> By the 1870s, the scientific community and a majority of the educated public had accepted ]. However, many initially favoured ] that gave only a minor role to natural selection, and it was not until the emergence of the ] from the 1930s to the 1950s that a broad consensus developed in which natural selection was the basic mechanism of evolution.<ref name="van Wyhe 2008">{{Harvnb|van Wyhe|2008}}.</ref><ref name="Bowler 2003">{{harvnb|Bowler|2003|pp=178–179, 338, 347}}.</ref> Darwin's scientific discovery is the unifying theory of the ], explaining the ].
			==Biography==
			===Early life and education===
			{{further|Charles Darwin's education|Darwin–Wedgwood family}}
			Darwin was born in ], Shropshire, on 12 February 1809, at his family's home, ].<ref>{{cite encyclopedia |last=Desmond |first=Adrian J. |title=Charles Darwin |encyclopedia=Encyclopædia Britannica |date=13 September 2002 |url=https://www.britannica.com/biography/Charles-Darwin |access-date=11 February 2018 |archive-date=6 February 2018 |archive-url=https://web.archive.org/web/20180206114419/https://www.britannica.com/biography/Charles-Darwin |url-status=live}}</ref><ref>{{cite web |url=http://darwin.baruch.cuny.edu/biography/shrewsbury/mount/ |title=The Mount House, Shrewsbury, England (Charles Darwin) |author=John H. Wahlert |date=11 June 2001 |work=Darwin and Darwinism |publisher=] |access-date=26 November 2008 |url-status=dead |archive-url=https://web.archive.org/web/20081206010149/http://darwin.baruch.cuny.edu/biography/shrewsbury/mount/ |archive-date=6 December 2008}}</ref> He was the fifth of six children of wealthy society doctor and financier ] and ] (née Wedgwood). His grandfathers ] and ] were both prominent ]. Erasmus Darwin had praised general concepts of evolution and ] in his '']'' (1794), a poetic fantasy of gradual creation including undeveloped ideas anticipating concepts his grandson expanded.<ref>{{cite book |last=Smith |first=Homer W. |url=https://archive.org/details/manhisgods00smit |title=Man and His Gods |date=1952 |publisher=] |location=New York |pages= |author-link=Homer W. Smith |url-access=registration}}</ref>
			]. Part of ] showing him together with his sister Catherine.]]
			Both families were largely ], though the Wedgwoods were adopting ]. Robert Darwin, a ], had baby Charles ] in November 1809 in the Anglican ], but Charles and his siblings attended the ] with their mother. The eight-year-old Charles already had a taste for natural history and collecting when he joined the day school run by its preacher in 1817. That July, his mother died. From September 1818, he joined his older brother ] in attending the nearby Anglican ] as a ].<ref name="Desmond-3">{{Harvnb|Desmond|Moore|1991|pp=12–15}};<br />{{harvnb|Darwin|1958|pp=}}.</ref>
			Darwin spent the summer of 1825 as an apprentice doctor, helping his father treat the poor of Shropshire, before going to the well-regarded ] with his brother Erasmus in October 1825. Darwin found lectures dull and surgery distressing, so he neglected his studies.{{sfn|Darwin|1958|pp=}} He learned ] in around 40 daily hour-long sessions from ], a freed black slave who had accompanied ] in the South American ].<ref name="Darwin 1958-6">{{harvnb|Darwin|1958|p=}};<br />{{harvnb|Desmond|Moore|2009|pp=}}.</ref>
			In Darwin's second year at the university, he joined the ], a student ] group featuring lively debates in which ] students with ] views challenged orthodox religious concepts of science.{{sfn|Desmond|Moore|1991|pp=31–34}} He assisted ]'s investigations of the anatomy and life cycle of ] in the ], and on 27 March 1827 presented at the Plinian his own discovery that black spores found in ] shells were the eggs of a skate ]. One day, Grant praised ]'s ]. Darwin was astonished by Grant's audacity, but had recently read similar ideas in his grandfather Erasmus' journals.<ref>{{Harvnb|Browne|1995|pp=72–88}}.</ref> Darwin was rather bored by ]'s natural-history course, which covered geology{{snd}}including the debate between ] and ]. He learned the ] of plants and assisted with work on the collections of the ], one of the largest museums in Europe at the time.<ref>{{Harvnb|Desmond|Moore|1991|pp=42–43}}.</ref>
			Darwin's neglect of medical studies annoyed his father, who sent him to ], in January 1828, to study for a Bachelor of Arts degree as the first step towards becoming an Anglican country ]. Darwin was unqualified for Cambridge's '']'' exams and was required instead to join the ordinary degree course.<ref>{{Harvnb|Browne|1995|pp=47–48, 89–91}};<br />{{harvnb|Desmond|Moore|2009|pp=47–48}}.</ref> He preferred ] and ] to studying.{{sfn|Darwin|1887|p=}}
			] portrait by ] of Darwin as a student, in the courtyard at ], where he had rooms.<ref name="BBC News-2009">{{cite web | title=Darwin statue unveiled at college | website=BBC News | date=12 February 2009 | url=http://news.bbc.co.uk/1/hi/england/cambridgeshire/7886278.stm | access-date=22 April 2022 | archive-date=16 February 2009 | archive-url=https://web.archive.org/web/20090216155859/http://news.bbc.co.uk/1/hi/england/cambridgeshire/7886278.stm | url-status=live }}</ref>]]
			During the first few months of Darwin's enrolment at Christ's College, his second cousin ] was still studying there. Fox impressed him with his butterfly collection, introducing Darwin to ] and influencing him to pursue ] collecting.<ref name="Smith-1952">{{cite book |last=Smith |first=Homer W. |url=https://archive.org/details/manhisgods00smit |title=Man and His Gods |date=1952 |publisher=] |location=New York |pages= |author-link=Homer W. Smith |url-access=registration}}</ref><ref name="Darwin 1887">{{harvnb|Darwin|1887|pp=}}.</ref> He did this zealously and had some of his finds published in ]' ''Illustrations of British entomology'' (1829–1932).<ref name="Darwin 1887" /><ref>{{Cite web |editor-last=van Wyhe |editor-first=John |title=Darwin's insects in Stephens' Illustrations of British entomology (1829–32) |url=http://darwin-online.org.uk/EditorialIntroductions/vanWyhe_Stephens.html |url-status=dead |access-date=3 July 2020 |website=Darwin Online |archive-date=1 September 2019 |archive-url=https://web.archive.org/web/20190901090213/http://darwin-online.org.uk/EditorialIntroductions/vanWyhe_Stephens.html}}</ref>
			Through Fox, Darwin became a close friend and follower of botany professor ].<ref name="Smith-1952" /> He met other leading ]s who saw scientific work as religious ], becoming known to these ] as "the man who walks with Henslow". When his own exams drew near, Darwin applied himself to his studies and was delighted by the language and logic of ]'s ''Evidences of Christianity'' (1795).<ref name="Desmond">{{Harvnb|Desmond|Moore|1991|pp=73–79, 763}};<br />{{Harvnb|Darwin|1958|pp=}}.</ref> In his final examination in January 1831, Darwin did well, coming tenth out of 178 candidates for the ''ordinary'' degree.<ref>{{Harvnb|Browne|1995|p=97}}.</ref>
			Darwin had to stay at Cambridge until June 1831. He studied Paley's '']'' (first published in 1802), which made an ], explaining ] as God acting through ].<ref name="von Sydow 2005">{{Harvnb|von Sydow|2005|pp=5–7}}.</ref> He read ]'s new book, ''Preliminary Discourse on the Study of Natural Philosophy'' (1831), which described the highest aim of ] as understanding such laws through ] based on observation, and ]'s ''Personal Narrative'' of scientific travels in 1799–1804.<ref>{{ cite book | last=Daum | first=Andreas W.|author-link=Andreas Daum | year=2024 | title=Alexander von Humboldt: A Concise Biography | location=Trans. Robert Savage. Princeton, N.J. | publisher=Princeton University Press |pages=137–138 | isbn=978-0-691-24736-6 }}</ref> Inspired with "a burning zeal" to contribute, Darwin planned to visit ] with some classmates after graduation to study natural history in the tropics. In preparation, he joined ]'s geology course, then on 4 August travelled with him to spend a fortnight mapping ] in Wales.<ref name="Darwin 1958-5">{{Harvnb|Darwin|1958|pp=}}.</ref><ref>{{Harvnb|Browne|1995|pp=128–129, 133–141}}.</ref>
	Puzzled by the geographical distribution of wildlife and ]s he collected on the voyage, Darwin investigated the ] and conceived his theory of natural selection in 1838.<ref>{{Harvnb|Desmond|Moore|1991|pp=263–274}}</ref> Although he discussed his ideas with several naturalists, he needed time for extensive research and his geological work had priority.<ref>{{harvnb|van Wyhe|2007|pp=184, 187}}</ref> He was writing up his theory in 1858 when ] sent him an essay which described the same idea, prompting immediate joint publication of ].<ref> ]. Retrieved on 2007-03-21</ref> Darwin's work established evolutionary descent with modification as the dominant scientific explanation of diversification in nature.<ref name = JvW/> In 1871 he examined ] and ] in '']'', followed by '']''. His research on plants was published in a series of books, and in his final book, he examined ]s and their effect on soil.<ref>{{Harvnb|Freeman|1977}}</ref>
			===Survey voyage on HMS ''Beagle''===
	In recognition of Darwin's pre-eminence as a scientist, he was one of only five 19th-century UK non-royal personages to be honoured by a ],<ref name=BBCstatefuneral>{{cite news|url=http://news.bbc.co.uk/2/hi/uk_news/politics/7538482.stm |title=BBC NEWS : Politics : Thatcher state funeral undecided|date=2008-08-02|accessdate=2008-08-10}}</ref> and was buried in ], close to ] and ].<ref name=DarwinsBurial>{{Harvnb|Leff|2000|loc=}}</ref>
			{{further|Second voyage of HMS Beagle}}
			], 1831–1836]]
	==Life of Darwin==
	===Childhood and education===
	{{seealso|Charles Darwin's education|Darwin-Wedgwood family}}
	]
			After leaving Sedgwick in Wales, Darwin spent a few days with student friends at ]. He returned home on 29 August to find a letter from Henslow proposing him as a suitable (if unfinished) naturalist for a self-funded ] place on {{HMS|Beagle}} with captain ], a position for a ] rather than "a mere collector". The ship was to leave in four weeks on an expedition to chart the coastline of South America.<ref name="Peter Lucas-2010">{{cite web |url=http://darwin-online.org.uk/EditorialIntroductions/Lucas_Lowe_Journal.html |title=The recovery of time past: Darwin at Barmouth on the eve of the Beagle |author=Peter Lucas |date=1 January 2010 |website=Darwin Online |access-date=5 December 2021 |archive-date=11 April 2010 |archive-url=https://web.archive.org/web/20100411090308/http://darwin-online.org.uk/EditorialIntroductions/Lucas_Lowe_Journal.html |url-status=live }}</ref><ref>{{cite web|url=https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-105.xml|title=Letter no. 105, Henslow, J. S. to Darwin, C. R., 24 Aug 1831|website=Darwin Correspondence Project|access-date=29 December 2021|archive-date=29 December 2021|archive-url=https://web.archive.org/web/20211229182043/https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-105.xml|url-status=live}}</ref> Robert Darwin objected to his son's planned two-year voyage, regarding it as a waste of time, but was persuaded by his brother-in-law, ], to agree to (and fund) his son's participation.<ref>{{Harvnb|Desmond|Moore|1991|pp=94–97}}</ref> Darwin took care to remain in a private capacity to retain control over his collection, intending it for a major scientific institution.<ref name="Browne 1995">{{Harvnb|Browne|1995|pp=204–210}}</ref>
	Charles Robert Darwin was born in ], ], England on 12 February 1809 at his family home, ].<ref>{{cite web|url=http://darwin.baruch.cuny.edu/biography/shrewsbury/mount/|title=The Mount House, Shrewsbury, England (Charles Darwin)|author=John H. Wahlert|date=11 June 2001|work=Darwin and Darwinism|publisher=]|accessdate=2008-11-26}}</ref> He was the fifth of six children of wealthy society doctor and financier ], and ] (''née'' Wedgwood). He was the grandson of ] on his father's side, and of ] on his mother's side. Both families were largely ], though the Wedgwoods were adopting ]. Robert Darwin, himself quietly a ], had baby Charles ] in the Anglican Church, but Charles and his siblings attended the Unitarian chapel with their mother. The eight year old Charles already had a taste for natural history and collecting when he joined the day school run by its preacher in 1817. That July, his mother died. From September 1818, he joined his older brother ] attending the nearby Anglican ] as a ].<ref name=skool>{{Harvnb|Desmond|Moore|1991|pp= 12–15}}<br />{{harvnb|Darwin|1958|pp=}}</ref>
			After delays, the voyage began on 27 December 1831; it lasted almost five years. As FitzRoy had intended, Darwin spent most of that time on land investigating geology and making natural history collections, while HMS ''Beagle'' ] coasts.<ref name="van Wyhe 2008" /><ref name="Keynes 2000">{{harvnb|Keynes|2000|pp=}}</ref> He kept careful notes of his observations and theoretical speculations. At intervals during the voyage, his specimens were sent to Cambridge together with letters including a copy of ] for his family.<ref>{{Harvnb|van Wyhe|2008b|pp=18–21}}</ref> He had some expertise in geology, beetle collecting and dissecting marine invertebrates, but in all other areas, was a novice and ably collected specimens for expert appraisal.<ref name="Randal Keynes-2006">{{cite web|url=http://darwin-online.org.uk/EditorialIntroductions/Chancellor_Keynes_Galapagos.html|title=Darwin's field notes on the Galapagos: 'A little world within itself'|author=Gordon Chancellor|author2=Randal Keynes|date=October 2006|publisher=]|access-date=16 September 2009|url-status=dead|archive-url=https://web.archive.org/web/20090901082402/http://darwin-online.org.uk/EditorialIntroductions/Chancellor_Keynes_Galapagos.html|archive-date=1 September 2009|author2-link=Randal Keynes}}</ref> Despite suffering badly from seasickness, Darwin wrote copious notes while on board the ship. Most of his zoology notes are about marine invertebrates, starting with ] collected during a calm spell.<ref name="Keynes 2000" /><ref name="Keynes 2001-4">{{Harvnb|Keynes|2001|pp=}}</ref>
	Darwin spent the summer of 1825 as an apprentice doctor, helping his father treat the poor of Shropshire, before going to the ] with his brother Erasmus in October 1825. He found lectures dull and ] distressing, so neglected his medical studies. He learned ] from ], a freed black slave who had accompanied ] in the ]n ], and often sat with this "very pleasant and intelligent man".<ref name=eddy>{{harvnb|Darwin|1958|pp=}}</ref>
			] in Argentina, with fossils; caricature by ], the initial ship's artist]]
	In Darwin's second year he joined the ], a student ] group whose debates strayed into ] ]. He assisted ]'s investigations of the anatomy and life cycle of ] in the ], and in March 1827 presented at the Plinian his own discovery that black spores found in ] shells were the eggs of a skate ]. One day, Grant praised ] ]. Darwin was astonished, but had recently read the similar ideas of his grandfather Erasmus and remained indifferent.<ref>{{Harvnb|Browne|1995|pp=72–88}}</ref> Darwin was rather bored by ]'s natural history course which covered ] including the debate between ] and ]. He learned ] of plants, and assisted with work on the collections of the ], one of the largest museums in Europe at the time.<ref>{{Harvnb|Desmond|Moore|1991|pp=42–43}}</ref>
			On their first stop ashore at ] in Cape Verde, Darwin found that a white band high in the ] cliffs included seashells. FitzRoy had given him the first volume of ]'s '']'', which set out ] concepts of land slowly rising or falling over immense periods,{{Ref label|B|II|none}} and Darwin saw things Lyell's way, theorising and thinking of writing a book on geology.<ref>{{Harvnb|Browne|1995|pp=183–190}}</ref> When they reached Brazil, Darwin was delighted by the ],<ref>{{harvnb|Keynes|2001|pp=}}</ref> but detested the sight of ], and disputed this issue with FitzRoy.<ref>{{harvnb|Darwin|1958|pp=}}</ref>
	This neglect of medical studies annoyed his father, who shrewdly sent him to ], for a ] degree as the first step towards becoming an Anglican ].<ref>{{Harvnb|Browne|1995|pp=47–48}}</ref> Darwin began there in January 1828, but preferred ] and ] to studying. His cousin ] introduced him to the popular craze for ] collecting which he pursued zealously, getting some of his finds published in ] ''Illustrations of British entomology''. He became a close friend and follower of botany professor ] and met other leading naturalists who saw scientific work as religious ], becoming known to these ] as "the man who walks with Henslow". When exams drew near, Darwin focused on his studies and was delighted by the language and logic of ]'s ''Evidences of Christianity''.<ref name=dar57>{{Harvnb|Darwin|1958|pp=}}</ref> In his final examination in January 1831 Darwin did well, coming tenth out of a pass list of 178.<ref>{{Harvnb|Browne|1995|p=97}}</ref>
			The survey continued to the south in ]. They stopped at ], and in cliffs near ] Darwin made a major find of fossil bones of huge ] beside modern seashells, indicating recent ] with no signs of change in climate or catastrophe. He found bony plates like a giant version of the armour on local ]s. From a jaw and tooth he identified the gigantic '']'', then from ] description thought the armour was from this animal. The finds were shipped to England, and scientists found the fossils of great interest.<ref>{{Harvnb|Browne|1995|pp=223–225}}<br />{{Harvnb|Darwin|1835|p=}}<br />{{cite web|url=https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-213.xml|title=Letter no. 213, Henslow, J. S. to Darwin, C. R., 31 August 1833|website=Darwin Correspondence Project|access-date=29 December 2021|archive-date=29 December 2021|archive-url=https://web.archive.org/web/20211229182046/https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-213.xml|url-status=live}}</ref><ref name="Keynes 2001">{{harvnb|Keynes|2001|pp=}}</ref> In Patagonia, Darwin came to wrongly believe the territory was devoid of reptiles.<ref name="Jaksic-2022">{{Cite journal |title=Historical account and current ecological knowledge of the southernmost lizard in the world, Liolaemus magellanicus (Squamata: Liolaemidae) |journal=] |last=Jaksic |first=Fabian M. |issue=7 |volume=95 |doi=10.1186/s40693-022-00112-y |year=2022|s2cid=252717680 |doi-access=free |bibcode=2022RvCHN..95....7J }}</ref>
	Darwin had to stay at Cambridge until June. He studied Paley's '']'' which made an ], explaining ] as God acting through laws of nature.<ref name=syd5-7>{{Harvnb|von Sydow|2005|pp=5–7}}</ref> He read ]'s new book which described the highest aim of ] as understanding such laws through ] based on observation, and ]'s ''Personal Narrative'' of scientific travels. Inspired with "a burning zeal" to contribute, Darwin planned to visit ] with some classmates after graduation to study natural history in the ]. In preparation, he joined ]'s geology course, then went with him in the summer to map strata in ].<ref name=db>{{Harvnb|Darwin|1958|pp=}}<br />{{Harvnb|Browne|1995|pp=128–129, 133–141}}</ref> After a fortnight with student friends at ], he returned home to find a letter from Henslow proposing Darwin as a suitable (if unfinished) gentleman naturalist for a self-funded place with captain ], more as a companion than a mere collector, on {{HMS|Beagle}} which was to leave in four weeks on an expedition to chart the coastline of South America.<ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-105.html|title=Darwin Correspondence Project - Letter 105 — Henslow, J. S. to Darwin, C. R., 24 Aug 1831|accessdate=2008-12-29}}</ref> His father objected to the planned two-year voyage, regarding it as a waste of time, but was persuaded by his brother-in-law, ], to agree to his son's participation.<ref>{{Harvnb|Desmond|Moore|1991|pp= 94–97}}</ref>
			On rides with ]s into the interior to explore geology and collect more fossils, Darwin gained social, political and ] insights into both native and colonial people at a time of revolution, and learnt that two types of ] had separate but overlapping territories.<ref>{{Harvnb|Desmond|Moore|1991|pp=189–192, 198}}</ref><ref>{{Harvnb|Eldredge|2006}}</ref> Further south, he saw stepped plains of shingle and seashells as ]es at a series of elevations. He read Lyell's second volume and accepted its view of "centres of creation" of species, but his discoveries and theorising challenged Lyell's ideas of smooth continuity and of extinction of species.<ref>{{Harvnb|Desmond|Moore|1991|pp=131, 159}}<br />{{harvnb|Herbert|1991|pp=}}</ref><ref name="Darwin Online-2">{{cite web|url=http://darwin-online.org.uk/EditorialIntroductions/Chancellor_fieldNotebooks1.8.html|title=Darwin Online: 'Hurrah Chiloe': an introduction to the Port Desire Notebook|access-date=24 October 2008|url-status=dead|archive-url=https://web.archive.org/web/20081204201413/http://www.darwin-online.org.uk./EditorialIntroductions/Chancellor_fieldNotebooks1.8.html|archive-date=4 December 2008}}</ref>
	===Journey of the ''Beagle''===
	{{details|Second voyage of HMS Beagle}}
	]
	Beginning on the 27th of December, 1831, the voyage lasted almost five years and, as FitzRoy had intended, Darwin spent most of that time on land investigating geology and making natural history collections, while the ''Beagle'' ] coasts.<ref name=JvW/><ref name=kix>{{harvnb|Keynes|2000|pp=}}</ref> He kept careful notes of his observations and theoretical speculations, and at intervals during the voyage his specimens were sent to Cambridge together with letters including a copy of ] for his family.<ref>{{Harvnb|van Wyhe|2008b|pp=18–21}}</ref> He had some expertise in geology, beetle collecting and dissecting ], but in all other areas was a novice and ably collected specimens for expert appraisal.<ref name=fnGal>{{cite web|url=http://darwin-online.org.uk/EditorialIntroductions/Chancellor_Keynes_Galapagos.html|title=Darwin's field notes on the Galapagos: 'A little world within itself'|author=Gordon Chancellor|coauthors=]|month=October| year=2006|publisher=]|accessdate=2009-09-16}}</ref> Despite repeatedly suffering badly from seasickness while at sea, most of his zoology notes are about marine invertebrates, starting with ] collected in a calm spell.<ref name=kix/><ref name=plankton>{{Harvnb|Keynes|2001|pp=}}</ref>
			Three ] on board, who had been seized during the ] then given Christian education in England, were returning with a missionary. Darwin found them friendly and civilised, yet at ] he met "miserable, degraded savages", as different as wild from domesticated animals.<ref>{{Harvnb|Darwin|1845|pp=}}</ref> He remained convinced that, despite this diversity, all humans were interrelated with ] and potential for improvement towards civilisation. Unlike his scientist friends, he now thought there was no unbridgeable gap between humans and animals.<ref>{{Harvnb|Browne|1995|pp=243–244, 248–250, 382–383}}</ref> A year on, the mission had been abandoned. The Fuegian they had named ] lived like the other natives, had a wife, and had no wish to return to England.<ref>{{harvnb|Keynes|2001|pp=}}</ref>
	On their first stop ashore at ], Darwin found that a white band high in the ] cliffs included seashells. FitzRoy had given him the first volume of ]'s ''Principles of Geology'' which set out ] concepts of land slowly rising or falling over immense periods,{{Ref_label|B|II|none}} and Darwin saw things Lyell's way, theorising and thinking of writing a book on geology.<ref>{{Harvnb|Browne|1995|pp=183–190}}</ref>
	In ], Darwin was delighted by the ],<ref>{{harvnb|Keynes|2001|pp=}}</ref> but detested the sight of ].<ref>{{harvnb|Darwin|1958|pp=}}</ref>
			] surveyed the coasts of South America, Darwin theorised about geology and the extinction of giant mammals; watercolour by the ship's artist ], who replaced Augustus Earle, in ]]]
	At ] in ] he made a major find of fossils of huge extinct ]s in cliffs beside modern seashells, indicating recent ] with no signs of change in climate or catastrophe. He identified the little known '']'', with bony armour which at first seemed to him like a giant version of the armour on local ]s. The finds brought great interest when they reached England.<ref>{{Harvnb|Browne|1995|pp= 223–235}}<br />{{Harvnb|Darwin|1835|p=}}<br />{{Harvnb|Desmond|Moore|1991|p= 210}}</ref> On rides with ]s into the interior to explore geology and collect more fossils he gained social, political and ] insights into both native and colonial people at a time of revolution, and learnt that two types of ] had separate but overlapping territories.<ref>{{Harvnb|Desmond|Moore|1991|pp= 189–192, 198}}</ref><ref>{{Harvnb|Eldredge|2006}}</ref> Further south he saw stepped plains of shingle and seashells as ]es showing a series of elevations. He read Lyell's second volume and accepted its view of "centres of creation" of species, but his discoveries and theorising challenged Lyell's ideas of smooth continuity and of extinction of species.<ref>{{Harvnb|Desmond|Moore|1991|pp= 131, 159}}<br />{{harvnb|Herbert|1991|pp=}}</ref><ref name=HurrahChiloe>{{cite web|url= http://darwin-online.org.uk/EditorialIntroductions/Chancellor_fieldNotebooks1.8.html|title=Darwin Online: 'Hurrah Chiloe': an introduction to the Port Desire Notebook|accessdate=2008-10-24}}</ref>
			Darwin experienced ] in 1835 and saw signs that the land had just been raised, including ]-beds stranded above high tide. High in the ] he saw seashells and several fossil trees that had grown on a sand beach. He theorised that as the land rose, ] sank, and ]s round them grew to form ]s.<ref>{{Harvnb|Desmond|Moore|1991|pp=160–168, 182}}<br />{{cite web | title=Letter no. 275 – Charles Darwin to Susan Elizabeth Darwin – 23 April 1835 | website=Darwin Correspondence Project | url=https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-275.xml | access-date=6 December 2021 | archive-date=6 December 2021 | archive-url=https://web.archive.org/web/20211206101229/https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-275.xml | url-status=live }}</ref><ref name="Darwin 1958">{{Harvnb|Darwin|1958|loc=}}</ref>
	] surveyed the coasts of ], Darwin theorised about geology and extinction of giant mammals.]]
	Three Fuegians on board, who had been seized during the ] and had spent a year in England, were taken back to ] as missionaries. Darwin found them friendly and civilised, yet their relatives seemed "miserable, degraded savages", as different as wild from domesticated animals.<ref>{{Harvnb|Darwin|1845|pp= }}</ref> To Darwin the difference showed cultural advances, not racial inferiority. Unlike his scientist friends, he now thought there was no unbridgeable gap between humans and animals.<ref>{{Harvnb|Browne|1995|pp= 244–250}}</ref> A year on, the mission had been abandoned. The Fuegian they'd named ] lived like the other natives, had a wife, and had no wish to return to England.<ref>{{harvnb|Keynes|2001|pp=}}</ref>
			On the geologically new ], Darwin looked for evidence attaching wildlife to an older "centre of creation", and found ]s allied to those in Chile but differing from island to island. He heard that slight variations in the shape of ] shells showed which island they came from, but failed to collect them, even after eating tortoises taken on board as food.<ref name="Keynes 2001-2">{{harvnb|Keynes|2001|pp=}}</ref><ref>{{harvnb|Sulloway|1982|p=19}}</ref> In Australia, the ] ] and the ] seemed so unusual that Darwin thought it was almost as though two distinct Creators had been at work.<ref name="Darwin Online">{{cite web|url=http://darwin-online.org.uk/EditorialIntroductions/Chancellor_fieldNotebooks1.3.html|title=Darwin Online: Coccatoos & Crows: An introduction to the Sydney Notebook|access-date=2 January 2009|url-status=live|archive-url=https://web.archive.org/web/20090114015611/http://darwin-online.org.uk/EditorialIntroductions/Chancellor_fieldNotebooks1.3.html|archive-date=14 January 2009}}</ref> He found the ] "good-humoured & pleasant", their numbers depleted by European settlement.<ref>{{harvnb|Keynes|2001|pp=.}}</ref>
	Darwin experienced an earthquake in ] and saw signs that the land had just been raised, including ]-beds stranded above high tide. High in the ] he saw seashells, and several fossil trees that had grown on a sand beach. He theorised that as the land rose, ] sank, and ]s round them grew to form ]s.<ref>{{Harvnb|Desmond|Moore|1991|pp= 160–168, 182}}<br />{{Harvnb|Darwin|1887|p= }}</ref><ref name=atolls>{{Harvnb|Darwin|1958|loc=}}</ref>
			FitzRoy investigated how the atolls of the ] had formed, and the survey supported Darwin's theorising.<ref name="Darwin 1958" /> FitzRoy began writing the official ''Narrative'' of the ''Beagle'' voyages, and after reading Darwin's diary, he proposed incorporating it into the account.<ref name="Darwin Correspondence Project-5">{{cite web | title=Letter no. 301, Charles Darwin to Caroline Darwin, 29 April 1836, Port Lewis, Mauritius. | website=Darwin Correspondence Project | url=https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-301.xml | access-date=12 February 2022 | archive-date=15 February 2022 | archive-url=https://web.archive.org/web/20220215113535/https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-301.xml | url-status=live }}</ref> Darwin's ''Journal'' was eventually rewritten as a separate third volume, on geology and natural history.<ref>{{Harvnb|Browne|1995|p=336}}</ref><ref name="Darwin 1839">{{Harvnb|Darwin|1839|p=}}</ref>
	On the geologically new ] Darwin looked for evidence attaching wildlife to an older "centre of creation", and found ]s allied to those in Chile but differing from island to island. He heard that slight variations in the shape of ] shells showed which island they came from, but failed to collect them, even after eating tortoises taken on board as food.<ref name=k356>{{harvnb|Keynes|2001|pp=}}</ref><ref>{{harvnb|Sulloway|1982|p=19}}</ref> In Australia, the ] ] and the ] seemed so unusual that Darwin thought it was almost as though two distinct Creators had been at work.<ref name=Crows>{{cite web|url=http://darwin-online.org.uk/EditorialIntroductions/Chancellor_fieldNotebooks1.3.html|title=Darwin Online: Coccatoos & Crows: An introduction to the Sydney Notebook|accessdate=2009-01-02}}</ref> He found the ] "good-humoured & pleasant", and noted their depletion by European settlement.<ref>{{harvnb|Keynes|2001|pp=.}}</ref>
			In ], South Africa, Darwin and FitzRoy met John Herschel, who had recently written to Lyell praising his ] as opening bold speculation on "that mystery of mysteries, the replacement of extinct species by others" as "a natural in contradistinction to a miraculous process".<ref name="van Wyhe 2007">{{harvnb|van Wyhe|2007|p=}}</ref>
	The ''Beagle'' investigated how the atolls of the ] had formed, and the survey supported Darwin's theorising.<ref name=atolls/> FitzRoy began writing the official ''Narrative'' of the ''Beagle'' voyages, and after reading Darwin's diary he proposed incorporating it into the account.<ref name=Letter301>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-301.html|title=Darwin Correspondence Project - Letter 301 — Darwin, C.R. to Darwin, C.S., 29 Apr 1836}}</ref> Darwin's '']'' was eventually rewritten as a separate third volume, on natural history.<ref>{{Harvnb|Browne|1995|p= 336}}</ref>
			When organising his notes as the ship sailed home, Darwin wrote that, if his growing suspicions about the mockingbirds, the tortoises and the ] were correct, "such facts undermine the stability of Species", then cautiously added "would" before "undermine".<ref name="Keynes 2000-2">{{Harvnb|Keynes|2000|pp=}}<br />{{Harvnb|Eldredge|2006}}</ref> He later wrote that such facts "seemed to me to throw some light on the origin of species".<ref>{{Harvnb|Darwin|1859|loc=}}</ref>
			Without telling Darwin, ] had been read to scientific societies, printed as a pamphlet for private distribution among members of the ], and reported in magazines,{{sfn|Darwin|1835|p=1}} including ].<ref name="Darwin Correspondence Project-3">{{cite web | title=Letter no. 291, Caroline Darwin to Charles Darwin, 29 December , | website=Darwin Correspondence Project | url=https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-291.xml | access-date=19 January 2022 | archive-date=11 February 2022 | archive-url=https://web.archive.org/web/20220211205926/https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-291.xml | url-status=live }}</ref> Darwin first heard of this at Cape Town,<ref name="Darwin Correspondence Project-4">{{cite web | title=Letter no. 302, Charles Darwin to Catherine Darwin, 3 June 1836, Cape of Good Hope | website=Darwin Correspondence Project | url=https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-302.xml | access-date=19 January 2022 | archive-date=27 January 2022 | archive-url=https://web.archive.org/web/20220127212812/https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-302.xml | url-status=live }}</ref> and at ] read of Sedgwick's prediction that Darwin "will have a great name among the Naturalists of Europe".<ref name="Darwin Correspondence Project-2">{{cite web | title=Letter no. 288, Susan Darwin to Charles Darwin, 22 November 1835, Shrewsbury | website=Darwin Correspondence Project | url=https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-288.xml | access-date=19 January 2022 | archive-date=13 February 2022 | archive-url=https://web.archive.org/web/20220213100832/https://www.darwinproject.ac.uk/letter/?docId=letters/DCP-LETT-288.xml | url-status=live }}</ref><ref>{{harvnb|Darwin|1958|pp=.}}</ref>
	In ] Darwin and FitzRoy met ], who had recently written to Lyell praising his ] as opening bold speculation on "that mystery of mysteries, the replacement of extinct species by others" as "a natural in contradistinction to a miraculous process".<ref name=Rascals>{{harvnb|van Wyhe|2007|p=}}</ref>
	When organising his notes as the ship sailed home, Darwin wrote that if his growing suspicions about the mockingbirds, the tortoises and the ] were correct, "such facts undermine the stability of Species", then cautiously added "would" before "undermine".<ref name=xix>{{Harvnb|Keynes|2000|pp=}}<br />{{Harvnb|Eldredge|2006}}</ref> He later wrote that such facts "seemed to me to throw some light on the origin of species".<ref>{{Harvnb|Darwin|1859|loc=}}</ref>
	===Inception of Darwin's evolutionary theory===		===Inception of Darwin's evolutionary theory===
	{{details|Inception of Darwin's theory}}		{{further|Inception of Darwin's theory}}
	]		].]]
	When the ''Beagle'' reached ], on 2 October 1836, Darwin was already a celebrity in scientific circles as in December 1835 ] had fostered his former pupil's reputation by giving selected naturalists a pamphlet of Darwin's geological letters.<ref>{{Harvnb|Darwin|1835|loc=}}</ref> Darwin visited his home in Shrewsbury and saw relatives, then hurried to ] to see Henslow, who advised on finding naturalists available to catalogue the collections and agreed to take on the botanical specimens. Darwin's father organised investments, enabling his son to be a self-funded ] scientist, and an excited Darwin went round the ] institutions being fêted and seeking experts to describe the collections. Zoologists had a huge backlog of work, and there was a danger of specimens just being left in storage.<ref>{{Harvnb|Desmond|Moore|1991|pp= 195–198}}</ref>		On 2 October 1836, ''Beagle'' anchored at ], Cornwall. Darwin promptly made the long coach journey to Shrewsbury to visit his home and see relatives. He then hurried to ] to see Henslow, who advised him on finding available naturalists to catalogue Darwin's animal collections and to take on the botanical specimens. Darwin's father organised investments, enabling his son to be a self-funded ], and an excited Darwin went around the London institutions being fêted and seeking experts to describe the collections. British zoologists at the time had a huge backlog of work, due to natural history collecting being encouraged throughout the British Empire, and there was a danger of specimens just being left in storage.<ref>{{Harvnb|Desmond|Moore|1991|pp=195–198}}</ref>
	] eagerly met Darwin for the first time on 29 October and soon introduced him to the up-and-coming anatomist ], who had the facilities of the ] to work on the fossil bones collected by Darwin. Owen's surprising results included gigantic extinct ]s, a near complete skeleton of the unknown '']'' and a ]-sized ]-like skull named '']'' resembling a giant ]. The armour fragments were from '']'', a huge armadillo-like creature as Darwin had initially thought.<ref>{{Harvnb|Owen|1840|pp=, , }}<br />{{Harvnb|Eldredge|2006}}</ref> These extinct creatures were related to living species in South America.<ref>{{Harvnb|Desmond|Moore|1991|pp= 201–205}}.<br />{{Harvnb|Browne|1995|pp=349–350}}</ref>		Charles Lyell eagerly met Darwin for the first time on 29 October and soon introduced him to the up-and-coming anatomist ], who had the facilities of the ] to work on the fossil bones collected by Darwin. Owen's surprising results included other gigantic extinct ]s as well as the ''Megatherium'' Darwin had identified, a near complete skeleton of the unknown '']'' and a ]-sized ]-like skull named '']'' resembling a giant ]. The armour fragments were actually from '']'', a huge armadillo-like creature, as Darwin had initially thought.<ref name="Keynes 2001" /><ref>{{Harvnb|Owen|1840|pp=, , }}<br />{{Harvnb|Eldredge|2006}}</ref> These extinct creatures were related to living species in South America.<ref>{{Harvnb|Desmond|Moore|1991|pp=201–205}}<br />{{Harvnb|Browne|1995|pp=349–350}}</ref>
			In mid-December, Darwin took lodgings in Cambridge to arrange expert classification of his collections, and prepare his own research for publication. Questions of how to combine his diary into the ''Narrative'' were resolved at the end of the month when FitzRoy accepted ] advice to make it a separate volume, and Darwin began work on his ''Journal and Remarks''.{{sfn|Browne|1995|pp=345–347}}{{sfn|Keynes|2001|pp=}}
	In mid-December Darwin took lodgings in Cambridge to organise work on his collections and rewrite his ''Journal''.<ref>{{Harvnb|Browne|1995|pp=345–347}}</ref> He wrote his first paper, showing that the South American landmass was slowly rising, and with Lyell's enthusiastic backing read it to the ] on 4 January 1837. On the same day, he presented his mammal and bird specimens to the ]. The ornithologist ] soon announced that the Galapagos birds that Darwin had thought a mixture of ], "]" and ]es, were, in fact, twelve ]. On 17 February Darwin was elected to the Council of the Geological Society, and Lyell's presidential address presented Owen's findings on Darwin's fossils, stressing geographical continuity of species as supporting his ] ideas.<ref>{{Harvnb|Desmond|Moore|1991|pp= 207–210}}<br />{{Harvnb|Sulloway|1982|pp=20–23}}</ref>
			Darwin's first paper showed that the South American landmass was slowly rising. With Lyell's enthusiastic backing, he read it to the ] on 4 January 1837. On the same day, he presented his mammal and bird specimens to the ]. The ornithologist ] soon announced that the Galápagos birds that Darwin had thought a mixture of ], "]" and ]es, were, in fact, twelve ]. On 17 February, Darwin was elected to the Council of the Geological Society, and Lyell's presidential address presented Owen's findings on Darwin's fossils, stressing geographical continuity of species as supporting his uniformitarian ideas.<ref>{{Harvnb|Desmond|Moore|1991|pp=207–210}}<br />{{Harvnb|Sulloway|1982|pp=20–23}}</ref>
	Early in March, Darwin moved to London to be near this work, joining Lyell's social circle of scientists and ]s such as ],<ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-346.html|title=Darwin Correspondence Project - Letter 346 — Darwin, C. R. to Darwin, C. S., 27 Feb 1837|accessdate=2008-12-19}} proposes a move on Friday 3 March 1837,<br />Darwin's Journal ({{harvnb|Darwin|2006|p=}}) backdated from August 1838 gives a date of 6 March 1837</ref> who described God as a programmer of laws. ]'s letter on the "mystery of mysteries" of new species was widely discussed, with explanations sought in ], not ] miracles. Darwin stayed with his ] brother ], part of this ] circle and close friend of writer ] who promoted ] underlying the controversial Whig ] to stop welfare from causing overpopulation and more poverty. As a ] she welcomed the ] implications of ], promoted by ] and younger surgeons influenced by ], but anathema to Anglicans defending social order.<ref name=Rascals/><ref>{{Harvnb|Desmond|Moore|1991|pp=201, 212–221}}</ref>
			].]]
	In their first meeting to discuss his detailed findings, Gould told Darwin that the Galápagos ]s from different islands were separate species, not just varieties, and the finch group included the "]s". Darwin had not labelled the finches by island, but from the notes of others on the ''Beagle'', including FitzRoy, he allocated species to islands.<ref>{{Harvnb|Sulloway|1982|pp=20–23}}</ref> The two ] were also distinct species, and on 14 March Darwin announced how their distribution changed going southwards.<ref>{{Harvnb|Browne|1995|p=360}}<br />{{cite web|url=http://darwin-online.org.uk/content/frameset?itemID=F1643&viewtype=text&pageseq=1|title=Darwin, C. R. (Read 14 March 1837) Notes on Rhea americana and Rhea darwinii, ''Proceedings of the Zoological Society of London''|accessdate=2008-12-17}}</ref>
			Early in March, Darwin moved to London to be near this work, joining Lyell's social circle of scientists and experts such as ],<ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-346.html|title=Darwin Correspondence Project&nbsp;– Letter 346&nbsp;– Darwin, C. R. to Darwin, C. S., 27 Feb 1837|access-date=19 December 2008|url-status=live|archive-url=https://web.archive.org/web/20090629192201/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-346.html|archive-date=29 June 2009}} proposes a move on Friday 3 March 1837,<br />Darwin's Journal ({{harvnb|Darwin|2006|pp=}}) backdated from August 1838 gives a date of 6 March 1837</ref> who described God as a programmer of laws. Darwin stayed with his ] brother Erasmus, part of this ] circle and a close friend of the writer ], who promoted the ] that underpinned the controversial Whig ] to stop welfare from causing overpopulation and more poverty. As a Unitarian, she welcomed the ] implications of ], promoted by Grant and younger surgeons influenced by ]. Transmutation was anathema to Anglicans defending social order,<ref>{{Harvnb|Desmond|Moore|1991|pp=201, 212–221}}</ref> but reputable scientists openly discussed the subject, and there was wide interest in John Herschel's letter praising Lyell's approach as a way to find a ] of the origin of new species.<ref name="van Wyhe 2007" />
			Gould met Darwin and told him that the Galápagos mockingbirds from different islands were separate species, not just varieties, and what Darwin had thought was a "]" was ]. Darwin had not labelled the finches by island, but from the notes of others on the ship, including FitzRoy, he allocated species to islands.<ref>{{Harvnb|Sulloway|1982|pp=9, 20–23}}</ref> The two rheas were distinct species, and on 14 March Darwin announced how their distribution changed going southwards.<ref>{{Harvnb|Browne|1995|p=360}}<br />{{cite web|url=http://darwin-online.org.uk/content/frameset?itemID=F1643&viewtype=text&pageseq=1|title=Darwin, C. R. (Read 14 March 1837) Notes on Rhea americana and Rhea darwinii, ''Proceedings of the Zoological Society of London''|access-date=17 December 2008|url-status=live|archive-url=https://web.archive.org/web/20090210085710/http://darwin-online.org.uk/content/frameset?itemID=F1643&viewtype=text&pageseq=1|archive-date=10 February 2009}}</ref>
	].]]
	By mid-March, Darwin was speculating in his ''Red Notebook'' on the possibility that "one species does change into another" to explain the geographical distribution of living species such as the rheas, and extinct ones such as the strange '']'', resembling a giant ]. His thoughts on lifespan, ] and ] developed in his "B" notebook around mid-July on to variation in offspring "to adapt & alter the race to ''changing'' world" explaining the ]s, mockingbirds and rheas. He sketched branching descent, then a ] branching of a single ], in which "It is absurd to talk of one animal being higher than another", discarding ] independent ] progressing to higher forms.<ref>{{harvnb|Herbert|1980|pp=}}<br />{{Harvnb|van Wyhe|2008b|p=44}}<br />{{harvnb|Darwin|1837|pp=}}<br />{{Harvnb|Desmond|Moore|1991|pp=229–232}}</ref>
			By mid-March 1837, barely six months after his return to England, Darwin was speculating in his ''Red Notebook'' on the possibility that "one species does change into another" to explain the geographical distribution of living species such as the rheas, and extinct ones such as the strange extinct mammal '']'', which resembled a giant ], a llama relative. Around mid-July, he recorded in his "B" notebook his thoughts on lifespan and variation across generations{{snd}}explaining the variations he had observed in ]s, mockingbirds, and rheas. He sketched branching descent, and then a ] branching of a single ], in which "It is absurd to talk of one animal being higher than another", thereby discarding Lamarck's idea of independent ] progressing to higher forms.<ref>{{harvnb|Herbert|1980|pp=}}<br />{{Harvnb|van Wyhe|2008b|p=44}}<br />{{harvnb|Darwin|1837|pp=}}<br />{{Harvnb|Desmond|Moore|1991|pp=229–232}}</ref>
	===Overwork, illness, and marriage===		===Overwork, illness, and marriage===
			{{further|Health of Charles Darwin}}
	While developing this intensive study of ], Darwin became mired in more work. Still rewriting his ''Journal'', he took on editing and publishing the expert reports on his collections, and with Henslow's help obtained a Treasury grant of ]1,000 to sponsor this multi-volume '']'', a sum equivalent to about £{{formatnum:{{inflation|UK|1000|1837|r=-3}}}} in present day terms.{{inflation-fn|UK}} He stretched the funding to include his planned books on geology, and agreed unrealistic dates with the publisher.<ref>{{Harvnb|Browne|1995|pp=367–369}}</ref> As the ] began, Darwin pressed on with writing his ''Journal'', and in August 1837 began correcting ].<ref>{{harvnb|Keynes|2001|p=}}</ref>
			While developing this intensive study of transmutation, Darwin became mired in more work. Still rewriting his ''Journal'', he took on editing and publishing the expert reports on his collections, and with Henslow's help obtained a Treasury grant of £1,000 to sponsor this multi-volume '']'', a sum equivalent to about £115,000 in 2021.<ref>{{cite web |title=£1,000 in 1832 → 2021 {{!}} UK Inflation Calculator |url=https://www.in2013dollars.com/uk/inflation/1832 |access-date=8 August 2021 |website=www.in2013dollars.com |archive-date=15 August 2021 |archive-url=https://web.archive.org/web/20210815234035/https://www.in2013dollars.com/uk/inflation/1832 |url-status=live }}</ref> He stretched the funding to include his planned books on geology, and agreed to unrealistic dates with the publisher.<ref>{{Harvnb|Browne|1995|pp=367–369}}</ref> As the ] began, Darwin pressed on with writing his ''Journal'', and in August 1837 began correcting ].<ref name="Keynes 2001-3">{{harvnb|Keynes|2001|p=}}</ref>
	Darwin's health suffered from the pressure. On 20 September he had "an uncomfortable palpitation of the heart", so his doctors urged him to "knock off all work" and live in the country for a few weeks. After visiting Shrewsbury he joined his Wedgwood relatives at ], ], but found them too eager for tales of his travels to give him much rest. His charming, intelligent, and cultured cousin ], nine months older than Darwin, was nursing his invalid aunt. His uncle ] pointed out an area of ground where cinders had disappeared under ] and suggested that this might have been the work of ]s, inspiring "a new & important theory" on their role in ] which Darwin presented at the Geological Society on 1 November.<ref>{{Harvnb|Desmond|Moore|1991|pp= 233–234}}<br />{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-404.html|title=Darwin Correspondence Project - Letter 404 — Buckland, William to Geological Society of London, 9 Mar 1838|accessdate=2008-12-23}}</ref>
			As Darwin worked under pressure, his health suffered. On 20 September, he had "an uncomfortable ] of the heart", so his doctors urged him to "knock off all work" and live in the country for a few weeks. After visiting Shrewsbury, he joined his Wedgwood relatives at ], Staffordshire, but found them too eager for tales of his travels to give him much rest. His charming, intelligent, and cultured cousin ], nine months older than Darwin, was nursing his invalid aunt. His uncle Josiah pointed out an area of ground where cinders had disappeared under ] and suggested that this might have been the work of ]s, inspiring "a new & important theory" on their role in ], which Darwin presented at the Geological Society on 1 November 1837.<ref>{{Harvnb|Desmond|Moore|1991|pp=233–234}}<br />{{cite web |url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-404.html |title=Darwin Correspondence Project&nbsp;– Letter 404&nbsp;– Buckland, William to Geological Society of London, 9 Mar 1838 |access-date=23 December 2008 |url-status=live|archive-url=https://web.archive.org/web/20090629192234/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-404.html |archive-date=29 June 2009}}</ref> His ''Journal'' was printed and ready for publication by the end of February 1838, as was the first volume of the ''Narrative'', but FitzRoy was still working hard to finish his own volume.<ref name="Keynes 2001-3" />
	] pushed Darwin to take on the duties of Secretary of the Geological Society. After initially declining the work, he accepted the post in March 1838.<ref>{{Harvnb|Desmond|Moore|1991|pp= 233–236}}.</ref> Despite the grind of writing and editing the ''Beagle'' reports, Darwin made remarkable progress on transmutation, taking every opportunity to question expert naturalists and, unconventionally, people with practical experience such as farmers and ].<ref name=JvW/><ref>{{Harvnb|Desmond|Moore|1991|pp= 241–244, 426}}</ref> Over time his research drew on information from his relatives and children, the family butler, neighbours, colonists and former shipmates.<ref>{{Harvnb|Browne|1995|p=xii}}</ref> He included mankind in his speculations from the outset, and on seeing an ] in the zoo on 28 March 1838 noted its child-like behaviour.<ref>{{Harvnb|Desmond|Moore|1991|pp= 241–244}}</ref>
			] pushed Darwin to take on the duties of Secretary of the Geological Society. After initially declining the work, he accepted the post in March 1838.<ref>{{Harvnb|Desmond|Moore|1991|pp=233–236}}.</ref> Despite the grind of writing and editing the ''Beagle'' reports, Darwin made remarkable progress on transmutation, taking every opportunity to question expert naturalists and, unconventionally, people with practical experience in ] such as farmers and ].<ref name="van Wyhe 2008" /><ref>{{Harvnb|Desmond|Moore|1991|pp=241–244, 426}}</ref> Over time, his research drew on information from his relatives and children, the family butler, neighbours, colonists and former shipmates.<ref>{{Harvnb|Browne|1995|p=xii}}</ref> He included mankind in his speculations from the outset, and on seeing an ] in the zoo on 28 March 1838 noted its childlike behaviour.<ref>{{Harvnb|Desmond|Moore|1991|pp=241–244}}</ref>
	The strain took a toll, and by June he was being laid up for days on end with stomach problems, headaches and heart symptoms. For the rest of his life, he was repeatedly incapacitated with episodes of stomach pains, vomiting, severe ]s, palpitations, trembling and other symptoms, particularly during times of stress such as attending meetings or making social visits. The cause of ] remained unknown, and attempts at treatment had little success.<ref>{{Harvnb|Desmond|Moore|1991|pp= 252, 476, 531}}<br />{{harvnb|Darwin|1958|p=}}</ref>
	On 23 June he took a break and went "geologising" in Scotland. He visited ] in glorious weather to see the parallel "roads" cut into the hillsides at three heights. He later published his view that these were marine ]es, but then had to accept that they were shorelines of a ].<ref>{{Harvnb|Desmond|Moore|1991|p= 254}}<br />{{Harvnb|Browne|1995|pp=377–378}}<br />{{Harvnb|Darwin|1958|p=}}</ref>		The strain took a toll, and by June he was being laid up for days on end with stomach problems, headaches and heart symptoms. For the rest of his life, he was repeatedly incapacitated with episodes of stomach pains, vomiting, severe ]s, palpitations, trembling and other symptoms, particularly during times of stress, such as attending meetings or making social visits. The cause of ] remained unknown, and attempts at treatment had only ephemeral success.<ref>{{Harvnb|Desmond|Moore|1991|pp=252, 476, 531}}<br />{{harvnb|Darwin|1958|p=}}</ref>
			On 23 June, he took a break and went "geologising" in Scotland. He visited ] in glorious weather to see the parallel "roads" cut into the hillsides at three heights. He later published his view that these were marine-raised beaches, but then had to accept that they were shorelines of a ].<ref>{{Harvnb|Desmond|Moore|1991|p=254}}<br />{{Harvnb|Browne|1995|pp=377–378}}<br />{{Harvnb|Darwin|1958|p=}}</ref>
	Fully recuperated, he returned to Shrewsbury in July. Used to jotting down daily notes on animal breeding, he scrawled rambling thoughts about career and prospects on two scraps of paper, one with columns headed ''"Marry"'' and ''"Not Marry"''. Advantages included "constant companion and a friend in old age&nbsp;... better than a dog anyhow", against points such as "less money for books" and "terrible loss of time."<ref>{{Harvnb|Darwin|1958|pp=}}</ref> Having decided in favour, he discussed it with his father, then went to visit Emma on 29 July. He did not get around to proposing, but against his father's advice he mentioned his ideas on transmutation.<ref>{{Harvnb|Desmond|Moore|1991|pp=256–259}}</ref>
			].]]
	Continuing his research in London, Darwin's wide reading now included the sixth edition of ] '']''
			Fully recuperated, he returned to Shrewsbury in July 1838. Used to jotting down daily notes on animal breeding, he scrawled rambling thoughts about marriage, career and prospects on two scraps of paper, one with columns headed ''"Marry"'' and ''"Not Marry"''. Advantages under "Marry" included "constant companion and a friend in old age&nbsp;... better than a dog anyhow", against points such as "less money for books" and "terrible loss of time".<ref>{{Harvnb|Darwin|1958|pp=}}</ref> Having decided in favour of marriage, he discussed it with his father, then went to visit his cousin Emma on 29 July. At this time he did not get around to proposing, but against his father's advice, he mentioned his ideas on transmutation.<ref>{{Harvnb|Desmond|Moore|1991|pp=256–259}}</ref>
	{{Quotation|In October 1838, that is, fifteen months after I had begun my systematic enquiry, I happened to read for amusement Malthus on Population, and being well prepared to appreciate the struggle for existence which everywhere goes on from long-continued observation of the habits of animals and plants, it at once struck me that under these circumstances favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species. Here, then, I had at last got a theory by which to work...<ref>{{harvnb|Darwin|1958|p=}}</ref>}}
			He married Emma on 29 January 1839 and they were the parents of ten children, seven of whom survived to adulthood.
			===Malthus and natural selection===
	Malthus asserted that unless human population is kept in check, it increases in a ] and soon exceeds food supply in what is known as a ].<ref name="JvW"/> Darwin was well prepared to see at once that this also applied to ] "warring of the species" of plants and the struggle for existence among wildlife, explaining how numbers of a species kept roughly stable. As species always breed beyond available resources, favourable variations would make organisms better at surviving and passing the variations on to their offspring, while unfavourable variations would be lost. This would result in the formation of new species.<ref name=JvW/><ref>{{Harvnb|Desmond|Moore|1991|pp= 264–265}}<br />{{Harvnb|Browne|1995|pp= 385–388}}<br />{{Harvnb|Darwin|1842|p=}}</ref> On 28 September 1838 he noted this insight, describing it as a kind of wedging, forcing adapted structures into gaps in the economy of nature as weaker structures were thrust out.<ref name=JvW/> By mid December he saw a similarity between farmers picking the best breeding stock and a Malthusian Nature selecting from chance variants so that "every part of newly acquired structure is fully practical and perfected",<ref>{{cite web |url=http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=CUL-DAR124.-&pageseq=63|title=Darwin transmutation notebook E p. 75|accessdate=2009-03-17 }}</ref> thinking this comparison "a beautiful part of my theory".<ref>{{cite web |url=http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=CUL-DAR124.-&pageseq=61|title=Darwin transmutation notebook E p. 71|accessdate=2009-03-17 }}</ref>
			Continuing his research in London, Darwin's wide reading now included the sixth edition of ]'s '']''. On 28 September 1838, he noted its assertion that human "population, when unchecked, goes on doubling itself every twenty-five years, or increases in a geometrical ratio", a ] so that population soon exceeds food supply in what is known as a ]. Darwin was well-prepared to compare this to ]'s "warring of the species" of plants and the struggle for existence among wildlife, explaining how numbers of a species kept roughly stable. As species always breed beyond available resources, favourable variations would make organisms better at surviving and passing the variations on to their offspring, while unfavourable variations would be lost. He wrote that the "final cause of all this wedging, must be to sort out proper structure, & adapt it to changes", so that "One may say there is a force like a hundred thousand wedges trying force into every kind of adapted structure into the gaps of in the economy of nature, or rather forming gaps by thrusting out weaker ones."<ref name="van Wyhe 2008" /><ref name="Darwin transmutation notebook D pp">{{cite web | title=Darwin transmutation notebook D pp. 134e–135e | url=http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=CUL-DAR123.-&pageseq=112 | access-date=4 June 2012 | url-status=live | archive-url=https://web.archive.org/web/20120718105154/http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=CUL-DAR123.-&pageseq=112 | archive-date=18 July 2012 | df=dmy-all }}</ref> This would result in the formation of new species.<ref name="van Wyhe 2008" /><ref>{{Harvnb|Desmond|Moore|1991|pp=264–265}}<br />{{Harvnb|Browne|1995|pp=385–388}}<br />{{Harvnb|Darwin|1842|p=}}</ref> As he later wrote in his '']'':
	].]]
	On 11 November, he returned to Maer and proposed to Emma, once more telling her his ideas. She accepted, then in exchanges of loving letters she showed how she valued his openness in sharing their differences, also expressing her strong ] beliefs and concerns that his honest doubts might separate them in the afterlife.<ref name=Belief>{{cite web|url=http://www.darwinproject.ac.uk/content/view/130/125/|title=Darwin Correspondence Project - Belief: historical essay|accessdate=2008-11-25}}</ref> While he was house-hunting in London, bouts of illness continued and Emma wrote urging him to get some rest, almost prophetically remarking "So don't be ill any more my dear Charley till I can be with you to nurse you." He found what they called "Macaw Cottage" (because of its gaudy interiors) in ], then moved his "museum" in over Christmas. On 24 January 1839 Darwin was elected a ] ].<ref>{{Harvnb|Desmond|Moore|1991|pp= 272–279}}</ref>
			{{Blockquote|In October 1838, that is, fifteen months after I had begun my systematic enquiry, I happened to read for amusement Malthus on Population, and being well prepared to appreciate the struggle for existence which everywhere goes on from long-continued observation of the habits of animals and plants, it at once struck me that under these circumstances favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species. Here, then, I had at last got a theory by which to work...<ref name="Darwin 1958-2">{{harvnb|Darwin|1958|p=}}</ref>}}
	On 29 January Darwin and Emma Wedgwood were married at Maer in an Anglican ceremony arranged to suit the Unitarians, then immediately caught the train to London and their new home.<ref>{{Harvnb|Desmond|Moore|1991|p= 279}}</ref>
			By mid-December, Darwin saw a similarity between farmers picking the best stock in selective breeding, and a Malthusian Nature selecting from chance variants so that "every part of newly acquired structure is fully practical and perfected",<ref>{{cite web |url=http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=CUL-DAR124.-&pageseq=63 |title=Darwin transmutation notebook E p. 75 |access-date=17 March 2009 |url-status=live |archive-url=https://web.archive.org/web/20090628082830/http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=CUL-DAR124.-&pageseq=63 |archive-date=28 June 2009 }}</ref> thinking this comparison "a beautiful part of my theory".<ref>{{cite web|url=http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=CUL-DAR124.-&pageseq=61|title=Darwin transmutation notebook E p. 71|access-date=17 March 2009|url-status=live|archive-url=https://web.archive.org/web/20090628080656/http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=CUL-DAR124.-&pageseq=61|archive-date=28 June 2009}}</ref> He later called his theory ], an analogy with what he termed the "artificial selection" of selective breeding.<ref name="van Wyhe 2008" />
	===Preparing the theory of natural selection for publication===
	{{details|Development of Darwin's theory}}
			On 11 November, he returned to Maer and proposed to Emma, once more telling her his ideas. She accepted, then in exchanges of loving letters showed how she valued his openness in sharing their differences, while expressing her strong Unitarian beliefs and concerns that his honest doubts might separate them in the afterlife.<ref name="Darwin Correspondence Project">{{cite web|url=http://www.darwinproject.ac.uk/content/view/130/125/|title=Darwin Correspondence Project&nbsp;– Belief: historical essay|access-date=25 November 2008 |archive-url=https://web.archive.org/web/20090225124103/http://www.darwinproject.ac.uk/content/view/130/125/ |archive-date=25 February 2009 }}</ref> While he was house-hunting in London, bouts of illness continued and Emma wrote urging him to get some rest, almost prophetically remarking "So don't be ill any more my dear Charley till I can be with you to nurse you." He found what they called "Macaw Cottage" (because of its gaudy interiors) in ], then moved his "museum" in over Christmas. On 24 January 1839, Darwin was ] (FRS).<ref name="catalogues.royalsociety.org-2015">{{cite web | title=Search Results: Record – Darwin; Charles Robert | website=The Royal Society Collections Catalogues | date=20 June 2015 | url=https://catalogues.royalsociety.org/CalmView/Record.aspx?src=CalmView.Persons&id=NA8196&pos=1 | access-date=2 December 2021 | archive-date=2 December 2021 | archive-url=https://web.archive.org/web/20211202200044/https://catalogues.royalsociety.org/CalmView/Record.aspx?src=CalmView.Persons&id=NA8196&pos=1 | url-status=live }}</ref><ref>{{Harvnb|Desmond|Moore|1991|pp=272–279}}</ref>
	Darwin now had the framework of his theory of ] "by which to work",<ref>{{Harvnb|Darwin|1958|p=}}</ref> as his "prime hobby".<ref name=Letter419>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-419.html|title=Darwin Correspondence Project - Letter 419 — Darwin, C. R. to Fox, W. D., (15 June 1838)|accessdate=2008-02-08}}</ref> His research included ] and extensive experiments with plants, finding evidence that species were not fixed and investigating many detailed ideas to refine and substantiate his theory.<ref name=JvW/> For fifteen years this work was in the background to his main occupation of writing on geology and publishing expert reports on the ''Beagle'' collections.<ref name=vw186>{{Harvnb|van Wyhe|2007|pp=}}</ref>
			On 29 January, Darwin and Emma Wedgwood were married at Maer in an Anglican ceremony arranged to suit the Unitarians, then immediately caught the train to London and their new home.<ref>{{Harvnb|Desmond|Moore|1991|p=279}}</ref>
	When FitzRoy's ''Narrative'' was published in May 1839, Darwin's '']'' was such a success as the third volume that later that year it was published on its own.<ref>{{Harvnb|Darwin|1887|loc=}}</ref> Early in 1842, Darwin wrote about his ideas to ], who noted that his ally "denies seeing a beginning to each crop of species".<ref>{{Harvnb|Desmond|Moore|1991|p=292}}</ref>
			===Geology books, barnacles, evolutionary research===
	Darwin's book '']'' on his theory of ] formation was published in May 1842 after more than three years of work, and he then wrote his first "pencil sketch" of his theory of natural selection.<ref>{{Harvnb|Desmond|Moore|1991|pp=292–293}}<br />{{Harvnb|Darwin|1842|pp=}}</ref> To escape the pressures of London, the family moved to rural ] in September.<ref>{{Harvnb|Darwin|1954|p=}}</ref> On 11 January 1844 Darwin mentioned his theorising to the botanist ], writing with melodramatic humour "it is like confessing a murder".<ref>{{harvnb|van Wyhe|2007|pp=}}</ref><ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-729.html#back-mark-729.f6|title=Darwin Correspondence Project - Letter 729 — Darwin, C. R. to Hooker, J. D., (11 January 1844)|accessdate=2008-02-08}}</ref> Hooker replied "There may in my opinion have been a series of productions on different spots, & also a gradual change of species. I shall be delighted to hear how you think that this change may have taken place, as no presently conceived opinions satisfy me on the subject."<ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-734.html|title=Darwin Correspondence Project - Letter 734 — Hooker, J. D. to Darwin, C. R., 29 January 1844|accessdate=2008-02-08}}</ref>
			{{further|Development of Darwin's theory}}
			].]]
	] was his usual "Thinking Path".<ref>{{harvnb|Darwin|1887|pp=}}</ref>]]
			Darwin now had the framework of his theory of natural selection "by which to work",<ref name="Darwin 1958-2" /> as his "prime hobby".<ref name="Darwin Correspondence Project-6">{{cite news |url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-419.html |title=Darwin Correspondence Project&nbsp;– Letter 419&nbsp;– Darwin, C. R. to Fox, W. D., (15 June 1838) |newspaper=Darwin Correspondence Project |access-date=8 February 2008 |url-status=live |archive-url=https://web.archive.org/web/20070904124133/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-419.html |archive-date=4 September 2007}}</ref> His research included extensive experimental selective breeding of plants and animals, finding evidence that species were not fixed and investigating many detailed ideas to refine and substantiate his theory.<ref name="van Wyhe 2008" /> For fifteen years this work was in the background to his main occupation of writing on geology and publishing expert reports on the ''Beagle'' collections, in particular, the ]s.<ref name="van Wyhe 2007-2">{{Harvnb|van Wyhe|2007|pp=}}</ref>
	By July, Darwin had expanded his "sketch" into a 230-page "Essay", to be expanded with his research results if he died prematurely.<ref>{{Harvnb|van Wyhe|2007|p= 188}}</ref> In November the anonymously published sensational best-seller '']'' brought wide interest in transmutation. Darwin scorned its amateurish geology and zoology, but carefully reviewed his own arguments. Controversy erupted, and it continued to sell well despite contemptuous dismissal by scientists.<ref>{{harvnb|Browne|1995|pp=461-465}}</ref><ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-814.html#back-mark-814.f5|title=Darwin Correspondence Project - Letter 814 — Darwin, C. R. to Hooker, J. D., (7 Jan 1845)|accessdate=2008-11-24}}</ref>
			The impetus of Darwin's barnacle research came from a collection of a barnacle colony from Chile in 1835, which he dubbed ]. His confusion over the relationship of this species (''Cryptophialus minutus'') to other barnacles caused him to fixate on the systematics of the taxa. He wrote his first examination of the species in 1846 but did not formally describe it until 1854.<ref name="buch">{{cite journal |last1=Buchanan |first1=Roderick D. |title=Darwin's "Mr. Arthrobalanus": Sexual Differentiation, Evolutionary Destiny and the Expert Eye of the Beholder |journal=Journal of the History of Biology |date=May 2017 |volume=50 |issue=2 |pages=315–355 |doi=10.1007/s10739-016-9444-9 |pmid=27098777 |url=https://pubmed.ncbi.nlm.nih.gov/27098777/ |access-date=3 September 2024 |issn=1573-0387}}</ref>
	Darwin completed his third geological book in 1846. He now renewed a fascination and expertise in ], dating back to his student days with ], by dissecting and classifying the ]s he had collected on the voyage, enjoying observing beautiful structures and thinking about comparisons with allied structures.<ref>{{Harvnb|van Wyhe|2007|pp=190–191}}</ref> In 1847, Hooker read the "Essay" and sent notes that provided Darwin with the calm critical feedback that he needed, but would not commit himself and questioned Darwin's opposition to continuing acts of ].<ref>{{Harvnb|Desmond|Moore|1991|pp= 320–323, 339–348}}</ref>
			FitzRoy's long-delayed ''Narrative'' was published in May 1839. Darwin's ''Journal and Remarks'' got good reviews as the third volume, and on 15 August it was published on its own. Early in 1842, Darwin wrote about his ideas to Charles Lyell, who noted that his ally "denies seeing a beginning to each crop of species".<ref name="Darwin 1839" /><ref>{{Harvnb|Desmond|Moore|1991|pp=284–285, 292}}</ref>
	In an attempt to improve his chronic ill health, Darwin went in 1849 to Dr. ]'s ] spa and was surprised to find some benefit from ].<ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-1236.html|title=Darwin Correspondence Project - Letter 1236 — Darwin, C. R. to Hooker, J. D., 28 Mar 1849|accessdate=2008-11-24}}</ref> Then in 1851 his treasured daughter ] fell ill, reawakening his fears that his illness might be hereditary, and after a long series of crises she died.<ref>{{harvnb|Browne|1995|pp=498–501}}</ref>
			Darwin's book '']'' on his theory of atoll formation was published in May 1842 after more than three years of work, and he then wrote his first "pencil sketch" of his theory of natural selection.<ref>{{Harvnb|Desmond|Moore|1991|pp=292–293}}<br />{{Harvnb|Darwin|1842|pp=}}</ref> To escape the pressures of London, the family moved to rural ] in Kent in September.<ref>{{Harvnb|Darwin|1958|p=}}</ref> On 11 January 1844, Darwin mentioned his theorising to the botanist ], writing with melodramatic humour "it is like confessing a murder".<ref>{{harvnb|van Wyhe|2007|pp=}}</ref><ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-729.html#back-mark-729.f6|title=Darwin Correspondence Project&nbsp;– Letter 729&nbsp;– Darwin, C. R. to Hooker, J. D., (11 January 1844)|access-date=8 February 2008|url-status=live|archive-url=https://web.archive.org/web/20080307235150/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-729.html#back-mark-729.f6|archive-date=7 March 2008}}</ref> Hooker replied, "There may, in my opinion, have been a series of productions on different spots, & also a gradual change of species. I shall be delighted to hear how you think that this change may have taken place, as no presently conceived opinions satisfy me on the subject."<ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-734.html|title=Darwin Correspondence Project&nbsp;– Letter 734&nbsp;– Hooker, J. D. to Darwin, C. R., 29 January 1844|access-date=8 February 2008|url-status=live|archive-url=https://web.archive.org/web/20090226141303/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-734.html|archive-date=26 February 2009}}</ref>
	In eight years of work on barnacles (Cirripedia), Darwin's theory helped him to find "]" showing that slightly changed body parts served different functions to meet new conditions, and in some ] he found minute males ] on ]s, showing an ] in evolution of ].<ref name=barlowbio117>{{Harvnb|Darwin|1954|pp=}}</ref> In 1853 it earned him the ]'s Royal Medal, and it made his reputation as a ].<ref>{{Harvnb|Desmond|Moore|1991|pp= 383–387}}</ref> He resumed work on his theory of species in 1854, and in November realised that divergence in the character of descendants could be explained by them becoming adapted to "diversified places in the economy of nature".<ref>{{Harvnb|Desmond|Moore|1991|pp= 419–420}}</ref>
			] in Kent was his usual "thinking path"<ref name="Darwin Online-3">{{cite web | title=Charles Darwin: a life in pictures, The Sand Walk near Down House, Darwin's thinking path | website=Darwin Online | url=http://darwin-online.org.uk/life19.html | access-date=1 October 2022 | archive-date=1 October 2022 | archive-url=https://web.archive.org/web/20221001173155/http://darwin-online.org.uk/life19.html | url-status=live }}</ref>]]
			By July, Darwin had expanded his "sketch" into a 230-page "Essay", to be expanded with his research results if he died prematurely.<ref>{{Harvnb|van Wyhe|2007|p=188}}</ref> In November, the anonymously published sensational best-seller '']'' brought wide interest in transmutation. Darwin scorned its amateurish geology and zoology, but carefully reviewed his own arguments. Controversy erupted, and it continued to sell well despite contemptuous dismissal by scientists.<ref>{{harvnb|Browne|1995|pp=461–465}}</ref><ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-814.html#back-mark-814.f5|title=Darwin Correspondence Project&nbsp;– Letter 814&nbsp;– Darwin, C. R. to Hooker, J. D., (7 Jan 1845)|access-date=24 November 2008|url-status=live|archive-url=https://web.archive.org/web/20081205084645/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-814.html#back-mark-814.f5|archive-date=5 December 2008}}</ref>
			Darwin completed his third geological book in 1846. He now renewed a fascination and expertise in marine invertebrates, dating back to his student days with Grant, by dissecting and classifying the barnacles he had collected on the voyage, enjoying observing beautiful structures and thinking about comparisons with allied structures.<ref>{{Harvnb|van Wyhe|2007|pp=190–191}}</ref> In 1847, Hooker read the "Essay" and sent notes that provided Darwin with the calm critical feedback that he needed, but would not commit himself and questioned Darwin's opposition to continuing acts of ].<ref>{{Harvnb|Desmond|Moore|1991|pp=320–323, 339–348}}</ref>
			In an attempt to improve his chronic ill health, Darwin went in 1849 to Dr. ]'s ] spa and was surprised to find some benefit from ].<ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-1236.html|title=Darwin Correspondence Project&nbsp;– Letter 1236&nbsp;– Darwin, C. R. to Hooker, J. D., 28 Mar 1849|access-date=24 November 2008|url-status=live|archive-url=https://web.archive.org/web/20081207005457/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-1236.html|archive-date=7 December 2008}}</ref> Then, in 1851, his treasured daughter ] fell ill, reawakening his fears that his illness might be hereditary. She died the same year after a long series of crises.<ref>{{harvnb|Browne|1995|pp=498–501}}</ref>
			In eight years of work on barnacles, Darwin's theory helped him to find "]" showing that slightly changed body parts served different functions to meet new conditions, and in some ] he found minute males ] on ]s, showing an ] in evolution of ].<ref name="Darwin 1958-3">{{Harvnb|Darwin|1958|pp=}}</ref> In 1853, it earned him the ]'s Royal Medal, and it made his reputation as a ].<ref>{{Harvnb|Desmond|Moore|1991|pp=383–387}}</ref> Upon the conclusion of his research, Darwin declared "I hate a barnacle as no man ever did before."<ref>{{Cite journal |last=Bromham |first=Lindell |date=1 October 2020 |title=Comparability in evolutionary biology: The case of Darwin's barnacles |url=https://www.degruyter.com/document/doi/10.1515/lingty-2020-2056/html?lang=en |journal=Linguistic Typology |language=en |volume=24 |issue=3 |pages=427–463 |doi=10.1515/lingty-2020-2056 |issn=1613-415X |s2cid=222319487 |hdl-access=free |hdl=1885/274303}}</ref>{{sfn|van Wyhe|2007}} In 1854, he became a Fellow of the ], gaining postal access to its library.<ref>{{harvnb|Freeman|2007|pp=107, 109}}</ref> He began a major reassessment of his theory of species, and in November realised that divergence in the character of descendants could be explained by them becoming adapted to "diversified places in the economy of nature".<ref>{{Harvnb|Desmond|Moore|1991|pp=419–420}}</ref>
	===Publication of the theory of natural selection===		===Publication of the theory of natural selection===
	{{details|Publication of Darwin's theory}}		{{further|Publication of Darwin's theory}}
	].]]
			]. He wrote to ] about this portrait, "if I really have as bad an expression, as my photograph gives me, how I can have one single friend is surprising."<ref> {{webarchive|url=https://web.archive.org/web/20120107045842/http://darwin-online.org.uk/EditorialIntroductions/vanWyhe_MaullandPolyblankPhoto.html |date=7 January 2012 }}, John van Wyhe, December 2006</ref>]]
			By the start of 1856, Darwin was investigating whether eggs and ]s could survive travel across seawater to spread species across oceans. Hooker increasingly doubted the traditional view that species were fixed, but their young friend ] was still firmly against the transmutation of species. Lyell was intrigued by Darwin's speculations without realising their extent. When he read a paper by ], "On the Law which has Regulated the Introduction of New Species", he saw similarities with Darwin's thoughts and urged him to publish to establish precedence.<ref name="Desmond-2">{{Harvnb|Desmond|Moore|1991|pp=412–441, 457–458, 462–463}}<br />{{harvnb | Desmond |Moore | 2009 | pp=283–284, 290–292, 295}}</ref>
			Though Darwin saw no threat, on 14 May 1856 he began writing a short paper. Finding answers to difficult questions held him up repeatedly, and he expanded his plans to a "big book on species" titled '']'', which was to include his "note on Man". He continued his research, ] and specimens from naturalists worldwide, including Wallace who was working in ].<ref name="Desmond-2" />
			In mid-1857, he added a section heading, "Theory applied to Races of Man", but did not add text on this topic. On 5 September 1857, Darwin sent the American botanist ] a detailed outline of his ideas, including an abstract of ''Natural Selection'', which omitted ] and ]. In December, Darwin received a letter from Wallace asking if the book would examine human origins. He responded that he would avoid that subject, "so surrounded with prejudices", while encouraging Wallace's theorising and adding that "I go much further than you."<ref name="Desmond-2" />
			Darwin's book was only partly written when, on 18 June 1858, he received a paper from Wallace describing natural selection. Shocked that he had been "forestalled", Darwin sent it on that day to Lyell, as requested by Wallace,<ref>Ball, P. (2011). Shipping timetables debunk Darwin plagiarism accusations: Evidence challenges claims that Charles Darwin stole ideas from Alfred Russel Wallace. Nature. {{webarchive|url=https://web.archive.org/web/20120222191430/http://www.nature.com/news/shipping-timetables-debunk-darwin-plagiarism-accusations-1.9613 |date=22 February 2012 }}</ref><ref>{{cite journal |doi=10.1111/j.1095-8312.2011.01808.x|title=A new theory to explain the receipt of Wallace's Ternate Essay by Darwin in 1858|journal=Biological Journal of the Linnean Society|volume=105|pages=249–252|year=2012|last1=van Wyhe|first1=John|last2=Rookmaaker|first2=Kees|doi-access=free}}</ref> and although Wallace had not asked for publication, Darwin suggested he would send it to any journal that Wallace chose. His family was in crisis, with children in the village dying of ], and he put matters in the hands of his friends. After some discussion, with no reliable way of involving Wallace, Lyell and Hooker decided on a joint presentation at the Linnean Society on 1 July of '']''. On the evening of 28 June, Darwin's baby son died of scarlet fever after almost a week of severe illness, and he was too distraught to attend.<ref>{{Harvnb|Desmond|Moore|1991|pp=466–470}}</ref>
			There was little immediate attention to this announcement of the theory; the president of the Linnean Society remarked in May 1859 that the year had not been marked by any revolutionary discoveries.<ref>{{Harvnb|Browne|2002|pp=40–42, 48–49}}</ref> Only one review rankled enough for Darwin to recall it later; Professor ] of Dublin claimed that "all that was new in them was false, and what was true was old".<ref>{{Harvnb|Darwin|1958|p=}}</ref> Darwin struggled for thirteen months to produce an abstract of his "big book", suffering from ill health but getting constant encouragement from his scientific friends. Lyell arranged to have it published by ].<ref>{{Harvnb|Desmond|Moore|1991|pp=374–474}}</ref>
			'']'' proved unexpectedly popular, with the entire stock of 1,250 copies oversubscribed when it went on sale to booksellers on 22 November 1859.<ref>{{Harvnb|Desmond|Moore|1991|p=477}}</ref> In the book, Darwin set out "one long argument" of detailed observations, inferences and consideration of anticipated objections.<ref>{{Harvnb|Darwin|1859|loc=}}</ref> In making the case for common descent, he included evidence of homologies between humans and other mammals.{{sfn|van Wyhe|2008}}{{Ref label|C|III|none}} Having outlined sexual selection, he hinted that it could explain differences between ].<ref name="Darwin 1859-2">{{Harvnb|Darwin|1859|p=}}<br />{{harvnb | Darwin |Costa | 2009 | p=199}}<br />{{harvnb | Desmond |Moore | 2009 | p=310}}</ref>{{Ref label|D|IV|1}} He avoided explicit discussion of human origins, but implied the significance of his work with the sentence; "Light will be thrown on the origin of man and his history."<ref name="Darwin 1859">{{harvnb|Darwin|1859|p=}}<br />{{harvnb | Darwin |Costa | 2009 | pp=199, 488}}<br />{{harvnb|van Wyhe|2008}}</ref>{{Ref label|D|IV|2}} His theory is simply stated in the introduction:
			{{blockquote|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.<ref>{{Harvnb|Darwin|1859|loc=}}</ref>}}
			At the end of the book, he concluded that:
			{{blockquote|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.<ref>{{Harvnb|Darwin|1859|loc=}}</ref>}}
			The last word was the only variant of "evolved" in the first five editions of the book. "]" at that time was associated with other concepts, most commonly with ]. Darwin first used the word ] in '']'' in 1871, before adding it in 1872 to the 6th edition of ''The Origin of Species''.<ref>{{harvnb|Browne|2002|p=59}}, {{harvnb|Freeman|1977|pp=}}</ref>
			===Responses to publication===
	By the start of 1856, Darwin was investigating whether eggs and ]s could survive travel across seawater to spread species across oceans. ] increasingly doubted the traditional view that species were fixed, but their young friend ] was firmly against evolution. ] was intrigued by Darwin's speculations without realising their extent. When he read a paper by ] on the ''Introduction'' of species, he saw similarities with Darwin's thoughts and urged him to publish to establish precedence. Though Darwin saw no threat, he began work on a short paper. Finding answers to difficult questions held him up repeatedly, and he expanded his plans to a "big book on species" titled ''Natural Selection''. He continued his researches, ] and specimens from naturalists worldwide including Wallace who was working in ]. The American botanist ] showed similar interests, and on 5 September 1857 Darwin sent Gray a detailed outline of his ideas including an abstract of ''Natural Selection''. In December, Darwin received a letter from Wallace asking if the book would examine ]. He responded that he would avoid that subject, "so surrounded with prejudices", while encouraging Wallace's theorising and adding that "I go much further than you."<ref>{{Harvnb|Desmond|Moore|1991|pp= 412–441, 457–458, 462–463}}</ref>
			{{further|Reactions to On the Origin of Species}}
			].<ref name="Browne 2002-2" />]]
			]'' was typical of many showing Darwin with an ] body, identifying him in popular culture as the leading author of evolutionary theory.<ref name="Browne 2002-2" />]]
			The book aroused international interest, with less controversy than had greeted the popular and less scientific ''Vestiges of the Natural History of Creation''.<ref>{{harvnb|van Wyhe|2008b|p=48}}</ref> Though Darwin's illness kept him away from the public debates, he eagerly scrutinised the scientific response, commenting on press cuttings, reviews, articles, satires and caricatures, and corresponded on it with colleagues worldwide.<ref>{{Harvnb|Browne|2002|pp=103–104, 379}}</ref> The book did not explicitly discuss human origins,<ref name="Darwin 1859" />{{Ref label|D|IV|3}} but included a number of hints about the animal ancestry of humans from which the inference could be made.<ref>{{harvnb|Radick|2013|pp=174–175}}<br />{{harvnb|Huxley|Kettlewell|1965|p=88}}</ref>
	Darwin's book was half way when, on 18 June 1858, he received a paper from Wallace describing natural selection. Shocked that he had been "forestalled", Darwin sent it on to Lyell, as requested, and, though Wallace had not asked for publication, he suggested he would send it to any journal that Wallace chose. His family was in crisis with children in the village dying of ], and he put matters in the hands of Lyell and Hooker. They decided on a joint presentation at the ] on 1 July of '']''; however, Darwin's baby son died of the scarlet fever and he was too distraught to attend.<ref>{{Harvnb|Desmond|Moore|1991|pp= 466–470}}</ref>
			The first review asked, "If a monkey has become a man – what may not a man become?" It said this should be left to theologians as being too dangerous for ordinary readers.<ref>{{harvnb|Browne|2002|p=87}}<br />{{harvnb|Leifchild|1859}}</ref> Among early favourable responses, Huxley's reviews swiped at ], leader of the scientific establishment which Huxley was trying to overthrow.<ref>{{Harvnb|Desmond|Moore|1991|pp=477–491}}</ref>
	There was little immediate attention to this announcement of the theory; the president of the Linnean Society remarked in May 1859 that the year had not been marked by any revolutionary discoveries.<ref>{{Harvnb|Browne|2002|pp=40–42, 48–49}}</ref> Only one review rankled enough for Darwin to recall it later; Professor ] of Dublin claimed that "all that was new in them was false, and what was true was old."<ref>{{Harvnb|Darwin|1958|p=}}</ref> Darwin struggled for thirteen months to produce an abstract of his "big book", suffering from ill health but getting constant encouragement from his scientific friends. Lyell arranged to have it published by ].<ref>{{Harvnb|Desmond|Moore|1991|pp= 374–474}}</ref>
			In April, Owen's review attacked Darwin's friends and condescendingly dismissed his ideas, angering Darwin,<ref>{{harvnb|Browne|2002|pp=110–112}}</ref> but Owen and others began to promote ideas of supernaturally guided evolution. ] drew attention to his 1831 book which had a brief appendix suggesting a concept of natural selection leading to new species, but he had not developed the idea.<ref>{{harvnb|Bowler|2003|pp=158, 186}}</ref>
	'']'' proved unexpectedly popular, with the entire stock of 1,250 copies oversubscribed when it went on sale to booksellers on 22 November 1859.<ref>{{Harvnb|Desmond|Moore|1991|p= 477}}</ref> In the book, Darwin set out "one long argument" of detailed observations, inferences and consideration of anticipated objections.<ref>{{Harvnb|Darwin|1859|loc= }}</ref> His only allusion to human evolution was the understatement that "light will be thrown on the origin of man and his history".<ref>{{Harvnb|Darwin|1859|loc= }}</ref> His theory is simply stated in the introduction:
	{{quotation|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.<ref>{{Harvnb|Darwin|1859|loc= }}</ref>}}
			The ]'s response was mixed. Darwin's old Cambridge tutors Sedgwick and Henslow dismissed the ideas, but ] interpreted natural selection as an instrument of God's design, with the cleric ] seeing it as "just as noble a conception of Deity".<ref name="Darwin Correspondence Project-2007">{{cite web|url=http://www.darwinproject.ac.uk/content/view/110/104/|title=Darwin and design: historical essay|year=2007|publisher=Darwin Correspondence Project|access-date=17 September 2008 |archive-url=https://web.archive.org/web/20090615191012/http://www.darwinproject.ac.uk/content/view/110/104/ |archive-date=15 June 2009}}</ref> In 1860, the publication of '']'' by seven liberal Anglican theologians diverted ] attention from Darwin. Its ideas, including ], were attacked by church authorities as ]. In it, ] argued that ]s broke God's laws, so belief in them was ], and praised "Mr Darwin's masterly volume &#91;supporting&#93; the grand principle of the self-evolving powers of nature".<ref>{{Harvnb|Desmond|Moore|1991|pp=487–488, 500}}</ref>
	He put a strong case for ], but avoided the then controversial term "]", and at the end of the book concluded that:
	{{quotation|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.<ref>{{Harvnb|Darwin|1859|loc= }}</ref>}}
			Asa Gray discussed ] with Darwin, who imported and distributed Gray's pamphlet on ], ''Natural Selection is not inconsistent with natural theology''.<ref name="Darwin Correspondence Project-2007" /><ref name="Miles 2001">{{Harvnb|Miles|2001}}</ref> The most famous confrontation was at the public ] during a meeting of the ], where the ] ], though not opposed to transmutation of species, argued against Darwin's explanation and human descent from apes. Joseph Hooker argued strongly for Darwin, and Thomas Huxley's legendary retort, that he would rather be descended from an ape than a man who misused his gifts, came to symbolise a triumph of science over religion.<ref name="Darwin Correspondence Project-2007" /><ref>{{harvnb|Bowler|2003|p=185}}</ref>
	===Responses to the publication===
	] cottage, ] took portraits showing the bushy beard Darwin had grown by 1866.]]
	]'' was typical of many showing Darwin with an ] body, identifying him in popular culture as the leading author of evolutionary theory.<ref name=b373/>]]
	{{details|Reaction to Darwin's theory}}
	The book aroused international interest, with less controversy than had greeted the popular '']''.<ref>{{harvnb|van Wyhe|2008b|p=48}}</ref> Though Darwin's illness kept him away from the public debates, he eagerly scrutinised the scientific response, commenting on press cuttings, reviews, articles, satires and caricatures, and ] with colleagues worldwide.<ref>{{Harvnb|Browne|2002|pp=103–104, 379}}</ref> Darwin had only said "Light will be thrown on the origin of man",<ref>{{harvnb|Darwin|1859|p=}}</ref> but the first review claimed it made a creed of the "men from monkeys" idea from ''Vestiges''.<ref>{{harvnb|Browne|2002|p=87}}<br />{{harvnb|Leifchild|1859}}</ref> Amongst early favourable responses, Huxley's reviews swiped at ], leader of the scientific establishment Huxley was trying to overthrow.<ref>{{Harvnb|Desmond|Moore|1991|pp=477–491}}</ref> In April, Owen's review attacked Darwin's friends and condescendingly dismissed his ideas, angering Darwin,<ref>{{harvnb|Browne|2002|pp=110–112}}</ref> but Owen and others began to promote ideas of supernaturally guided evolution.<ref>{{harvnb|Bowler|2003|p=186}}</ref>
			Even Darwin's close friends Gray, Hooker, Huxley and Lyell still expressed various reservations but gave strong support, as did many others, particularly younger naturalists. Gray and Lyell sought reconciliation with faith, while Huxley portrayed a polarisation between religion and science. He campaigned pugnaciously against the authority of the clergy in education,<ref name="Darwin Correspondence Project-2007" /> aiming to overturn the dominance of clergymen and aristocratic amateurs under Owen in favour of a new generation of professional scientists. Owen's claim that brain anatomy proved humans to be a separate ] from apes was shown to be false by Huxley in a long-running dispute parodied by Kingsley as the "]", and discredited Owen.<ref>{{Harvnb|Browne|2002|pp=156–159}}</ref>
	The ]'s response was mixed. Darwin's old Cambridge tutors ] and ] dismissed the ideas, but ] interpreted natural selection as an instrument of God's design, with the cleric ] seeing it as "just as noble a conception of Deity".<ref name=Darwinanddesign>{{cite web|url=http://www.darwinproject.ac.uk/content/view/110/104/|title=Darwin and design: historical essay|year=2007|publisher=Darwin Correspondence Project|accessdate=2008-09-17}}</ref> In 1860, the publication of '']'' by seven liberal Anglican theologians diverted ] attention from Darwin, with its ideas including ] attacked by church authorities as ]. In it, ] argued that ]s broke God's laws, so belief in them was ], and praised "Mr Darwin's masterly volume &#91;supporting&#93; the grand principle of the self-evolving powers of nature".<ref>{{Harvnb|Desmond|Moore|1991|pp= 487–488, 500}}</ref> ] discussed ] with Darwin, who imported and distributed Gray's pamphlet on ], ''Natural Selection is not inconsistent with ]''.<ref name=Darwinanddesign/><ref name=miles>{{Harvnb|Miles|2001}}</ref> The most famous confrontation was at the public ] during a meeting of the ], where the ] ], though not opposed to ], argued against Darwin's explanation and human descent from apes. ] argued strongly for Darwin, and ]'s legendary retort, that he would rather be descended from an ape than a man who misused his gifts, came to symbolise a triumph of science over religion.<ref name=Darwinanddesign/><ref>{{harvnb|Bowler|2003|p=185}}</ref>
			In response to objections that the ] was unexplained, Darwin pointed to acceptance of ] even though the cause of gravity was unknown.<ref name="National University of Singapore News-2022">{{cite web | title=Science ahead of its time: Secret of 157-year old Darwin manuscript | website=] News | date=24 November 2022 | url=https://news.nus.edu.sg/secret-of-157-year-old-darwin-manuscript/ | access-date=25 November 2022 | archive-date=25 November 2022 | archive-url=https://web.archive.org/web/20221125100051/https://news.nus.edu.sg/secret-of-157-year-old-darwin-manuscript/ | url-status=live }}</ref> Despite criticisms and reservations related to this topic, he nevertheless proposed a prescient idea in an 1871 letter to Hooker in which he suggested the origin of life may have occurred in a "]".<ref>{{Cite journal |last1=Peretó |first1=Juli |last2=Bada |first2=Jeffrey L. |last3=Lazcano |first3=Antonio |date=1 October 2009 |title=Charles Darwin and the Origin of Life |url=https://doi.org/10.1007/s11084-009-9172-7 |journal=Origins of Life and Evolution of Biospheres |language=en |volume=39 |issue=5 |pages=395–406 |doi=10.1007/s11084-009-9172-7 |issn=1573-0875 |pmc=2745620 |pmid=19633921 |access-date=8 December 2023 |archive-date=18 December 2023 |archive-url=https://web.archive.org/web/20231218114111/https://link.springer.com/article/10.1007/s11084-009-9172-7 |url-status=live }}</ref>
			] became a movement covering a wide range of evolutionary ideas. In 1863, Lyell's '']'' popularised prehistory, though his caution on evolution disappointed Darwin. Weeks later Huxley's '']'' showed that anatomically, humans are apes, then '']'' by ] provided empirical evidence of natural selection.<ref name="Browne 2002">{{harvnb|Browne|2002|pp=217–226}}</ref> Lobbying brought Darwin Britain's highest scientific honour, the Royal Society's ], awarded on 3 November 1864.<ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-4652.html|title=Darwin Correspondence Project&nbsp;– Letter 4652&nbsp;– Falconer, Hugh to Darwin, C. R., 3 Nov (1864)|access-date=1 December 2008|url-status=live|archive-url=https://web.archive.org/web/20081205084616/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-4652.html|archive-date=5 December 2008}}</ref> That day, Huxley held the first meeting of what became the influential "]" devoted to "science, pure and free, untrammelled by religious dogmas".<ref name="Darwin Correspondence Project-7">{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-4807.html#mark-4807.f8|title=Darwin Correspondence Project&nbsp;– Letter 4807&nbsp;– Hooker, J. D. to Darwin, C. R., (7–8 Apr 1865)|access-date=1 December 2008|url-status=live|archive-url=https://web.archive.org/web/20081205084621/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-4807.html#mark-4807.f8|archive-date=5 December 2008}}</ref> By the end of the decade, most scientists agreed that evolution occurred, but only a minority supported Darwin's view that the chief mechanism was natural selection.<ref>{{harvnb|Bowler|2003|p=196}}</ref>
	Even Darwin's close friends Gray, Hooker, Huxley and Lyell still expressed various reservations but gave strong support, as did many others, particularly younger naturalists. Gray and Lyell sought reconciliation with faith, while Huxley portrayed a polarisation between religion and science. He campaigned pugnaciously against the authority of the clergy in education,<ref name=Darwinanddesign/> aiming to overturn the dominance of clergymen and aristocratic amateurs under Owen in favour of a new generation of professional scientists. Owen's claim that ] anatomy proved humans to be a separate ] from apes was shown to be false by Huxley in a long running dispute parodied by Kingsley as the "]", and discredited Owen.<ref>{{Harvnb|Browne|2002|pp=156–159}}</ref>
			The ''Origin of Species'' was translated into many languages, becoming a staple scientific text attracting thoughtful attention from all walks of life, including the "working men" who flocked to Huxley's lectures.<ref>{{harvnb|Desmond|Moore|1991|pp=507–508}}<br />{{Harvnb|Browne|2002|pp=128–129, 138}}</ref> Darwin's theory resonated with various movements at the time{{Ref label|E|V|none}} and became a key fixture of popular culture.{{Ref label|F|VI|none}} Cartoonists parodied animal ancestry in an old tradition of showing humans with animal traits, and in Britain, these droll images served to popularise Darwin's theory in an unthreatening way. While ill in 1862, Darwin began growing a beard, and when he reappeared in public in 1866, caricatures of him as an ] helped to identify all forms of evolutionism with Darwinism.<ref name="Browne 2002-2">{{harvnb|Browne|2002|pp=373–379}}</ref>
	] became a movement covering a wide range of evolutionary ideas. In 1863 ] '']'' popularised prehistory, though his caution on evolution disappointed Darwin. Weeks later Huxley's '']'' showed that anatomically, humans are apes, then '']'' by ] provided empirical evidence of natural selection.<ref name=B217>{{harvnb|Browne|2002|pp=217–226}}</ref> Lobbying brought Darwin Britain's highest scientific honour, the ]'s ], awarded on 3 November 1864.<ref>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-4652.html|title=Darwin Correspondence Project - Letter 4652 — Falconer, Hugh to Darwin, C. R., 3 Nov (1864)|accessdate=2008-12-01}}</ref> That day, Huxley held the first meeting of what became the influential '']'' devoted to "science, pure and free, untrammelled by religious dogmas".<ref name=Letter4807>{{cite web|url=http://www.darwinproject.ac.uk/darwinletters/calendar/entry-4807.html#mark-4807.f8|title=Darwin Correspondence Project - Letter 4807 — Hooker, J. D. to Darwin, C. R., (7–8 Apr 1865)|accessdate=2008-12-01}}</ref> By the end of the decade most scientists agreed that evolution occurred, but only a minority supported Darwin's view that the chief mechanism was natural selection.<ref>{{harvnb|Bowler|2003|p=196}}</ref>
			], America's first palaeontologist, was the first to provide solid fossil evidence to support Darwin's theory of evolution by unearthing the ancestors of the modern horse.<ref>Plate, Robert. ''The Dinosaur Hunters: Othniel C. Marsh and Edward D. Cope,'' pp. 69, 203-5, David McKay Company, Inc., New York, 1964.</ref> In 1877, Marsh delivered a very influential speech before the annual meeting of the American Association for the Advancement of Science, providing a demonstrative argument for evolution. For the first time, Marsh traced the evolution of vertebrates from fish all the way through humans. Sparing no detail, he listed a wealth of fossil examples of past life forms. The significance of this speech was immediately recognized by the scientific community, and it was printed in its entirety in several scientific journals.<ref>McCarren, Mark J. ''The Scientific Contributions of Othniel Charles Marsh,'' pp. 37-9, Peabody Museum of Natural History, Yale University, New Haven, Connecticut, 1993. {{ISBN|0-912532-32-7}}</ref><ref>Plate, Robert. ''The Dinosaur Hunters: Othniel C. Marsh and Edward D. Cope,'' pp. 188-9, David McKay Company, Inc., New York, 1964.</ref>
	The ''Origin of Species'' was translated into many languages, becoming a staple scientific text attracting thoughtful attention from all walks of life, including the "working men" who flocked to Huxley's lectures.<ref>{{harvnb|Desmond|Moore|1991|pp=507–508}}<br />{{Harvnb|Browne|2002|pp=128–129, 138}}</ref> Darwin's theory also resonated with various movements at the time{{Ref_label|C|III|none}} and became a key fixture of ].{{Ref_label|D|IV|none}} Cartoonists parodied animal ancestry in an old tradition of showing humans with animal traits, and in Britain these droll images served to popularise Darwin's theory in an unthreatening way. While ill in 1862 Darwin began growing a beard, and when he reappeared in public in 1866 caricatures of him as an ] helped to identify all forms of ] with Darwinism.<ref name=b373>{{harvnb|Browne|2002|pp=373–379}}</ref>
	===''Descent of Man'', sexual selection, and botany===		===''Descent of Man'', sexual selection, and botany===
			{{further|Darwin from Orchids to Variation|Darwin from Descent of Man to Emotions|Darwin from Insectivorous Plants to Worms|label 1=Orchids to Variation|label 2=Descent of Man to Emotions|label 3=Insectivorous Plants to Worms}}
	]
			]
	:''More detailed articles cover Darwin's life from ], from ] and from ]''
	Despite repeated bouts of illness during the last twenty-two years of his life, Darwin's work continued. Having published '']'' as an ] of his theory, he pressed on with experiments, research, and writing of his "big book". He covered ] from earlier animals including evolution of society and of mental abilities, as well as explaining decorative beauty in wildlife and diversifying into innovative plant studies.		Despite repeated bouts of illness during the last twenty-two years of his life,{{sfn|Desmond|Moore|Browne|2007|pp=73–75}} Darwin's work continued. Having published ''On the Origin of Species'' as an ] of his theory, he pressed on with experiments, research, and writing of his "]". He covered human descent from earlier animals, including the evolution of society and of mental abilities, as well as explaining decorative beauty in ] and diversifying into innovative plant studies.{{sfn|Desmond|Moore|Browne|2007|pp=78–83, 86–90}}
	Enquiries about insect ] led in 1861 to novel studies of wild ]s, showing adaptation of their flowers to ] to each species and ensure ]. In 1862 '']'' gave his first detailed demonstration of the power of natural selection to explain complex ] relationships, making testable predictions. As his health declined, he lay on his sickbed in a room filled with inventive experiments to trace the movements of ].<ref>{{harvnb|van Wyhe|2008b|pp=50–55}}</ref> Admiring visitors included ], a zealous proponent of ''Darwinismus'' incorporating ] and ]'s idealism.<ref> Cambridge University Press. Retrieved on 2008-11-28</ref> Wallace remained supportive, though he increasingly turned to ].<ref>{{harvnb|Smith|1999}}.</ref>		Enquiries about insect ] led in 1861 to novel studies of wild ]s, showing adaptation of their flowers to ] to each species and ensure ]. In 1862 '']'' gave his first detailed demonstration of the power of natural selection to explain complex ecological relationships, making testable predictions. As his health declined, he lay on his sickbed in a room filled with inventive experiments to trace the movements of ].<ref>{{harvnb|van Wyhe|2008b|pp=50–55}}</ref> Admiring visitors included ], a zealous proponent of Darwinism incorporating Lamarckism and ]'s idealism.<ref>{{cite web |url=https://www.darwinproject.ac.uk/letters/darwins-life-letters/darwin-letters1866-survival-fittest |title=The correspondence of Charles Darwin, volume 14: 1866 |access-date=6 March 2009 |archive-url=https://web.archive.org/web/20100605110511/http://www.darwinproject.ac.uk/correspondence-volume-14 |archive-date=5 June 2010 }} Cambridge University Press. Retrieved 25 June 2012</ref> Wallace remained supportive, though he increasingly turned to ].<ref>{{harvnb|Smith|1999}}.</ref>
	'']'' of 1868 was the first part of Darwin's planned "big book", and included his unsuccessful hypothesis of ] attempting to explain ]. It sold briskly at first, despite its size, and was translated into many languages. He wrote most of a second part, on natural selection, but it remained unpublished in his lifetime.<ref>{{Harvnb|Freeman|1977|p=}}</ref>		Darwin's book '']'' (1868) was the first part of his planned "big book", and included his unsuccessful hypothesis of ] attempting to explain ]. It sold briskly at first, despite its size, and was translated into many languages. He wrote most of a second part, on natural selection, but it remained unpublished in his lifetime.<ref>{{Harvnb|Freeman|1977|p=}}</ref>
			Lyell had already popularised human prehistory, and Huxley had shown that anatomically humans are apes.<ref name="Browne 2002"/> With ''The Descent of Man, and Selection in Relation to Sex'' published in 1871, Darwin set out evidence from numerous sources that humans are animals, showing continuity of physical and mental attributes, and presented sexual selection to explain impractical animal features such as the ]'s plumage as well as human evolution of culture, differences between sexes, and physical and cultural racial classification, while emphasising that humans are all one species.<ref>{{Harvnb|Darwin|1871|pp=}}<br />{{Harvnb|Browne|2002|pp=339–343}}</ref> According to an editorial in Nature journal: "Although Charles Darwin opposed slavery and proposed that humans have a common ancestor, he also advocated a hierarchy of races, with white people higher than others."<ref>{{Cite journal |last1=Nobles |first1=Melissa |last2=Womack |first2=Chad |last3=Wonkam |first3=Ambroise |last4=Wathuti |first4=Elizabeth |date=8 June 2022 |title=Science must overcome its racist legacy: Nature's guest editors speak |journal=Nature |language=en |volume=606 |issue=7913 |pages=225–227 |doi=10.1038/d41586-022-01527-z |pmid=35676434 |bibcode=2022Natur.606..225N |s2cid=249520597 |quote="In The Descent of Man, Darwin describes what he calls the gradations between the highest men of the highest races and the lowest savages*. He uses the word 'savages' to describe Black and Indigenous people." (*see Darwin, C. R. The Descent of Man, and Selection in Relation to Sex (John Murray, 1871))|doi-access=free }}</ref>
	] ] for 1882, published shortly before Darwin's death, depicts him amidst evolution from chaos to Victorian gentleman with the title ''Man Is But A Worm''.]]
	] had already popularised human prehistory, and ] had shown that anatomically humans are apes.<ref name=B217/> With '']'' published in 1871, Darwin set out evidence from numerous sources that humans are animals, showing continuity of physical and mental attributes, and presented ] to explain impractical animal features such as the ]'s plumage as well as human evolution of ], differences between sexes, and physical and cultural ], while emphasising that humans are all one species.<ref>{{Harvnb|Darwin|1871|pp=}}<br />{{Harvnb|Browne|2002|pp=339–343}}</ref> His research using images was expanded in his 1872 book '']'', one of the first books to feature printed photographs, which discussed the ] and its continuity with the ]. Both books proved very popular, and Darwin was impressed by the general assent with which his views had been received, remarking that "everybody is talking about it without being shocked."<ref>{{Harvnb|Browne|2002|pp=359–369}}<br />{{harvnb|Darwin|1887|p=}}</ref> His conclusion was "that man with all his noble qualities, with sympathy which feels for the most debased, with benevolence which extends not only to other men but to the humblest living creature, with his god-like intellect which has penetrated into the movements and constitution of the solar system–with all these exalted powers–Man still bears in his bodily frame the indelible stamp of his lowly origin."<ref>{{Harvnb|Darwin|1871|p=}}</ref>
			{{Multiple image|total_width=400
	His evolution-related experiments and investigations led to books on ''], ], ]'', different forms of flowers on plants of the same species, and '']''. In his last book he returned to '']''.
			|image1=Darwin letter.jpg
			|alt1=handwritten letter from Charles Darwin to John Burdon-Sanderson dated 9 October 1874
			|caption1=Letter of enquiry from Charles Darwin to the physiologist ]
			|image2=Man is But a Worm.jpg
			|alt2=Darwin's figure is shown seated, dressed in a toga, in a circular frame labelled "TIME'S METER" around which a succession of figures spiral, starting with an earthworm emerging from the broken letters "CHAOS" then worms with head and limbs, followed by monkeys, apes, primitive men, a loin cloth clad hunter with a club, and a gentleman who tips his top hat to Darwin
			|caption2='']''{{'}}s ] for 1882, published shortly before Darwin's death, depicts him amidst evolution from chaos to Victorian gentleman with the title ''Man Is But A Worm''
			}}
			His research using images was expanded in his 1872 book '']'', one of the first books to feature printed photographs, which discussed the ] and its continuity with the ]. Both books proved very popular, and Darwin was impressed by the general assent with which his views had been received, remarking that "everybody is talking about it without being shocked."<ref>{{Harvnb|Browne|2002|pp=359–369}}<br />{{harvnb|Darwin|1887|p=}}</ref> His conclusion was "that man with all his noble qualities, with sympathy which feels for the most debased, with benevolence which extends not only to other men but to the humblest living creature, with his god-like intellect which has penetrated into the movements and constitution of the solar system{{snd}}with all these exalted powers{{snd}}Man still bears in his bodily frame the indelible stamp of his lowly origin."<ref>{{Harvnb|Darwin|1871|p=}}</ref>
	He died at ] on 19 April 1882. He had expected to be buried in St Mary's churchyard at ], but at the request of Darwin's colleagues, ] (President of the ]) arranged for Darwin to be given a ] and buried in ], close to ] and ].<ref>{{Harvnb|Browne|2002|pp=495–497}}</ref> Only five non-royal personages were granted that honour of a UK state funeral during the 19th century.<ref name=BBCstatefuneral/>
			His evolution-related experiments and investigations led to books on ''], ]'', different forms of flowers on plants of the same species, and '']''. He continued to collect information and exchange views from scientific correspondents all over the world, including ], whom he encouraged to persevere in her scientific work.<ref> {{Webarchive|url=https://web.archive.org/web/20200212213901/https://www.cam.ac.uk/research/news/darwins-women |date=12 February 2020 }} at ]</ref> He was the first person to recognise the significance of carnivory in plants.<ref name="Hedrich-2021">{{cite journal |last1=Hedrich |first1=Rainer |last2=Fukushima |first2=Kenji |title=On the Origin of Carnivory: Molecular Physiology and Evolution of Plants on an Animal Diet |journal=Annual Review of Plant Biology |date=17 June 2021 |volume=72 |issue=1 |pages=133–153 |doi=10.1146/annurev-arplant-080620-010429 |pmid=33434053 |s2cid=231595236 |issn=1543-5008|doi-access=free }}</ref> His botanical work{{Ref label|I|IX|none}} was interpreted and popularised by various writers including ] and ], and helped transform plant science in the late 19th century and early 20th century.<ref name="Pain-2022">{{cite journal |last1=Pain |first1=Stephanie |title=How plants turned predator |journal=Knowable Magazine |date=2 March 2022 |doi=10.1146/knowable-030122-1 |doi-access=free |url=https://knowablemagazine.org/article/living-world/2022/how-plants-turned-predator |access-date=11 March 2022 |archive-date=8 March 2022 |archive-url=https://web.archive.org/web/20220308103952/https://knowablemagazine.org/article/living-world/2022/how-plants-turned-predator |url-status=live }}</ref><ref name="Endersby-2016">{{cite journal |last1=Endersby |first1=Jim |title=Deceived by orchids: sex, science, fiction and Darwin |journal=The British Journal for the History of Science |date=June 2016 |volume=49 |issue=2 |pages=205–229 |doi=10.1017/S0007087416000352 |pmid=27278105 |s2cid=23027055 |url=http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=10359900&fulltextType=RA&fileId=S0007087416000352 |access-date=17 March 2022 |language=en |issn=0007-0874}}</ref>
	Darwin was perceived as a national hero who had changed thinking, and scientists now accepted ] as ], but few agreed with him that "natural selection has been the main but not the exclusive means of modification".<ref name=b222>{{harvnb|Bowler|2003|pp=222–225}}<br>{{Harvnb|van Wyhe|2008}}<br>{{harvnb|Darwin|1872|p=}}</ref> In "]" most favoured alternative evolutionary mechanisms, but these proved untenable, and the development of the ] with ] and ] ] from the 1930s to the 1950s brought a broad scientific consensus that ] was the basic mechanism of evolution. Research and debate has continued within this frame of reference.<ref name=b3847/>
	{{-}}
			===Death and funeral===
	==Darwin's children==
			{{See also|Darwin from Insectivorous Plants to Worms#Death}}
	{| class="toccolours" style="float: right; margin-left: 1em; margin-right: 1em; font-size: 85%; background:#e0e0ee; color:black; width:32em; max-width:50%" cellspacing="5"
	|colspan=2|<div class="center">]]]</div>
			] and Charles Darwin in the nave of ], London.]]
	<div class="center">Darwin and his eldest son ] in 1842.</div>
	|-
			In 1882, he was diagnosed with what was called "]" which then meant ] and disease of the heart. At the time of his death, the physicians diagnosed "anginal attacks", and "heart-failure"; there has since been scholarly speculation about his ].<ref name="Colp-2008">{{cite book |title=Darwin's Illness |pages=116–120 |first=Ralph |last=Colp |doi=10.5744/florida/9780813032313.003.0014 |chapter=The Final {{sic|nolink=y|reason=error in source|Illnes}} |year=2008 |isbn=978-0-8130-3231-3 }}</ref><ref name="Clayton-2010">{{cite journal |last=Clayton |first=Julie |date=24 June 2010 |title=Chagas disease 101 |journal=Nature |volume=465 |issue=n7301_supp |pages=S4–S5 |doi=10.1038/nature09220 |pmid=20571553 |bibcode=2010Natur.465S...3C |s2cid=205221512 |doi-access=free}}</ref>
	!<div class="center">Darwin's Children</div>
	|-
			He died at Down House on 19 April 1882. His last words were to his family, telling Emma, "I am not the least afraid of death{{mdash}}Remember what a good wife you have been to me{{mdash}}Tell all my children to remember how good they have been to me". While she rested, he repeatedly told Henrietta and Francis, "It's almost worthwhile to be sick to be nursed by you".<ref>{{cite web |url=http://darwin-online.org.uk/content/frameset?viewtype=side&itemID=CUL-DAR210.9&pageseq=16 |title= CUL-DAR210.9|author=Darwin, Emma|author-link=Emma Darwin |year=1882 |access-date=8 January 2009 |url-status=live |archive-url=https://web.archive.org/web/20090628080442/http://darwin-online.org.uk/content/frameset?viewtype=side&itemID=CUL-DAR210.9&pageseq=16 |archive-date=28 June 2009}}</ref>
	|]||(27 December 1839–1914)
			He had expected to be buried in St Mary's churchyard at ], but at the request of Darwin's colleagues, after public and parliamentary petitioning, ] (President of the Royal Society) arranged for Darwin to be honoured by ], close to John Herschel and ]. The funeral, held on Wednesday 26 April, was attended by thousands of people, including family, friends, scientists, philosophers and dignitaries.<ref>{{Harvnb|Desmond|Moore|1991|pp=664–677}}</ref><ref name="Westminster Abbey-2016">{{cite web | title=Westminster Abbey » Charles Darwin | website=Westminster Abbey | date=2 January 2016 | url=http://www.westminster-abbey.org/our-history/people/charles-darwin | access-date=2 January 2016 | url-status=live | archive-url=https://web.archive.org/web/20160304200905/http://www.westminster-abbey.org/our-history/people/charles-darwin | archive-date=4 March 2016 | df=dmy-all }}<br />{{Harvnb|Leff|2000|loc=}}</ref>
			==Children==
			{{further|Darwin–Wedgwood family#Charles Darwin}}
			{|class=toccolours style=float:right;clear:right;font-size:small;margin-left:1em;
			|]||style=text-align:right;|27 December 1839 –||8 September 1914
	|-		|-
	|]||(2 March 1841–23 April 1851)		|]||style=text-align:right;|2 March 1841 –||23 April 1851
	|-		|-
	|Mary Eleanor Darwin||(23 September 1842–16 October 1842)		|Mary Eleanor Darwin||style=text-align:right;|23 September 1842 –||16 October 1842
	|-		|-
	|]||(25 September 1843–1929)		|]||style=text-align:right;|{{nobr|25 September 1843 –}}||17 December 1927
	|-		|-
	|]||(9 July 1845–7 December 1912)		|]||style=text-align:right;|9 July 1845 –||7 December 1912
	|-		|-
	|]||(8 July 1847–1926)		|Elizabeth Darwin||style=text-align:right;|8 July 1847 –||8 June 1926
	|-		|-
	|]||(16 August 1848–19 September 1925)		|]||style=text-align:right;|16 August 1848 –||{{nobr|19 September 1925}}
	|-		|-
	|]||(15 January 1850–26 March 1943)		|]||style=text-align:right;|15 January 1850 –||26 March 1943
	|-		|-
	|]||(13 May 1851–29 September 1928)		|]||style=text-align:right;|13 May 1851 –||29 September 1928
	|-		|-
	|]||(6 December 1856–28 June 1858)		|Charles Waring Darwin||style=text-align:right;|6 December 1856 –||28 June 1858
	|}		|}
	] died. By then his faith in ] had dwindled, and he had stopped going to church.<ref name=jvw41>{{harvnb|van Wyhe|2008b|p=41}}</ref>]]
	The Darwins had ten children: two died in infancy, and ] death at the age of ten had a devastating effect on her parents. Charles was a devoted father and uncommonly attentive to his children.<ref name=whowas/> Whenever they fell ill he feared that they might have inherited weaknesses from ] due to the close family ties he shared with his wife and cousin, ]. He examined this topic in his writings, contrasting it with the advantages of crossing amongst many organisms.<ref>{{Harvnb|Desmond|Moore|1991|p=447}}.</ref> Despite his fears, most of the surviving children went on to have distinguished careers as notable members of the prominent ].<ref>{{harvnb|Leff|2000|loc=}}</ref>
			The Darwins had ten children: two died in infancy, and Annie's death at the age of ten had a devastating effect on her parents. Charles was a devoted father and uncommonly attentive to his children.<ref name="Leff 2000" /> Whenever they fell ill, he feared that they might have inherited weaknesses from ] due to the close family ties he shared with his ], Emma Wedgwood. He examined inbreeding in his writings, contrasting it with the advantages of ] in many species.{{sfn|Desmond|Moore|1991|p=447}}
	Of his surviving children, ], ] and ] became Fellows of the Royal Society,<ref>{{cite web |title=List of Fellows of the Royal Society / 1660 - 2006 / A-J |url=http://royalsociety.org/trackdoc.asp?id=4274&pId=1727 |accessdate=2009-09-16 |format=PDF}}</ref> distinguished as ],<ref>{{MacTutor Biography|id=Darwin}}</ref> ] and ], respectively. His son ], on the other hand, went on to be a ], ], ], ] and mentor of the statistician and ] ].<ref>Edwards, A. W. F. 2004. Darwin, Leonard (1850–1943). In: ''Oxford Dictionary of National Biography'', Oxford University Press.</ref>
			]
	==Religious views==
	{{Details|Charles Darwin's views on religion}}
	Darwin's family tradition was ] ], while his father and grandfather were ], and his ] and ] were ].<ref name=skool/> When going to Cambridge to become an ] clergyman, he did not doubt the ] of the ].<ref name=dar57/> He learnt ]'s science which, like ]'s ], sought explanations in laws of nature rather than miracles and saw ] of species as ].<ref name=syd5-7/><ref name=db/> On board the ''Beagle'', Darwin was quite ] and would quote the Bible as an authority on ].<ref name=biorelig>{{Harvnb|Darwin|1958|pp=}}</ref> He looked for "centres of creation" to explain distribution,<ref name=k356/> and related the ] found near ]s to distinct "periods of Creation".<ref name=Crows/>
			Charles Waring Darwin, born in December 1856, was the tenth and last of the children. Emma Darwin was aged 48 at the time of the birth, and the child was mentally subnormal and never learnt to walk or talk. He probably had ], which had not then been medically described. The evidence is a photograph by William Erasmus Darwin of the infant and his mother, showing a characteristic head shape, and the family's observations of the child.<ref>David P. Steensma (15 March 2005). "Down syndrome in Down House: trisomy 21, GATA1 mutations, and Charles Darwin". '']'' 105 (6) 2614–2616.</ref> Charles Waring died of scarlet fever on 28 June 1858,<ref>Freeman, R. B. (1984), ''Darwin Pedigrees'', London, p. 43.</ref> when Darwin wrote in his journal: "Poor dear Baby died."<ref>Darwin, C. R. ''Journal (1809–1881)'', p. 37.</ref> <!--The death kept Darwin from attending the first ] at the Linnean Society meeting on 1 July 1858.-->
	By his return he was ], and wondered why all religions should not be equally valid.<ref name=biorelig/> In the next few years, while intensively speculating on geology and ], he gave much thought to religion and openly discussed this with ], whose beliefs also came from intensive study and questioning.<ref name=Belief/> The ] of Paley and ] vindicated evils such as starvation as a result of a benevolent creator's laws which had an overall good effect. To Darwin, ] produced the good of adaptation but removed the need for design,<ref>{{harvnb|von Sydow|2005|pp=8–14}}</ref> and he could not see the work of an omnipotent deity in all the pain and suffering such as the ] paralysing ]s as live food for its eggs.<ref name=miles/> He still viewed organisms as perfectly adapted, and '']'' reflects theological views. Though he thought of religion as a ] survival strategy, Darwin still believed that God was the ultimate lawgiver.<ref>{{harvnb|von Sydow|2005|pp=4–5}}</ref><ref>{{Harvnb|Moore|2006}}</ref>
			Of his surviving children, ], Francis and ] became ],<ref>{{cite web|title=List of Fellows of the Royal Society, 1660–2006, A–J|url=http://royalsociety.org/trackdoc.asp?id=4274&pId=1727|access-date=16 September 2009|format=PDF|archive-date=3 February 2017|url-status=dead|archive-url=https://web.archive.org/web/20170203000229/https://royalsociety.org/trackdoc.asp?id=4274&pId=1727}}</ref> distinguished as an astronomer,<ref>{{MacTutor Biography|id=Darwin}}</ref> botanist and civil engineer, respectively. All three were knighted.<ref>Berra, Tim M. ''Darwin and His Children: His Other Legacy,'' (Oxford: 2013, Oxford UP), 101, 129, 168. George became a knight commander of the Order of the Bath in 1905. Francis was knighted in 1912. Horace became a knight commander of the KBE in 1918.</ref> Another son, ], went on to be a soldier, politician, economist, ], and mentor of the statistician and evolutionary biologist Ronald Fisher.<ref>Edwards, A. W. F. 2004. Darwin, Leonard (1850–1943). In: ''Oxford Dictionary of National Biography'', Oxford University Press.</ref>
	Darwin remained close friends with the ] of Downe, ], and continued to play a leading part in the parish work of the church,<ref>{{cite web|url=http://www.darwinproject.ac.uk/content/view/152/144/|title=Darwin Correspondence Project - Darwin and the church: historical essay|accessdate=2009-01-04}}</ref> but from around 1849 would go for a walk on Sundays while his family attended church.<ref name=jvw41/> He considered it "absurd to doubt that a man might be an ardent theist and an evolutionist"<ref name=Fordyce> — Darwin, C. R. to Fordyce, John, 7 May 1879</ref><ref name=spencer> ] 17-Sept-2009</ref> and, though reticent about his religious views, in 1879 he wrote that "I have never been an atheist in the sense of denying the existence of a God. – I think that generally ... an ] would be the most correct description of my state of mind."<ref name=Belief/><ref name=Fordyce/>
			==Views and opinions==
	The "]", published in 1915, claimed that Darwin had reverted back to Christianity on his sickbed. The claims were refuted by Darwin's children and have been dismissed as false by historians.<ref>{{harvnb|Moore|2005}}<br />{{Harvnb|Yates|2003}}</ref> His last words were to his family, telling Emma "I am not the least afraid of death – Remember what a good wife you have been to me – Tell all my children to remember how good they have been to me", then as she laid down for a rest, he repeatedly told Henrietta and Francis "It's almost worth while to be sick to be nursed by you".<ref>{{cite web|url=http://darwin-online.org.uk/content/frameset?viewtype=side&itemID=CUL-DAR210.9&pageseq=16|title= CUL-DAR210.9|author=Darwin, Emma|authorlink=Emma Darwin|year= 1882|accessdate=2009-01-08}}</ref>
			===Religious views===
	==Political interpretations==
			{{Further|Religious views of Charles Darwin}}
	Darwin's fame and popularity led to his name being associated with ideas and movements which at times had only an indirect relation to his writings, and sometimes went directly against his express comments.
			Darwin's family tradition was ] ], while his father and grandfather were ], and his baptism and boarding school were ].<ref name="Desmond-3" /> When going to Cambridge to become an Anglican clergyman, he did not "in the least doubt the strict and ] of every word in the Bible".<ref name="Desmond" /> He learned John Herschel's science which, like William Paley's natural theology, sought explanations in laws of nature rather than miracles and saw adaptation of species as evidence of design.<ref name="von Sydow 2005" /><ref name="Darwin 1958-5" /> On board HMS ''Beagle'', Darwin was quite ] and would quote the Bible as an authority on ].<ref name="Darwin 1958-4">{{Harvnb|Darwin|1958|pp=}}</ref> He looked for "centres of creation" to explain distribution,<ref name="Keynes 2001-2" /> and suggested that the very similar ]s found in Australia and England were evidence of a divine hand.<ref name="Darwin Online" />
	]'']]
			] died; by then his faith in Christianity had dwindled, and he had stopped going to church.<ref name="van Wyhe 2008b">{{harvnb|van Wyhe|2008b|p=41}}</ref>]]
	===Eugenics===
	{{details|Eugenics}}
	Darwin was interested by his ] ]'s argument, introduced in 1865, that ] of ] showed that moral and mental human traits could be inherited, and principles of animal breeding could apply to humans. In '']'' Darwin noted that aiding the weak to survive and have families could lose the benefits of ], but cautioned that withholding such aid would endanger the instinct of sympathy, "the noblest part of our nature", and factors such as education could be more important. When Galton suggested that publishing research could encourage intermarriage within a "caste" of "those who are naturally gifted", Darwin foresaw practical difficulties, and thought it "the sole feasible, yet I fear ], plan of procedure in improving the human race", preferring to simply publicise the importance of inheritance and leave decisions to individuals.<ref>{{harvnb|Desmond|Moore|1991|pp=556–557, 572, 598}}<br />{{Harvnb|Darwin|1871|pp=, }}<br />{{cite web|url=http://www.galton.org/letters/darwin/correspondence.htm|title=Corespondence between Francis Galton and Charles Darwin|accessdate=2008-11-08}}</ref>
			Upon his return, he expressed a ] and questioned the basis for considering one religion more valid than another.<ref name="Darwin 1958-4" /> In the next few years, while intensively speculating on geology and the transmutation of species, he gave much thought to religion and openly discussed this with his wife Emma, whose beliefs similarly came from intensive study and questioning.<ref name="Darwin Correspondence Project" />
	Galton named the field of study '']'' in 1883, after Darwin's death, and developed ]. Eugenics movements were widespread at a time when Darwin's ] by ] ], and in some countries including the United States, ] laws were imposed. Following the use of ] it has been largely abandoned throughout the world.{{Ref_label|E|V|none}}
			The ] of Paley and ] vindicated evils such as starvation as a result of a benevolent creator's laws, which had an overall good effect. To Darwin, natural selection produced the good of adaptation but removed the need for design,<ref>{{harvnb|von Sydow|2005|pp=8–14}}</ref> and he could not see the work of an omnipotent deity in all the pain and suffering, such as the ] paralysing ]s as live food for its eggs.<ref name="Miles 2001" /> Though he thought of religion as a ] survival strategy, Darwin was reluctant to give up the idea of ]. He was increasingly troubled by the ].<ref>{{harvnb|von Sydow|2005|pp=4–5, 12–14}}</ref><ref>{{Harvnb|Moore|2006}}</ref>
	===Social Darwinism===
	{{details|Social Darwinism}}
	Taking descriptive ideas as moral and social justification creates the ethical ]. When ] argued that ] was ordained by God to get humans to ] and show restraint in getting families, this was used in the 1830s to justify ]s and ].<ref name=wm>{{harvnb|Wilkins|1997}}<br />{{Harvnb|Moore|2006}}</ref> Evolution was seen as having social implications, and ]'s 1851 book ''Social Statics'' based ideas of human freedom and individual liberties on his ] evolutionary theory.<ref>{{Harvnb|Sweet|2004}}</ref>
			Darwin remained close friends with the ] of Downe, ], and continued to play a leading part in the parish work of the church,<ref>{{cite web |url=https://www.darwinproject.ac.uk/commentary/religion/darwin-and-church |title=Darwin Correspondence Project&nbsp;– Darwin and the church: historical essay |access-date=26 November 2016 |url-status=live |archive-url=https://web.archive.org/web/20161128133709/https://www.darwinproject.ac.uk/commentary/religion/darwin-and-church |archive-date=28 November 2016 |date=5 June 2015 }}</ref> but from {{circa|1849}} would go for a walk on Sundays while his family attended church.<ref name="van Wyhe 2008b" /> He considered it "absurd to doubt that a man might be an ardent theist and an evolutionist"<ref name="Letter 12041 Archived 7 November 2009 at the Wayback Machine"> {{webarchive|url=https://web.archive.org/web/20091107174817/http://www.darwinproject.ac.uk/darwinletters/calendar/entry-12041.html |date=7 November 2009 }}&nbsp;– Darwin, C. R. to Fordyce, John, 7 May 1879</ref><ref name="Darwin"> {{Webarchive|url=https://web.archive.org/web/20170211082018/https://www.theguardian.com/commentisfree/belief/2009/sep/17/darwin-evolution-religion |date=11 February 2017 }} ] 17 September 2009</ref> and, though reticent about his religious views, in 1879 he wrote that "I have never been an atheist in the sense of denying the existence of a God.&nbsp;– I think that generally ... an agnostic would be the most correct description of my state of mind".<ref name="Darwin Correspondence Project"/><ref name="Letter 12041 Archived 7 November 2009 at the Wayback Machine"/>
	Darwin's theory of evolution was a matter of explanation. He thought it "absurd to talk of one animal being higher than another" and saw evolution as having no goal, but soon after the ''Origin'' was published in 1859, critics derided his description of a struggle for existence as a ] justification for the English industrial ] of the time. The term '']'' was used for the evolutionary ideas of others, including Spencer's "]" as free-market progress, and ]'s ] ideas of ]. Darwin did not share the racism common at that time: a point examined by the philosopher ], who is at pains to distance Darwin's attitudes from those later attributed to him.<ref>{{cite book|last=Flew|first=Antony|authorlink=Antony Flew|title=Darwinian Evolution|publisher=Transaction|location=Piscataway, NJ|year=1997|edition=2|isbn=1-56000-948-9|quote=...there seem to be absolutely no grounds for pillorying Darwin as a racist. On the contrary... he shared...principled hatred...for Negro slavery}}</ref> Darwin was strongly against slavery, against "ranking the so-called races of man as distinct species", and against ill-treatment of native people.<ref>{{harvnb|Wilkins|2008|pp=408–413}}</ref>{{Ref_label|F|VI|none}}
			The "]", published in 1915, claimed that Darwin had reverted to Christianity on his sickbed. The claims were repudiated by Darwin's children and have been dismissed as false by historians.<ref>{{harvnb|Moore|2005}}<br/>{{Harvnb|Yates|2003}}</ref>
	Darwin's views on social and political issues reflected his time and social position. He thought men's eminence over women was the outcome of sexual selection, a view disputed by ] in '']''.<ref name=Vandermassen>{{cite journal|author=Vandermassen, Griet|title=Sexual Selection: A Tale of Male Bias and Feminist Denial|url=http://ejw.sagepub.com/cgi/content/abstract/11/1/9|journal=European Journal of Women's Studies|year=2004|volume=11|issue=9|doi=10.1177/1350506804039812|accessdate=2009-11-24|pages=11–13}}</ref> He valued European civilisation and saw colonisation as spreading its benefits, with the sad but inevitable effect of extermination of savage peoples who did not become civilised. Darwin's theories presented this as natural, and were cited to promote policies which went against his humanitarian principles.<ref>{{cite web
	|first=Tony
	|last=Barta
	|title=Mr Darwin's shooters: on natural selection and the naturalizing of genocide
	|url= http://www.informaworld.com/smpp/section?content=a713721865&fulltext=713240928
	|work=Patterns of Prejudice, Volume 39, Issue 2
	|publisher=Routledge
	|pages=116–137
	|doi=10.1080/00313220500106170
	|date=2 June 2005
	|accessdate=2009-05-20
	}}</ref> Writers used ] to argue for various, often contradictory, ideologies such as laissez-faire dog-eat dog capitalism, racism, warfare, ] and ]. However, Darwin's holistic view of nature included "dependence of one being on another", thus ], ], ] social reformers and ] such as Prince ] stressed the value of co-operation over struggle within a species.<ref>{{Harvnb|Paul|2003|pp=223–225}}</ref> Darwin himself insisted that social policy should not simply be guided by concepts of struggle and selection in nature.<ref>{{Harvnb|Bannister|1989}}</ref>
			===Human society===
	The term "]" was used infrequently from around the 1890s, but became popular as a derogatory term in the 1940s when used by ] to attack the laissez-faire ] of those like ] who opposed reform and socialism. Since then it has been used as a term of abuse by those opposed to what they think are the moral consequences of evolution.<ref>{{Harvnb|Paul|2003}}<br />{{Harvnb|Kotzin|2004}}</ref><ref name=wm/>
			Darwin's views on social and political issues reflected his time and social position. He grew up in a family of ] reformers who, like his uncle Josiah Wedgwood, supported ] and the emancipation of slaves. Darwin was passionately opposed to slavery, while seeing no problem with the working conditions of English factory workers or servants.{{sfn|Browne|1995|pp=196–197}}
	==Commemoration==
	]
			Taking taxidermy lessons in 1826 from the freed slave John Edmonstone, whom Darwin long recalled as "a very pleasant and intelligent man", reinforced his belief that black people shared the same feelings, and could be as intelligent as people of other races. He took the same attitude to native people he met on the ''Beagle'' voyage.{{sfn|Browne|1995|pp=66, 198, 240}} Though commonplace in Britain at the time, ] and ] noticed the contrast with slave-owning America. Around twenty years later, racism became a feature of British society,<ref name="Darwin 1958-6"/><ref name="Silliman-1810">{{cite book | last=Silliman | first=B. | title=A Journal of Travels in England, Holland and Scotland: And of Two Passages Over the Atlantic, in the Years 1805 and 1806 ... | publisher=D. & G. Bruce | year=1810 | url=https://books.google.com/books?id=z4pCAAAAYAAJ&pg=PA216 | access-date=29 August 2022 | pages=216–217 | quote=As there are no slaves in England, perhaps the English have not learned to regard negroes as a degraded class of men, as we do in the United States | archive-date=18 December 2023 | archive-url=https://web.archive.org/web/20231218114110/https://books.google.com/books?id=z4pCAAAAYAAJ&pg=PA216#v=onepage&q&f=false | url-status=live }}<br/>{{cite book | last=Bachman | first=J. | title=The Doctrine of the Unity of the Human Race Examined on the Principles of Science | publisher=C. Canning | series=American culture series | year=1850 | isbn=978-0-608-43507-7 | url=https://books.google.com/books?id=qKYdfBlX-GMC&pg=PA105 | access-date=29 August 2022 | page=105 | archive-date=18 September 2023 | archive-url=https://web.archive.org/web/20230918120545/https://books.google.com/books?id=qKYdfBlX-GMC&pg=PA105 | url-status=live }}</ref> but Darwin remained strongly against slavery, against "ranking the so-called races of man as distinct species", and against ill-treatment of native people.<ref>{{harvnb|Wilkins|2008|pp=408–413}}</ref>{{Ref label|G|VII|none}}
	During Darwin's lifetime, many geographical features were given his name. An expanse of water adjoining the ] was named '']'' by ] after Darwin's prompt action, along with two or three of the men, saved them from being marooned on a nearby shore when a collapsing ] caused a large wave that would have swept away their boats,<ref>{{Harvnb|FitzRoy|1839|pp=}}</ref> and the nearby ] in the ] was named in celebration of Darwin's 25th birthday.<ref>{{harvnb|Leff|2000|loc=}}</ref> Another ] in ]'s ], between ] and ], was named in 1878 by Canada's then-chief geographer ] for Darwin.<ref>{{BCGNIS|29755}}</ref> When the '']'' was surveying Australia in 1839, Darwin's friend ] sighted a natural harbour which the ship's captain ] named '']''.<ref name=NTDoPaI>{{cite web|url=http://www.ipe.nt.gov.au/whatwedo/landinformation/place/origins/palmdarwin.html|archiveurl=http://web.archive.org/web/20060918153343/http://www.ipe.nt.gov.au/whatwedo/landinformation/place/origins/palmdarwin.html|archivedate=2006-09-18|title=Territory origins| accessdate=2006-12-15|publisher=Northern Territory Department of Planning and Infrastructure, Australia}}</ref> The settlement of ] ] was officially renamed ] in 1911. It became the capital city of Australia's ],<ref name=NTDoPaI /> which also boasts ]<ref>{{cite web|url=http://www.cdu.edu.au/|title=Charles Darwin University Homepage|accessdate=2006-12-15}}</ref> and ].<ref> Northern Territory, Australia Government. Retrieved on 2006-12-15.</ref> However, ], founded in 1964, was named in honour of the Darwin family, in part because they owned some of the site.<ref> Darwin College, Cambridge University website. Retrieved on 2006-12-10.</ref>
			Darwin's interaction with ] (Fuegians) such as Jemmy Button during the second voyage of HMS ''Beagle'' had a profound impact on his view of indigenous peoples. At his arrival in Tierra del Fuego he made a colourful description of "] savages".<ref name="Rozzi-2018" /> This view changed as he came to know Yaghan people more in detail. By studying the Yaghans, Darwin concluded that a number of basic emotions by different human groups were the same and that mental capabilities were roughly the same as for Europeans.<ref name="Rozzi-2018">{{cite journal |last1=Rozzi |first1=Ricardo|author-link=Ricardo Rozzi |date=2018 |title=Transformaciones del pensamiento de Darwin en cabo de hornos: Un legado para la ciencia y la etica ambiental|trans-title=Transformations of Darwin's thought in cape horn: A legacy for science and environmental ethics |language=es |journal=] |volume=46 |issue=1 |pages=267–277|doi=10.4067/S0718-22442018000100267 |doi-access=free }}</ref> While interested in Yaghan culture, Darwin failed to appreciate their deep ecological knowledge and elaborate cosmology until the 1850s when he inspected a dictionary of ] detailing 32,000 words.<ref name="Rozzi-2018" /> He saw that European colonisation would often lead to the extinction of native civilisations, and "tr to integrate colonialism into an evolutionary history of civilization analogous to natural history".<ref name="Barta-2005">{{cite journal |first=Tony|last=Barta|title=Mr Darwin's shooters: on natural selection and the naturalizing of genocide|journal=Patterns of Prejudice |volume=39|issue=2|pages=116–137 |doi=10.1080/00313220500106170 |date=2 June 2005 |s2cid=159807728}}</ref>
	The ] has commemorated Darwin's achievements by the award of the ] since 1908.
			] was that men's eminence over them was the outcome of sexual selection, a view disputed by ] in her 1875 book '']''.<ref name="Vandermassen, Griet-2004">{{cite journal |author=Vandermassen, Griet |title=Sexual Selection: A Tale of Male Bias and Feminist Denial |journal=European Journal of Women's Studies |year=2004 |volume=11 |issue=9 |doi=10.1177/1350506804039812 |pages=11–13 |citeseerx=10.1.1.550.3672 |s2cid=145221350 }}</ref>
	More than 120 ] and nine ] have been named after Darwin.<ref>{{cite web |url=http://www.darwinfacts.com/ |title=Charles Darwin 200 years - Things you didn't know about Charles Darwin |accessdate=2009-05-23}}</ref> In 2009, a remarkably complete fossil ] from 47 million years ago was announced as a significant ], and named '']'' to celebrate Darwin's bicentenary.<ref>{{cite news |url=http://news.bbc.co.uk/2/hi/science/nature/8057465.stm |title=Science & Environment; Scientists hail stunning fossil |author=Christine McGourty |date=19 May 2009 |publisher=BBC News |accessdate=2009-05-20}}</ref>
	Although related to American ] or ]s rather than ]es, the group of species related to those Darwin found in the ] became popularly known as "]" following publication of ]'s book of that name in 1947, fostering inaccurate legends about their significance to his work.<ref>{{Harvnb|Sulloway|1982|pp=45–47}}</ref>
			Darwin was intrigued by his ] ]'s argument, introduced in 1865, that ] of ] showed that moral and mental human traits could be inherited, and principles of animal breeding could apply to humans. In ''The Descent of Man'', Darwin noted that aiding the weak to survive and have families could lose the benefits of natural selection, but cautioned that withholding such aid would endanger the instinct of sympathy, "the noblest part of our nature", and factors such as education could be more important. When Galton suggested that publishing research could encourage intermarriage within a "caste" of "those who are naturally gifted", Darwin foresaw practical difficulties and thought it "the sole feasible, yet I fear ], plan of procedure in improving the human race", preferring to simply publicise the importance of inheritance and leave decisions to individuals.<ref>{{harvnb|Desmond|Moore|1991|pp=556–557, 572, 598}}<br />{{Harvnb|Darwin|1871|pp=, }}<br />{{cite web|url=http://www.galton.org/letters/darwin/correspondence.htm|title=Correspondence between Francis Galton and Charles Darwin|access-date=8 November 2008|url-status=live|archive-url=https://web.archive.org/web/20090102150434/http://galton.org/letters/darwin/correspondence.htm|archive-date=2 January 2009}}</ref> Francis Galton named this field of study "eugenics" in 1883,{{Ref label|H|VIII|1}} after Darwin's death, and his theories were cited to promote eugenic policies.<ref name="Barta-2005" />
	In 1992, Darwin was ranked #16 on ]'s ].<ref>{{Harvnb|Hart|2000|pp=82ff}}</ref> Darwin came fourth in the '']'' poll sponsored by the ] and voted for by the public.<ref>{{citation|url=http://news.bbc.co.uk/2/hi/entertainment/2341661.stm |title= Ten greatest Britons chosen |date= 20 October 2002 |publisher=BBC News |accessdate=2009-08-18}}</ref><ref> ]. Retrieved on 2006-12-15.</ref> In 2000 Darwin's image appeared on the ] ], replacing ]. His impressive, luxuriant beard (which was reportedly difficult to forge) was said to be a contributory factor to the bank's choice.<ref>"" '']'' (7 November 2000). Retrieved on 2006-12-15.</ref>
			==Evolutionary social movements==
	In the ], the ] based at the ] does research and conservation. To mark 2009 they are helping to reintroduce to ] (Charles Island) the specific ] which first alerted Darwin to species being unique to islands. It was eradicated from the main island by European species, mainly rats and goats, but survived on two small islands nearby.<ref name=Mockingbirdorigins>{{cite news|url=http://www.timesonline.co.uk/tol/news/environment/article5434999.ece|title=Mockingbird goes back to its origins in honour of Charles Darwin - Times Online|author=Lewis Smith|date=3 January 2009|accessdate=2009-02-13}}</ref>
			{{further|Eugenics|Social effects of evolutionary theory|Degeneration theory}}
			]
			].]]
			Darwin's fame and popularity led to his name being associated with ideas and movements that, at times, had only an indirect relation to his writings, and sometimes went directly against his express comments.
	As a humorous celebration of evolution, the annual ] is bestowed on individuals who "improve our ] by removing themselves from it."<ref> ''DarwinAwards.com''. Retrieved on 2007-12-11.</ref>
			Thomas Malthus had argued that population growth beyond resources was ordained by God to get humans to ] and show restraint in getting families; this was used in the 1830s to justify ]s and ].<ref name="Wilkins 1997Moore 2006">{{harvnb|Wilkins|1997}}<br />{{Harvnb|Moore|2006}}</ref> Evolution was by then seen as having social implications, and ]'s 1851 book ''Social Statics'' based ideas of human freedom and individual liberties on his Lamarckian evolutionary theory.<ref>{{Harvnb|Sweet|2004}}</ref>
	Numerous biographies of Darwin have been written, and the 1980 biographical novel '']'' by ] gives a closely researched fictional account of Darwin's life from the age of 22 onwards.
			Soon after the ''Origin'' was published in 1859, critics derided his description of a struggle for existence as a Malthusian justification for the English industrial capitalism of the time. The term ''Darwinism'' was used for the evolutionary ideas of others, including Spencer's "]" as free-market progress, and ]'s ] ideas of ]. Writers used natural selection to argue for various, often contradictory, ideologies such as laissez-faire dog-eat-dog capitalism, ] and ]. However, Darwin's holistic view of nature included "dependence of one being on another"; thus ], socialists, liberal social reformers and anarchists such as ] stressed the value of cooperation over struggle within a species.<ref>{{Harvnb|Paul|2003|pp=223–225}}</ref> Darwin himself insisted that social policy should not simply be guided by concepts of struggle and selection in nature.<ref>{{Harvnb|Bannister|1989}}</ref>
	===Darwin 2009 commemorations===
	] commemorating Darwin's birth and publication of '']''.]]
	] has become an annual celebration, and the bicentenary of Darwin's birth and the 150th anniversary of the publication of '']'' were celebrated by events and publications around the world.<ref>{{cite web|url=http://darwin-online.org.uk/2009.html|title=Darwin Online: Darwin 2009 commemorations around the world|publisher=Darwin Online|accessdate=2008-11-23}}</ref> The Darwin exhibition, after opening at the ] in ] in 2006, was shown at the ], the ] in ], the ] in ],<ref>{{cite web|url=http://www.amnh.org/exhibitions/darwin/curator/#|title=Darwin &#124; American Museum of Natural History|work=Meet the curator|accessdate=2008-11-28}}</ref> then from 14 November 2008 to 19 April 2009 in the ], ], as part of the ''Darwin200'' programme of events across the ].<ref>{{cite web|url=http://www.darwin200.org/|title=Darwin 200: Celebrating Charles Darwin's bicentenary|publisher=]|accessdate=2008-11-23}}</ref> It also appears at the ] in ] from 12 February to 3 May 2009.<ref>{{cite web |url=http://www.palazzoesposizioni.it/canale.asp?id=236 |title=Palazzo delle Esposizioni |work=Darwin 1809 - 2009, A cura di Niles Eldredge, Ian Tattersall e Telmo Pievani |accessdate=2009-02-22}}</ref> The ] featured a festival in July 2009.<ref>{{cite web|url=http://www.darwin2009.cam.ac.uk/|title=Darwin 2009 - The Festival|publisher=]|accessdate=2008-11-23}}</ref> His birthplace is celebrating with "Darwin's Shrewsbury 2009 Festival" events during the year.<ref>{{cite web|url=http://www.darwinshrewsbury.org/|title=Darwin's Shrewsbury 2009 Festival, Birthplace, History and Information|accessdate=2008-12-16}}</ref>
			After the 1880s, a eugenics movement developed on ideas of biological inheritance, and for scientific justification of their ideas appealed to some concepts of Darwinism. In Britain, most shared Darwin's cautious views on voluntary improvement and sought to encourage those with good traits in "positive eugenics". During the "Eclipse of Darwinism", a scientific foundation for eugenics was provided by ] ]. Negative eugenics to remove the "feebleminded" were popular in America, Canada and Australia, and ] introduced ] laws, followed by several other countries. Subsequently, ] brought the field into disrepute.{{Ref label|H|VIII|2}}
	In the United Kingdom a special commemorative issue of the ] shows a portrait of Darwin facing a ] surrounded by the inscription <small>1809 DARWIN 2009</small>, with the edge inscription <small>ON THE ORIGIN OF SPECIES 1859</small>. Collector versions of the coin have been released at a premium, and during the year the coins will be available from banks and post offices at face value.<ref name="nine">{{cite web| url = http://www.publications.parliament.uk/pa/cm200607/cmhansrd/cm070712/wmstext/70712m0001.htm| title = House of Commons Hansard Ministerial Statements for 12 July 2007|accessdate=2008-11-23}}</ref> To celebrate Darwin's life and achievements, the BBC has commissioned numerous television and radio programmes known collectively as the ].
			The term "]" was used infrequently from around the 1890s, but became popular as a derogatory term in the 1940s when used by ] to attack the ] conservatism of those like ] who opposed reform and socialism. Since then, it has been used as a term of abuse by those opposed to what they think are the moral consequences of evolution.<ref>{{Harvnb|Paul|2003}}<br />{{Harvnb|Kotzin|2004}}</ref><ref name="Wilkins 1997Moore 2006"/>
	In September 2008, the ] issued an article saying that the 200th anniversary of his birth was a fitting time to apologise to Darwin "for misunderstanding you and, by getting our first reaction wrong, encouraging others to misunderstand you still".<ref>'''' Rev Dr Malcolm Brown, Director of Mission and Public Affairs, Church of England. Retrieved 17 September 2008.</ref>
	] in London]]
	A dramatic motion picture entitled '']'' was released in 2009, joining a short list of film dramas about Darwin, including '']'', released in 1972.
	==Works==		==Works==
	{{details|List of works by Charles Darwin}}		{{further|Charles Darwin bibliography}}
	Darwin was a prolific writer. Even without publication of his works on evolution, he would have had a considerable reputation as the author of '']'', as a geologist who had published extensively on ] and had solved the puzzle of the formation of ]s, and as a biologist who had published the definitive work on ]s. While '']'' dominates perceptions of his work, '']'' and '']'' had considerable impact, and his books on plants including '']'' were innovative studies of great importance, as was his final work on '']''.<ref>{{Harvnb|Balfour|1882}}<br />{{Harvnb|van Wyhe|2008}}<br />{{Harvnb|Anonymous|1882}}</ref>		Darwin was a prolific writer. Even without the publication of his works on evolution, he would have had a considerable reputation as the author of ''The Voyage of the Beagle'', as a geologist who had published extensively on South America and had solved the puzzle of the formation of ]s, and as a biologist who had published the definitive work on barnacles. While ''On the Origin of Species'' dominates perceptions of his work, ''The Descent of Man'' and ''The Expression of the Emotions in Man and Animals'' had considerable impact, and his books on plants including ''The Power of Movement in Plants'' were innovative studies of great importance, as was his final work on '']''.<ref>{{Harvnb|Balfour|1882}}<br />{{Harvnb|van Wyhe|2008}}<br />{{Harvnb|Anonymous|1882}}</ref><ref>{{cite book|last=Brummitt|first=R. K.|author2=C. E. Powell|title=Authors of Plant Names |publisher=] |year=1992 |isbn=978-1-84246-085-6}}</ref>
			== Legacy and commemoration ==
	This botanist is denoted by the ] '''<span class="vcard"><span class="fn n"><span class="nickname">Darwin</span></span></span>''' when ] a ].<ref>{{cite book|last = Brummitt|first = R. K.|coauthors = C. E. Powell|title = Authors of Plant Names |publisher = ] |year = 1992 |isbn = 1-84246-085-4}}</ref>
			{{further|List of things named after Charles Darwin|List of taxa described by Charles Darwin|Commemoration of Charles Darwin}}
			{{Anchor|Legacy}}
			] building where he had studied]]
			As Alfred Russel Wallace put it, Darwin had "wrought a greater revolution in human thought within a quarter of a century than any man of our time{{snd}}or perhaps any time", having "given us a new conception of the world of life, and a theory which is itself a powerful instrument of research; has shown us how to combine into one consistent whole the facts accumulated by all the separate classes of workers, and has thereby revolutionised the whole study of nature".<ref name="Van Wyhe-2021">{{cite book | last=Van Wyhe | first=J. | title=Charles Darwin: The Man, His Great Voyage, and His Theory of Evolution | publisher=Rosen Publishing Group, Incorporated | series=Pioneers of Science | year=2021 | isbn=978-1-4994-7110-6 | url=https://books.google.com/books?id=JnZeEAAAQBAJ&pg=PA155 | access-date=23 May 2022 | pages=154–155 | archive-date=18 December 2023 | archive-url=https://web.archive.org/web/20231218114110/https://books.google.com/books?id=JnZeEAAAQBAJ&pg=PA155#v=onepage&q&f=false | url-status=live }}</ref> The paleoanthropologist ] states that "Darwin is rightly considered to be the preeminent evolutionary scientist of all time".{{sfn|Holliday|2021|p=5}}
			By around 1880, most scientists were convinced of evolution as descent with modification, though few agreed with Darwin that natural selection "has been the main but not the exclusive means of modification".<ref name="van Wyhe 2008Darwin 1872">{{Harvnb|van Wyhe|2008}}<br />{{harvnb|Darwin|1872|p=}}.</ref> During "]" scientists explored alternative mechanisms. Then ] incorporated ] in '']'',<ref>{{cite journal |url=http://www.genetics.org/content/154/4/1419.full |journal=Genetics |title=The Genetical Theory of Natural Selection |first=A. W. F. |last=Edwards |author-link=A. W. F. Edwards |date=1 April 2000 |volume=154 |issue=4 |pages=1419–1426 |doi=10.1093/genetics/154.4.1419 |pmid=10747041 |pmc=1461012 |url-status=live |archive-url=https://web.archive.org/web/20150924041631/http://www.genetics.org/content/154/4/1419.full |archive-date=24 September 2015 }}</ref> leading to ] and the ], which continues to develop.<ref name="Bowler 2003" /> Scientific discoveries have confirmed and validated Darwin's key insights.<ref name="Van Wyhe-2021" />
			Geographical features given his name include ]<ref>{{Harvnb|FitzRoy|1839|pp=}}.</ref> and ],<ref>{{harvnb|Leff|2000|loc=}}</ref> both named while he was on ], and ], named by his former shipmates on ], which eventually became the location of ], the capital city of Australia's ].<ref name="Northern Territory Department of Planning and Infrastructure, Australia">{{cite web|url=http://www.ipe.nt.gov.au/whatwedo/landinformation/place/origins/palmdarwin.html|archive-url=https://web.archive.org/web/20060918153343/http://www.ipe.nt.gov.au/whatwedo/landinformation/place/origins/palmdarwin.html|archive-date=18 September 2006|title=Territory origins| access-date=15 December 2006|publisher=Northern Territory Department of Planning and Infrastructure, Australia}}</ref> Darwin's name was given, ] or ], to numerous plants and animals, including many he had collected on the voyage.<ref>{{Cite book |last=Heard, Stephen B. |title=Charles Darwin's barnacle and David Bowie's spider : how scientific names celebrate adventurers, heroes, and even a few scoundrels |others=Damstra, Emily S. |date=2020 |isbn=978-0-300-25269-9 |publisher=Yale University Press |location=New Haven |oclc=1143645266}}</ref><ref>{{cite web |url=http://www.darwinfacts.com/ |title=Charles Darwin 200 years&nbsp;– Things you didn't know about Charles Darwin |access-date=23 May 2009 |url-status=live |archive-url=https://web.archive.org/web/20090528033253/http://darwinfacts.com/ |archive-date=28 May 2009 }}</ref> The Linnean Society of London began awards of the ] in 1908, to mark fifty years from the joint reading on 1 July 1858 of papers by Darwin and Wallace publishing their theory. Further awards were made in 1958 and 2008; since 2010, the awards have been annual.<ref name="The Linnean Society-2016">{{cite web |date=1 February 2016 |title=The Darwin-Wallace Medal |url=https://www.linnean.org/the-society/medals-awards-prizes-grants/the-darwin-wallace-medal |access-date=22 April 2022 |website=The Linnean Society |archive-date=29 March 2019 |archive-url=https://web.archive.org/web/20190329223140/https://www.linnean.org/the-society/medals-awards-prizes-grants/the-darwin-wallace-medal |url-status=live }}</ref> ], a postgraduate college at ] founded in 1964, is named after the Darwin family.<ref>{{cite web |title=Darwin College – Maps and directions – University of Kent |url=https://www.kent.ac.uk/maps/canterbury/canterbury-campus/building/darwin-college |url-status=live |archive-url=https://web.archive.org/web/20161031024930/https://www.kent.ac.uk/maps/canterbury/canterbury-campus/building/darwin-college |archive-date=31 October 2016 |access-date=30 October 2016 |website=www.kent.ac.uk}}</ref> From 2000 to 2017, UK £10 banknotes issued by the ] featured Darwin's portrait printed on the reverse,<ref name="BBC News-2000">{{cite web |date=7 November 2000 |title=How to join the noteworthy |url=http://news.bbc.co.uk/1/hi/uk/1009901.stm |access-date=24 April 2022 |website=BBC News |archive-date=30 June 2006 |archive-url=https://web.archive.org/web/20060630102007/http://news.bbc.co.uk/1/hi/uk/1009901.stm |url-status=live }}</ref><ref name="CBBC Newsround-2013">{{cite web |date=24 July 2013 |title=Author Jane Austen to feature on new £10 note |url=https://www.bbc.co.uk/newsround/23434198 |access-date=24 April 2022 |website=CBBC Newsround |archive-date=24 April 2022 |archive-url=https://web.archive.org/web/20220424113045/https://www.bbc.co.uk/newsround/23434198 |url-status=live }}</ref> along with a ] and ].<ref name="bankofengland.co.uk-2005">{{cite web |date=25 May 2005 |title=Bank of England – Banknotes – Current Banknotes – £10 |url=http://www.bankofengland.co.uk/banknotes/current/current_10.htm |url-status=dead |archive-url=https://web.archive.org/web/20050525230931/http://www.bankofengland.co.uk/banknotes/current/current_10.htm |archive-date=25 May 2005 |access-date=24 April 2022 |website=bankofengland.co.uk}}</ref>
	==See also==		==See also==
			{{Portal|Biology}}
	<!-- Please avoid repeating links above -->		<!-- Please avoid repeating links above -->
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	*]
	*"]"		* '']''
			* '']'' (biographical drama film)
	*]
			* ]
	*]
			* ]
	*]
			* ]
	*]
			* ]
	*]
	*]		* ]
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	==Notes==		==Notes==
			{{refbegin}}
	<div class="references-small">
			'''{{small|I}}.''' {{Note label|B|II|none}} ] was to become known after the voyage for ], but at this time he had considerable interest in Lyell's ideas, and they met before the voyage when Lyell asked for observations to be made in South America. FitzRoy's diary during the ascent of the River Santa Cruz in ] recorded his opinion that the plains were ]es, but on return, newly married to a very religious lady, he recanted these ideas.{{Harv|Browne|1995|pp=186, 414}}
	'''<small>I</small>.''' {{Note_label|A|I|none}} Darwin was eminent as a ], ], ], and ]; after working as a physician's assistant and two years as a ] was educated as a ]; and was trained in ].<ref name=ODNB>{{Harvnb|Desmond|Moore|Browne|2004|}}</ref>
			'''{{small|II}}.''' {{Note label|C|III|none}} In the section ] of Chapter XIII of ''On the Origin of Species'', Darwin commented on ] bone patterns between humans and other mammals, writing: "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?"<ref>{{harvnb|Darwin|1859|p=}}</ref> and in the concluding chapter: "The framework of bones being the same in the hand of a man, wing of a bat, fin of the porpoise, and leg of the horse … at once explain themselves on the theory of descent with slow and slight successive modifications."<ref>{{harvnb|Darwin|1859|p=}}</ref>
	'''<small>II</small>.''' {{Note_label|B|II|none}} ] was to become known after the voyage for biblical literalism, but at this time he had considerable interest in Lyell's ideas, and they met before the voyage when Lyell asked for observations to be made in South America. FitzRoy's diary during the ascent of the River Santa Cruz in ] recorded his opinion that the plains were ]es, but on return, newly married to a very religious lady, he recanted these ideas. {{Harv|Browne|1995|pp= 186, 414}}
			'''{{small|III}}.''' {{Note label|D|IV|1}}{{Note label|D|IV|2}}{{Note label|D|IV|3}}
	'''<small>III</small>.''' {{Note_label|C|III|none}} See, for example, WILLA volume 4, '''' by Deborah M. De Simone: "Gilman shared many basic educational ideas with the generation of thinkers who matured during the period of "intellectual chaos" caused by Darwin's Origin of the Species. Marked by the belief that individuals can direct human and social evolution, many progressives came to view education as the panacea for advancing social progress and for solving such problems as urbanisation, poverty, or immigration."
			In '']'' Darwin mentioned ] in his concluding remark that "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."<ref name="Darwin 1859"/>
			In "Chapter VI: Difficulties on Theory" he referred to ]: "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."<ref name="Darwin 1859-2"/>
	'''<small>IV</small>.''' {{Note_label|D|IV|none}} See, for example, the song "A lady fair of lineage high" from ]'s '']'', which describes the descent of man (but not woman!) from apes.
			In '']'' of 1871, Darwin discussed the first passage:
	'''<small>V</small>.''' {{Note_label|E|V|none}} ] studied human heredity as ], while ] movements sought to manage society, with a focus on ] in the United Kingdom, and on disability and ethnicity in the United States, leading to geneticists seeing this as impractical ]. A shift from voluntary arrangements to "negative" eugenics included ] laws in the United States, copied by ] as the basis for ] based on virulent racism and "]".<br />({{Citation
			"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."<ref>{{Harvnb|Darwin|1871|p=}}</ref> In a preface to the 1874 second edition, he added a reference to the second point: "it has been said by several critics, that when I found that many details of structure in man could not be explained through natural selection, I invented sexual selection; I gave, however, a tolerably clear sketch of this principle in the first edition of the 'Origin of Species,' and I there stated that it was applicable to man."<ref>{{Harvnb|Darwin|1874|p=}}</ref>
	| last = Thurtle
	| first =Phillip
	| publication-date =
	| date =Updated 17 December 1996
	| title =the creation of genetic identity
	| periodical =SEHR
	| volume = 5
	| issue =Supplement: Cultural and Technological Incubations of Fascism
	| url =http://www.stanford.edu/group/SHR/5-supp/text/thurtle.html
	| accessdate =2008-11-11}}<br />{{Citation
	| last = Edwards
	| first =A. W. F.
	| date = April 1, 2000 | title =The Genetical Theory of Natural Selection
	| periodical = Genetics
	| volume = 154
	| issue =April 2000
	| pages = 1419–1426
	| url =http://www.genetics.org/cgi/content/full/154/4/1419#The_Eclipse_of_Darwinism
	| accessdate =2008-11-11
	| pmid = 10747041
	}}<br />{{cite web | last = Wilkins | first = John | title = Evolving Thoughts: Darwin and the Holocaust 3: eugenics | url = http://scienceblogs.com/evolvingthoughts/2006/09/darwin_and_the_holocaust_3_eug_1.php | accessdate =2008-11-11}})
			'''{{small|IV}}.''' {{Note label|E|V|none}} See, for example, WILLA volume 4, '''' by Deborah M. De Simone: "Gilman shared many basic educational ideas with the generation of thinkers who matured during the period of "intellectual chaos" caused by Darwin's Origin of the Species. Marked by the belief that individuals can direct human and social evolution, many progressives came to view education as the panacea for advancing social progress and for solving such problems as urbanisation, poverty, or immigration."
	'''<small>VI</small>.''' {{Note_label|F|VI|none}} Darwin did not share the then common view that other races are inferior, and said of his ] tutor ], a freed black slave, "I used often to sit with him, for he was a very pleasant and intelligent man".<ref name=eddy/>
			'''{{small|V}}.''' {{Note label|F|VI|none}} See, for example, the song "A lady fair of lineage high" from ]'s '']'', which describes the descent of man (but not woman!) from apes.
	Early in the ''Beagle'' voyage he nearly lost his position on the ship when he criticised FitzRoy's defence and praise of slavery. {{Harv|Darwin|1958|p=}} He wrote home about "how steadily the general feeling, as shown at elections, has been rising against Slavery. What a proud thing for England if she is the first European nation which utterly abolishes it! I was told before leaving England that after living in slave countries all my opinions would be altered; the only alteration I am aware of is forming a much higher estimate of the negro character." {{harv|Darwin|1887|p=}} Regarding ], he "could not have believed how wide was the difference between savage and civilized man: it is greater than between a wild and domesticated animal, inasmuch as in man there is a greater power of improvement", but he knew and liked civilised Fuegians like ]: "It seems yet wonderful to me, when I think over all his many good qualities, that he should have been of the same race, and doubtless partaken of the same character, with the miserable, degraded savages whom we first met here."{{Harv|Darwin|1845|pp= }}
			'''{{small|VI}}.''' {{Note label|G|VII|none}} Darwin's belief that black people had the same essential humanity as Europeans, and had many mental similarities, was reinforced by the lessons he had from ] in 1826.<ref name="Darwin 1958-6" /> Early in the ''Beagle'' voyage, Darwin nearly lost his position on the ship when he criticised FitzRoy's defence and praise of slavery. {{Harv|Darwin|1958|p=}} He wrote home about "how steadily the general feeling, as shown at elections, has been rising against Slavery. What a proud thing for England if she is the first European nation which utterly abolishes it! I was told before leaving England that after living in slave countries all my opinions would be altered; the only alteration I am aware of is forming a much higher estimate of the negro character." {{harv|Darwin|1887|p=}} Regarding ], he "could not have believed how wide was the difference between savage and civilized man: it is greater than between a wild and domesticated animal, inasmuch as in man there is a greater power of improvement", but he knew and liked civilised Fuegians like ]: "It seems yet wonderful to me, when I think over all his many good qualities, that he should have been of the same race, and doubtless partaken of the same character, with the miserable, degraded savages whom we first met here." {{Harv|Darwin|1845|pp=}}
	In the '']'' he mentioned the Fuegians and Edmonstone when arguing against "ranking the so-called races of man as distinct species".<ref>{{Harvnb|Darwin|1871|pp=, .}}</ref>
	He rejected the ill-treatment of native people, and for example wrote of massacres of ]n men, women, and children, "Every one here is fully convinced that this is the most just war, because it is against barbarians. Who would believe in this age that such atrocities could be committed in a Christian civilized country?" {{harv|Darwin|1845|p-}}		In the ''Descent of Man'', he mentioned the similarity of Fuegians' and Edmonstone's minds to Europeans' when arguing against "ranking the so-called races of man as distinct species".<ref>{{Harvnb|Darwin|1871|pp=, }}</ref>
			He rejected the ill-treatment of native people, and for example wrote of massacres of ]n men, women, and children, "Every one here is fully convinced that this is the most just war, because it is against barbarians. Who would believe in this age that such atrocities could be committed in a Christian civilized country?"{{harv|Darwin|1845|p=}}
	</div>
			'''{{small|VII}}.''' {{Note label|H|VIII|1}}{{Note label|H|VIII|2}} ]s studied human heredity as ], while ] movements sought to manage society, with a focus on social class in the United Kingdom, and on disability and ethnicity in the United States, leading to geneticists seeing this movement as impractical ]. A shift from voluntary arrangements to "negative" eugenics included ] laws in the United States, copied by ] as the basis for ] based on virulent racism and "]".<br />({{cite news | url=http://www.stanford.edu/group/SHR/5-supp/text/thurtle.html | title=the creation of genetic identity | last=Thurtle | first=Phillip | date=17 December 1996 | issue=Supplement: Cultural and Technological Incubations of Fascism | volume=5 | periodical=SEHR | access-date=11 November 2008 |ref=none}} {{Cite news |url=http://www.genetics.org/cgi/content/full/154/4/1419#The_Eclipse_of_Darwinism| title=The Genetical Theory of Natural Selection | last=Edwards | first=A. W. F. |author-link=A. W. F. Edwards | date=1 April 2000 | issue=April 2000 | volume=154 | pages=1419–1426 | pmc=1461012 | pmid=10747041 | periodical=Genetics | access-date=11 November 2008 | ref=none}}{{cite web |url=http://scienceblogs.com/evolvingthoughts/2006/09/darwin_and_the_holocaust_3_eug_1.php |title=Evolving Thoughts: Darwin and the Holocaust 3: eugenics |last=Wilkins |first=John |access-date=11 November 2008 |url-status=dead |archive-url=https://web.archive.org/web/20081205154013/http://scienceblogs.com/evolvingthoughts/2006/09/darwin_and_the_holocaust_3_eug_1.php |archive-date=5 December 2008 |ref=none}})
			'''{{small|VIII}}.''' {{Note label|I|IX|none}} ] writes of his "theory that turned to these arcane botanical studies – producing more than one book that was solidly empirical, discreetly evolutionary, yet a 'horrid bore' – at least partly so that the clamorous controversialists, fighting about apes and angels and souls, would leave him... alone". ], "The Brilliant Plodder" (review of Ken Thompson, ''Darwin's Most Wonderful Plants: A Tour of His Botanical Legacy'', ], 255 pp.; Elizabeth Hennessy, ''On the Backs of Tortoises: Darwin, the Galápagos, and the Fate of an Evolutionary Eden'', ], 310 pp.; Bill Jenkins, ''Evolution Before Darwin: Theories of the Transmutation of Species in Edinburgh, 1804–1834'', ], 222 pp.), '']'', vol. LXVII, no. 7 (23 April 2020), pp.&nbsp;22–24. Quammen, quoted from p.&nbsp;24 of his review.
			{{refend}}
	==Citations==		==Citations==
	{{reflist|colwidth=30em}}		{{reflist|30em}}
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			| issue=7 (24 April)
			| pages=1–164
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			| doi-access=free
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			| pages=159–192
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	| first = Momme
	| year = 2005
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	| editor-first =David M.
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	| pages =141–156
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	*{{Citation
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	{{refend}}		{{refend}}
	==External links==		==External links==
			{{Library resources box
	{{sisterlinks|Charles Darwin}}
			| onlinebooks = yes
	*] – ; Darwin's publications, private papers and bibliography, supplementary works including biographies, obituaries and reviews.
			| by = yes
	*{{gutenberg author| id=Charles+Darwin| name=Charles Darwin}}; public domain
			| viaf = 27063124
	* Full text and notes for complete correspondence to 1867, with summaries of all the rest
			| label = Charles Darwin
	*
			}}
	*{{dmoz|Science/Biology/History/People/Darwin,_Charles/}}
			* {{cite web |title=The Complete Work of Charles Darwin Online |url=https://darwin-online.org.uk/ |access-date=4 March 2024}}
	*{{worldcat id|id=lccn-n78-95637}}
			* {{StandardEbooks|Standard Ebooks URL=https://standardebooks.org/ebooks/charles-darwin}}
	*{{NRA|P7461}}
			* {{Gutenberg author |id=485| name=Charles Darwin}}
	*, ]
			* {{Internet Archive author |name=Charles Robert Darwin}}
	*
			* {{Librivox author |id=166}}
	*
			* ]&nbsp;– ; Darwin's publications, private papers and bibliography, supplementary works including biographies, obituaries and reviews
	*
			* Full text and notes for complete correspondence to 1867, with summaries of all the rest, and pages of commentary
	*{{Wikisource1911Enc Citation|Darwin, Charles Robert}}
			*
	* (3 min 20 sec).
			* {{UK National Archives ID}}
	*
			* View books owned and annotated by at the online Biodiversity Heritage Library.
	* - a short video discussing Darwin and Agassiz' coral reef formation debate
			* {{BHL author}}
	* - A 3 part drama-documentary exploring Charles Darwin and the significant contributions of his colleagues Joseph Hooker, Thomas Huxley and Alfred Russel Wallace also featuring interviews with ], David Suzuki, Jared Diamond and Iain McCalman.
			* in ]
	* Account of the ''Beagle'' voyage using animation, in English from ], Paris.
	{{Darwin}}		* {{NPG name}}
			* {{PM20|FID=pe/003703}}
	{{evolution}}
			* – Occasional Papers from RHS Lindley Library, volume 3 July 2010
			* ], 29 April 1882, pp.&nbsp;256,
			* {{IEP|/darwin/}}
			{{Darwin}}
	<!-- Metadata: see ] -->
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			{{Ethology}}
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	|PLACE OF BIRTH= ], ], ], England
			{{Evolutionary psychologists}}
	|DATE OF DEATH= 1882-04-19
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	|PLACE OF DEATH= ], ], England
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			{{Subject bar
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	{{DEFAULTSORT:Darwin, Charles}}		{{DEFAULTSORT:Darwin, Charles}}
	]		]
	]		]
	]		]
	]		]
	]		]
			]
			]
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English naturalist and biologist (1809–1882) For other people named Charles Darwin, see Charles Darwin (disambiguation).

Charles DarwinJP FRS FRGS FLS FZS
Darwin, c. 1854, when he was preparing On the Origin of Species
Born	Charles Robert Darwin (1809-02-12)12 February 1809 Shrewsbury, Shropshire, England
Died	19 April 1882(1882-04-19) (aged 73) Down House, Down, Kent, England
Resting place	Westminster Abbey
Education	University of Edinburgh Christ's College, Cambridge (BA, MA)
Known for	Natural selection
Spouse	Emma Wedgwood ​(m. 1839)​
Children	10, including William, Henrietta, George, Francis, Leonard and Horace
Parents	Robert Darwin Susannah Wedgwood
Family	Darwin–Wedgwood
Awards	FRS (1839) Royal Medal (1853) Wollaston Medal (1859) Copley Medal (1864) Pour le Mérite (1867) Baly Medal (1879)
Writing career
Notable works	The Voyage of the Beagle On the Origin of Species The Descent of Man
Scientific career
Fields	Natural history Geology
Institutions	Geological Society of London
Academic advisors	John Stevens Henslow Adam Sedgwick
Author abbrev. (botany)	Darwin
Author abbrev. (zoology)	Darwin
Signature

Charles Robert Darwin (/ˈdɑːrwɪn/ DAR-win; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended from a common ancestor is now generally accepted and considered a fundamental scientific concept. In a joint presentation with Alfred Russel Wallace, he introduced his scientific theory that this branching pattern of evolution resulted from a process he called natural selection, in which the struggle for existence has a similar effect to the artificial selection involved in selective breeding. Darwin has been described as one of the most influential figures in human history and was honoured by burial in Westminster Abbey.

Darwin's early interest in nature led him to neglect his medical education at the University of Edinburgh; instead, he helped to investigate marine invertebrates. His studies at the University of Cambridge's Christ's College from 1828 to 1831 encouraged his passion for natural science. However, it was his five-year voyage on HMS Beagle from 1831 to 1836 that truly established Darwin as an eminent geologist. The observations and theories he developed during his voyage supported Charles Lyell's concept of gradual geological change. Publication of his journal of the voyage made Darwin famous as a popular author.

Puzzled by the geographical distribution of wildlife and fossils he collected on the voyage, Darwin began detailed investigations and, in 1838, devised his theory of natural selection. Although he discussed his ideas with several naturalists, he needed time for extensive research, and his geological work had priority. He was writing up his theory in 1858 when Alfred Russel Wallace sent him an essay that described the same idea, prompting the immediate joint submission of both their theories to the Linnean Society of London. Darwin's work established evolutionary descent with modification as the dominant scientific explanation of natural diversification. In 1871, he examined human evolution and sexual selection in The Descent of Man, and Selection in Relation to Sex, followed by The Expression of the Emotions in Man and Animals (1872). His research on plants was published in a series of books, and in his final book, The Formation of Vegetable Mould, through the Actions of Worms (1881), he examined earthworms and their effect on soil.

Darwin published his theory of evolution with compelling evidence in his 1859 book On the Origin of Species. By the 1870s, the scientific community and a majority of the educated public had accepted evolution as a fact. However, many initially favoured competing explanations that gave only a minor role to natural selection, and it was not until the emergence of the modern evolutionary synthesis from the 1930s to the 1950s that a broad consensus developed in which natural selection was the basic mechanism of evolution. Darwin's scientific discovery is the unifying theory of the life sciences, explaining the diversity of life.

Biography

Early life and education

Further information: Charles Darwin's education and Darwin–Wedgwood family

Darwin was born in Shrewsbury, Shropshire, on 12 February 1809, at his family's home, The Mount. He was the fifth of six children of wealthy society doctor and financier Robert Darwin and Susannah Darwin (née Wedgwood). His grandfathers Erasmus Darwin and Josiah Wedgwood were both prominent abolitionists. Erasmus Darwin had praised general concepts of evolution and common descent in his Zoonomia (1794), a poetic fantasy of gradual creation including undeveloped ideas anticipating concepts his grandson expanded.

Both families were largely Unitarian, though the Wedgwoods were adopting Anglicanism. Robert Darwin, a freethinker, had baby Charles baptised in November 1809 in the Anglican St Chad's Church, Shrewsbury, but Charles and his siblings attended the local Unitarian Church with their mother. The eight-year-old Charles already had a taste for natural history and collecting when he joined the day school run by its preacher in 1817. That July, his mother died. From September 1818, he joined his older brother Erasmus in attending the nearby Anglican Shrewsbury School as a boarder.

Darwin spent the summer of 1825 as an apprentice doctor, helping his father treat the poor of Shropshire, before going to the well-regarded University of Edinburgh Medical School with his brother Erasmus in October 1825. Darwin found lectures dull and surgery distressing, so he neglected his studies. He learned taxidermy in around 40 daily hour-long sessions from John Edmonstone, a freed black slave who had accompanied Charles Waterton in the South American rainforest.

In Darwin's second year at the university, he joined the Plinian Society, a student natural-history group featuring lively debates in which radical democratic students with materialistic views challenged orthodox religious concepts of science. He assisted Robert Edmond Grant's investigations of the anatomy and life cycle of marine invertebrates in the Firth of Forth, and on 27 March 1827 presented at the Plinian his own discovery that black spores found in oyster shells were the eggs of a skate leech. One day, Grant praised Lamarck's evolutionary ideas. Darwin was astonished by Grant's audacity, but had recently read similar ideas in his grandfather Erasmus' journals. Darwin was rather bored by Robert Jameson's natural-history course, which covered geology – including the debate between neptunism and plutonism. He learned the classification of plants and assisted with work on the collections of the University Museum, one of the largest museums in Europe at the time.

Darwin's neglect of medical studies annoyed his father, who sent him to Christ's College, Cambridge, in January 1828, to study for a Bachelor of Arts degree as the first step towards becoming an Anglican country parson. Darwin was unqualified for Cambridge's Tripos exams and was required instead to join the ordinary degree course. He preferred riding and shooting to studying.

During the first few months of Darwin's enrolment at Christ's College, his second cousin William Darwin Fox was still studying there. Fox impressed him with his butterfly collection, introducing Darwin to entomology and influencing him to pursue beetle collecting. He did this zealously and had some of his finds published in James Francis Stephens' Illustrations of British entomology (1829–1932).

Through Fox, Darwin became a close friend and follower of botany professor John Stevens Henslow. He met other leading parson-naturalists who saw scientific work as religious natural theology, becoming known to these dons as "the man who walks with Henslow". When his own exams drew near, Darwin applied himself to his studies and was delighted by the language and logic of William Paley's Evidences of Christianity (1795). In his final examination in January 1831, Darwin did well, coming tenth out of 178 candidates for the ordinary degree.

Darwin had to stay at Cambridge until June 1831. He studied Paley's Natural Theology or Evidences of the Existence and Attributes of the Deity (first published in 1802), which made an argument for divine design in nature, explaining adaptation as God acting through laws of nature. He read John Herschel's new book, Preliminary Discourse on the Study of Natural Philosophy (1831), which described the highest aim of natural philosophy as understanding such laws through inductive reasoning based on observation, and Alexander von Humboldt's Personal Narrative of scientific travels in 1799–1804. Inspired with "a burning zeal" to contribute, Darwin planned to visit Tenerife with some classmates after graduation to study natural history in the tropics. In preparation, he joined Adam Sedgwick's geology course, then on 4 August travelled with him to spend a fortnight mapping strata in Wales.

Survey voyage on HMS Beagle

Further information: Second voyage of HMS Beagle

After leaving Sedgwick in Wales, Darwin spent a few days with student friends at Barmouth. He returned home on 29 August to find a letter from Henslow proposing him as a suitable (if unfinished) naturalist for a self-funded supernumerary place on HMS Beagle with captain Robert FitzRoy, a position for a gentleman rather than "a mere collector". The ship was to leave in four weeks on an expedition to chart the coastline of South America. Robert Darwin objected to his son's planned two-year voyage, regarding it as a waste of time, but was persuaded by his brother-in-law, Josiah Wedgwood II, to agree to (and fund) his son's participation. Darwin took care to remain in a private capacity to retain control over his collection, intending it for a major scientific institution.

After delays, the voyage began on 27 December 1831; it lasted almost five years. As FitzRoy had intended, Darwin spent most of that time on land investigating geology and making natural history collections, while HMS Beagle surveyed and charted coasts. He kept careful notes of his observations and theoretical speculations. At intervals during the voyage, his specimens were sent to Cambridge together with letters including a copy of his journal for his family. He had some expertise in geology, beetle collecting and dissecting marine invertebrates, but in all other areas, was a novice and ably collected specimens for expert appraisal. Despite suffering badly from seasickness, Darwin wrote copious notes while on board the ship. Most of his zoology notes are about marine invertebrates, starting with plankton collected during a calm spell.

On their first stop ashore at St Jago in Cape Verde, Darwin found that a white band high in the volcanic rock cliffs included seashells. FitzRoy had given him the first volume of Charles Lyell's Principles of Geology, which set out uniformitarian concepts of land slowly rising or falling over immense periods, and Darwin saw things Lyell's way, theorising and thinking of writing a book on geology. When they reached Brazil, Darwin was delighted by the tropical forest, but detested the sight of slavery there, and disputed this issue with FitzRoy.

The survey continued to the south in Patagonia. They stopped at Bahía Blanca, and in cliffs near Punta Alta Darwin made a major find of fossil bones of huge extinct mammals beside modern seashells, indicating recent extinction with no signs of change in climate or catastrophe. He found bony plates like a giant version of the armour on local armadillos. From a jaw and tooth he identified the gigantic Megatherium, then from Cuvier's description thought the armour was from this animal. The finds were shipped to England, and scientists found the fossils of great interest. In Patagonia, Darwin came to wrongly believe the territory was devoid of reptiles.

On rides with gauchos into the interior to explore geology and collect more fossils, Darwin gained social, political and anthropological insights into both native and colonial people at a time of revolution, and learnt that two types of rhea had separate but overlapping territories. Further south, he saw stepped plains of shingle and seashells as raised beaches at a series of elevations. He read Lyell's second volume and accepted its view of "centres of creation" of species, but his discoveries and theorising challenged Lyell's ideas of smooth continuity and of extinction of species.

Three Fuegians on board, who had been seized during the first Beagle voyage then given Christian education in England, were returning with a missionary. Darwin found them friendly and civilised, yet at Tierra del Fuego he met "miserable, degraded savages", as different as wild from domesticated animals. He remained convinced that, despite this diversity, all humans were interrelated with a shared origin and potential for improvement towards civilisation. Unlike his scientist friends, he now thought there was no unbridgeable gap between humans and animals. A year on, the mission had been abandoned. The Fuegian they had named Jemmy Button lived like the other natives, had a wife, and had no wish to return to England.

Darwin experienced an earthquake in Chile in 1835 and saw signs that the land had just been raised, including mussel-beds stranded above high tide. High in the Andes he saw seashells and several fossil trees that had grown on a sand beach. He theorised that as the land rose, oceanic islands sank, and coral reefs round them grew to form atolls.

On the geologically new Galápagos Islands, Darwin looked for evidence attaching wildlife to an older "centre of creation", and found mockingbirds allied to those in Chile but differing from island to island. He heard that slight variations in the shape of tortoise shells showed which island they came from, but failed to collect them, even after eating tortoises taken on board as food. In Australia, the marsupial rat-kangaroo and the platypus seemed so unusual that Darwin thought it was almost as though two distinct Creators had been at work. He found the Aborigines "good-humoured & pleasant", their numbers depleted by European settlement.

FitzRoy investigated how the atolls of the Cocos (Keeling) Islands had formed, and the survey supported Darwin's theorising. FitzRoy began writing the official Narrative of the Beagle voyages, and after reading Darwin's diary, he proposed incorporating it into the account. Darwin's Journal was eventually rewritten as a separate third volume, on geology and natural history.

In Cape Town, South Africa, Darwin and FitzRoy met John Herschel, who had recently written to Lyell praising his uniformitarianism as opening bold speculation on "that mystery of mysteries, the replacement of extinct species by others" as "a natural in contradistinction to a miraculous process". When organising his notes as the ship sailed home, Darwin wrote that, if his growing suspicions about the mockingbirds, the tortoises and the Falkland Islands fox were correct, "such facts undermine the stability of Species", then cautiously added "would" before "undermine". He later wrote that such facts "seemed to me to throw some light on the origin of species".

Without telling Darwin, extracts from his letters to Henslow had been read to scientific societies, printed as a pamphlet for private distribution among members of the Cambridge Philosophical Society, and reported in magazines, including The Athenaeum. Darwin first heard of this at Cape Town, and at Ascension Island read of Sedgwick's prediction that Darwin "will have a great name among the Naturalists of Europe".

Inception of Darwin's evolutionary theory

Further information: Inception of Darwin's theory

On 2 October 1836, Beagle anchored at Falmouth, Cornwall. Darwin promptly made the long coach journey to Shrewsbury to visit his home and see relatives. He then hurried to Cambridge to see Henslow, who advised him on finding available naturalists to catalogue Darwin's animal collections and to take on the botanical specimens. Darwin's father organised investments, enabling his son to be a self-funded gentleman scientist, and an excited Darwin went around the London institutions being fêted and seeking experts to describe the collections. British zoologists at the time had a huge backlog of work, due to natural history collecting being encouraged throughout the British Empire, and there was a danger of specimens just being left in storage.

Charles Lyell eagerly met Darwin for the first time on 29 October and soon introduced him to the up-and-coming anatomist Richard Owen, who had the facilities of the Royal College of Surgeons to work on the fossil bones collected by Darwin. Owen's surprising results included other gigantic extinct ground sloths as well as the Megatherium Darwin had identified, a near complete skeleton of the unknown Scelidotherium and a hippopotamus-sized rodent-like skull named Toxodon resembling a giant capybara. The armour fragments were actually from Glyptodon, a huge armadillo-like creature, as Darwin had initially thought. These extinct creatures were related to living species in South America.

In mid-December, Darwin took lodgings in Cambridge to arrange expert classification of his collections, and prepare his own research for publication. Questions of how to combine his diary into the Narrative were resolved at the end of the month when FitzRoy accepted Broderip's advice to make it a separate volume, and Darwin began work on his Journal and Remarks.

Darwin's first paper showed that the South American landmass was slowly rising. With Lyell's enthusiastic backing, he read it to the Geological Society of London on 4 January 1837. On the same day, he presented his mammal and bird specimens to the Zoological Society. The ornithologist John Gould soon announced that the Galápagos birds that Darwin had thought a mixture of blackbirds, "gros-beaks" and finches, were, in fact, twelve separate species of finches. On 17 February, Darwin was elected to the Council of the Geological Society, and Lyell's presidential address presented Owen's findings on Darwin's fossils, stressing geographical continuity of species as supporting his uniformitarian ideas.

Early in March, Darwin moved to London to be near this work, joining Lyell's social circle of scientists and experts such as Charles Babbage, who described God as a programmer of laws. Darwin stayed with his freethinking brother Erasmus, part of this Whig circle and a close friend of the writer Harriet Martineau, who promoted the Malthusianism that underpinned the controversial Whig Poor Law reforms to stop welfare from causing overpopulation and more poverty. As a Unitarian, she welcomed the radical implications of transmutation of species, promoted by Grant and younger surgeons influenced by Geoffroy. Transmutation was anathema to Anglicans defending social order, but reputable scientists openly discussed the subject, and there was wide interest in John Herschel's letter praising Lyell's approach as a way to find a natural cause of the origin of new species.

Gould met Darwin and told him that the Galápagos mockingbirds from different islands were separate species, not just varieties, and what Darwin had thought was a "wren" was in the finch group. Darwin had not labelled the finches by island, but from the notes of others on the ship, including FitzRoy, he allocated species to islands. The two rheas were distinct species, and on 14 March Darwin announced how their distribution changed going southwards.

By mid-March 1837, barely six months after his return to England, Darwin was speculating in his Red Notebook on the possibility that "one species does change into another" to explain the geographical distribution of living species such as the rheas, and extinct ones such as the strange extinct mammal Macrauchenia, which resembled a giant guanaco, a llama relative. Around mid-July, he recorded in his "B" notebook his thoughts on lifespan and variation across generations – explaining the variations he had observed in Galápagos tortoises, mockingbirds, and rheas. He sketched branching descent, and then a genealogical branching of a single evolutionary tree, in which "It is absurd to talk of one animal being higher than another", thereby discarding Lamarck's idea of independent lineages progressing to higher forms.

Overwork, illness, and marriage

Further information: Health of Charles Darwin

While developing this intensive study of transmutation, Darwin became mired in more work. Still rewriting his Journal, he took on editing and publishing the expert reports on his collections, and with Henslow's help obtained a Treasury grant of £1,000 to sponsor this multi-volume Zoology of the Voyage of H.M.S. Beagle, a sum equivalent to about £115,000 in 2021. He stretched the funding to include his planned books on geology, and agreed to unrealistic dates with the publisher. As the Victorian era began, Darwin pressed on with writing his Journal, and in August 1837 began correcting printer's proofs.

As Darwin worked under pressure, his health suffered. On 20 September, he had "an uncomfortable palpitation of the heart", so his doctors urged him to "knock off all work" and live in the country for a few weeks. After visiting Shrewsbury, he joined his Wedgwood relatives at Maer Hall, Staffordshire, but found them too eager for tales of his travels to give him much rest. His charming, intelligent, and cultured cousin Emma Wedgwood, nine months older than Darwin, was nursing his invalid aunt. His uncle Josiah pointed out an area of ground where cinders had disappeared under loam and suggested that this might have been the work of earthworms, inspiring "a new & important theory" on their role in soil formation, which Darwin presented at the Geological Society on 1 November 1837. His Journal was printed and ready for publication by the end of February 1838, as was the first volume of the Narrative, but FitzRoy was still working hard to finish his own volume.

William Whewell pushed Darwin to take on the duties of Secretary of the Geological Society. After initially declining the work, he accepted the post in March 1838. Despite the grind of writing and editing the Beagle reports, Darwin made remarkable progress on transmutation, taking every opportunity to question expert naturalists and, unconventionally, people with practical experience in selective breeding such as farmers and pigeon fanciers. Over time, his research drew on information from his relatives and children, the family butler, neighbours, colonists and former shipmates. He included mankind in his speculations from the outset, and on seeing an orangutan in the zoo on 28 March 1838 noted its childlike behaviour.

The strain took a toll, and by June he was being laid up for days on end with stomach problems, headaches and heart symptoms. For the rest of his life, he was repeatedly incapacitated with episodes of stomach pains, vomiting, severe boils, palpitations, trembling and other symptoms, particularly during times of stress, such as attending meetings or making social visits. The cause of Darwin's illness remained unknown, and attempts at treatment had only ephemeral success.

On 23 June, he took a break and went "geologising" in Scotland. He visited Glen Roy in glorious weather to see the parallel "roads" cut into the hillsides at three heights. He later published his view that these were marine-raised beaches, but then had to accept that they were shorelines of a proglacial lake.

Fully recuperated, he returned to Shrewsbury in July 1838. Used to jotting down daily notes on animal breeding, he scrawled rambling thoughts about marriage, career and prospects on two scraps of paper, one with columns headed "Marry" and "Not Marry". Advantages under "Marry" included "constant companion and a friend in old age ... better than a dog anyhow", against points such as "less money for books" and "terrible loss of time". Having decided in favour of marriage, he discussed it with his father, then went to visit his cousin Emma on 29 July. At this time he did not get around to proposing, but against his father's advice, he mentioned his ideas on transmutation. He married Emma on 29 January 1839 and they were the parents of ten children, seven of whom survived to adulthood.

Malthus and natural selection

Continuing his research in London, Darwin's wide reading now included the sixth edition of Malthus's An Essay on the Principle of Population. On 28 September 1838, he noted its assertion that human "population, when unchecked, goes on doubling itself every twenty-five years, or increases in a geometrical ratio", a geometric progression so that population soon exceeds food supply in what is known as a Malthusian catastrophe. Darwin was well-prepared to compare this to Augustin de Candolle's "warring of the species" of plants and the struggle for existence among wildlife, explaining how numbers of a species kept roughly stable. As species always breed beyond available resources, favourable variations would make organisms better at surviving and passing the variations on to their offspring, while unfavourable variations would be lost. He wrote that the "final cause of all this wedging, must be to sort out proper structure, & adapt it to changes", so that "One may say there is a force like a hundred thousand wedges trying force into every kind of adapted structure into the gaps of in the economy of nature, or rather forming gaps by thrusting out weaker ones." This would result in the formation of new species. As he later wrote in his Autobiography:

In October 1838, that is, fifteen months after I had begun my systematic enquiry, I happened to read for amusement Malthus on Population, and being well prepared to appreciate the struggle for existence which everywhere goes on from long-continued observation of the habits of animals and plants, it at once struck me that under these circumstances favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species. Here, then, I had at last got a theory by which to work...

By mid-December, Darwin saw a similarity between farmers picking the best stock in selective breeding, and a Malthusian Nature selecting from chance variants so that "every part of newly acquired structure is fully practical and perfected", thinking this comparison "a beautiful part of my theory". He later called his theory natural selection, an analogy with what he termed the "artificial selection" of selective breeding.

On 11 November, he returned to Maer and proposed to Emma, once more telling her his ideas. She accepted, then in exchanges of loving letters showed how she valued his openness in sharing their differences, while expressing her strong Unitarian beliefs and concerns that his honest doubts might separate them in the afterlife. While he was house-hunting in London, bouts of illness continued and Emma wrote urging him to get some rest, almost prophetically remarking "So don't be ill any more my dear Charley till I can be with you to nurse you." He found what they called "Macaw Cottage" (because of its gaudy interiors) in Gower Street, then moved his "museum" in over Christmas. On 24 January 1839, Darwin was elected a Fellow of the Royal Society (FRS).

On 29 January, Darwin and Emma Wedgwood were married at Maer in an Anglican ceremony arranged to suit the Unitarians, then immediately caught the train to London and their new home.

Geology books, barnacles, evolutionary research

Further information: Development of Darwin's theory

Darwin now had the framework of his theory of natural selection "by which to work", as his "prime hobby". His research included extensive experimental selective breeding of plants and animals, finding evidence that species were not fixed and investigating many detailed ideas to refine and substantiate his theory. For fifteen years this work was in the background to his main occupation of writing on geology and publishing expert reports on the Beagle collections, in particular, the barnacles.

The impetus of Darwin's barnacle research came from a collection of a barnacle colony from Chile in 1835, which he dubbed Mr. Arthrobalanus. His confusion over the relationship of this species (Cryptophialus minutus) to other barnacles caused him to fixate on the systematics of the taxa. He wrote his first examination of the species in 1846 but did not formally describe it until 1854.

FitzRoy's long-delayed Narrative was published in May 1839. Darwin's Journal and Remarks got good reviews as the third volume, and on 15 August it was published on its own. Early in 1842, Darwin wrote about his ideas to Charles Lyell, who noted that his ally "denies seeing a beginning to each crop of species".

Darwin's book The Structure and Distribution of Coral Reefs on his theory of atoll formation was published in May 1842 after more than three years of work, and he then wrote his first "pencil sketch" of his theory of natural selection. To escape the pressures of London, the family moved to rural Down House in Kent in September. On 11 January 1844, Darwin mentioned his theorising to the botanist Joseph Dalton Hooker, writing with melodramatic humour "it is like confessing a murder". Hooker replied, "There may, in my opinion, have been a series of productions on different spots, & also a gradual change of species. I shall be delighted to hear how you think that this change may have taken place, as no presently conceived opinions satisfy me on the subject."

By July, Darwin had expanded his "sketch" into a 230-page "Essay", to be expanded with his research results if he died prematurely. In November, the anonymously published sensational best-seller Vestiges of the Natural History of Creation brought wide interest in transmutation. Darwin scorned its amateurish geology and zoology, but carefully reviewed his own arguments. Controversy erupted, and it continued to sell well despite contemptuous dismissal by scientists.

Darwin completed his third geological book in 1846. He now renewed a fascination and expertise in marine invertebrates, dating back to his student days with Grant, by dissecting and classifying the barnacles he had collected on the voyage, enjoying observing beautiful structures and thinking about comparisons with allied structures. In 1847, Hooker read the "Essay" and sent notes that provided Darwin with the calm critical feedback that he needed, but would not commit himself and questioned Darwin's opposition to continuing acts of creation.

In an attempt to improve his chronic ill health, Darwin went in 1849 to Dr. James Gully's Malvern spa and was surprised to find some benefit from hydrotherapy. Then, in 1851, his treasured daughter Annie fell ill, reawakening his fears that his illness might be hereditary. She died the same year after a long series of crises.

In eight years of work on barnacles, Darwin's theory helped him to find "homologies" showing that slightly changed body parts served different functions to meet new conditions, and in some genera he found minute males parasitic on hermaphrodites, showing an intermediate stage in evolution of distinct sexes. In 1853, it earned him the Royal Society's Royal Medal, and it made his reputation as a biologist. Upon the conclusion of his research, Darwin declared "I hate a barnacle as no man ever did before." In 1854, he became a Fellow of the Linnean Society of London, gaining postal access to its library. He began a major reassessment of his theory of species, and in November realised that divergence in the character of descendants could be explained by them becoming adapted to "diversified places in the economy of nature".

Publication of the theory of natural selection

Further information: Publication of Darwin's theory

By the start of 1856, Darwin was investigating whether eggs and seeds could survive travel across seawater to spread species across oceans. Hooker increasingly doubted the traditional view that species were fixed, but their young friend Thomas Henry Huxley was still firmly against the transmutation of species. Lyell was intrigued by Darwin's speculations without realising their extent. When he read a paper by Alfred Russel Wallace, "On the Law which has Regulated the Introduction of New Species", he saw similarities with Darwin's thoughts and urged him to publish to establish precedence.

Though Darwin saw no threat, on 14 May 1856 he began writing a short paper. Finding answers to difficult questions held him up repeatedly, and he expanded his plans to a "big book on species" titled Natural Selection, which was to include his "note on Man". He continued his research, obtaining information and specimens from naturalists worldwide, including Wallace who was working in Borneo.

In mid-1857, he added a section heading, "Theory applied to Races of Man", but did not add text on this topic. On 5 September 1857, Darwin sent the American botanist Asa Gray a detailed outline of his ideas, including an abstract of Natural Selection, which omitted human origins and sexual selection. In December, Darwin received a letter from Wallace asking if the book would examine human origins. He responded that he would avoid that subject, "so surrounded with prejudices", while encouraging Wallace's theorising and adding that "I go much further than you."

Darwin's book was only partly written when, on 18 June 1858, he received a paper from Wallace describing natural selection. Shocked that he had been "forestalled", Darwin sent it on that day to Lyell, as requested by Wallace, and although Wallace had not asked for publication, Darwin suggested he would send it to any journal that Wallace chose. His family was in crisis, with children in the village dying of scarlet fever, and he put matters in the hands of his friends. After some discussion, with no reliable way of involving Wallace, Lyell and Hooker decided on a joint presentation at the Linnean Society on 1 July of On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection. On the evening of 28 June, Darwin's baby son died of scarlet fever after almost a week of severe illness, and he was too distraught to attend.

There was little immediate attention to this announcement of the theory; the president of the Linnean Society remarked in May 1859 that the year had not been marked by any revolutionary discoveries. Only one review rankled enough for Darwin to recall it later; Professor Samuel Haughton of Dublin claimed that "all that was new in them was false, and what was true was old". Darwin struggled for thirteen months to produce an abstract of his "big book", suffering from ill health but getting constant encouragement from his scientific friends. Lyell arranged to have it published by John Murray.

On the Origin of Species proved unexpectedly popular, with the entire stock of 1,250 copies oversubscribed when it went on sale to booksellers on 22 November 1859. In the book, Darwin set out "one long argument" of detailed observations, inferences and consideration of anticipated objections. In making the case for common descent, he included evidence of homologies between humans and other mammals. Having outlined sexual selection, he hinted that it could explain differences between human races. He avoided explicit discussion of human origins, but implied the significance of his work with the sentence; "Light will be thrown on the origin of man and his history." His theory is simply stated in the introduction:

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.

At the end of the book, he concluded that:

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.

The last word was the only variant of "evolved" in the first five editions of the book. "Evolutionism" at that time was associated with other concepts, most commonly with embryological development. Darwin first used the word evolution in The Descent of Man in 1871, before adding it in 1872 to the 6th edition of The Origin of Species.

Responses to publication

Further information: Reactions to On the Origin of Species

The book aroused international interest, with less controversy than had greeted the popular and less scientific Vestiges of the Natural History of Creation. Though Darwin's illness kept him away from the public debates, he eagerly scrutinised the scientific response, commenting on press cuttings, reviews, articles, satires and caricatures, and corresponded on it with colleagues worldwide. The book did not explicitly discuss human origins, but included a number of hints about the animal ancestry of humans from which the inference could be made.

The first review asked, "If a monkey has become a man – what may not a man become?" It said this should be left to theologians as being too dangerous for ordinary readers. Among early favourable responses, Huxley's reviews swiped at Richard Owen, leader of the scientific establishment which Huxley was trying to overthrow.

In April, Owen's review attacked Darwin's friends and condescendingly dismissed his ideas, angering Darwin, but Owen and others began to promote ideas of supernaturally guided evolution. Patrick Matthew drew attention to his 1831 book which had a brief appendix suggesting a concept of natural selection leading to new species, but he had not developed the idea.

The Church of England's response was mixed. Darwin's old Cambridge tutors Sedgwick and Henslow dismissed the ideas, but 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 1860, the publication of Essays and Reviews by seven liberal Anglican theologians diverted clerical attention from Darwin. Its ideas, including higher criticism, were attacked by church authorities as heresy. In it, Baden Powell argued that miracles broke God's laws, so belief in them was atheistic, and praised "Mr Darwin's masterly volume [supporting] the grand principle of the self-evolving powers of nature".

Asa Gray discussed teleology with Darwin, who imported and distributed Gray's pamphlet on theistic evolution, Natural Selection is not inconsistent with natural theology. The most famous confrontation was at the public 1860 Oxford evolution debate during a meeting of the British Association for the Advancement of Science, where the Bishop of Oxford Samuel Wilberforce, though not opposed to transmutation of species, argued against Darwin's explanation and human descent from apes. Joseph Hooker argued strongly for Darwin, and Thomas Huxley's legendary retort, that he would rather be descended from an ape than a man who misused his gifts, came to symbolise a triumph of science over religion.

Even Darwin's close friends Gray, Hooker, Huxley and Lyell still expressed various reservations but gave strong support, as did many others, particularly younger naturalists. Gray and Lyell sought reconciliation with faith, while Huxley portrayed a polarisation between religion and science. He campaigned pugnaciously against the authority of the clergy in education, aiming to overturn the dominance of clergymen and aristocratic amateurs under Owen in favour of a new generation of professional scientists. Owen's claim that brain anatomy proved humans to be a separate biological order from apes was shown to be false by Huxley in a long-running dispute parodied by Kingsley as the "Great Hippocampus Question", and discredited Owen. In response to objections that the origin of life was unexplained, Darwin pointed to acceptance of Newton's law even though the cause of gravity was unknown. Despite criticisms and reservations related to this topic, he nevertheless proposed a prescient idea in an 1871 letter to Hooker in which he suggested the origin of life may have occurred in a "warm little pond".

Darwinism became a movement covering a wide range of evolutionary ideas. In 1863, Lyell's Geological Evidences of the Antiquity of Man popularised prehistory, though his caution on evolution disappointed Darwin. Weeks later Huxley's Evidence as to Man's Place in Nature showed that anatomically, humans are apes, then The Naturalist on the River Amazons by Henry Walter Bates provided empirical evidence of natural selection. Lobbying brought Darwin Britain's highest scientific honour, the Royal Society's Copley Medal, awarded on 3 November 1864. That day, Huxley held the first meeting of what became the influential "X Club" devoted to "science, pure and free, untrammelled by religious dogmas". By the end of the decade, most scientists agreed that evolution occurred, but only a minority supported Darwin's view that the chief mechanism was natural selection.

The Origin of Species was translated into many languages, becoming a staple scientific text attracting thoughtful attention from all walks of life, including the "working men" who flocked to Huxley's lectures. Darwin's theory resonated with various movements at the time and became a key fixture of popular culture. Cartoonists parodied animal ancestry in an old tradition of showing humans with animal traits, and in Britain, these droll images served to popularise Darwin's theory in an unthreatening way. While ill in 1862, Darwin began growing a beard, and when he reappeared in public in 1866, caricatures of him as an ape helped to identify all forms of evolutionism with Darwinism.

Othniel C. Marsh, America's first palaeontologist, was the first to provide solid fossil evidence to support Darwin's theory of evolution by unearthing the ancestors of the modern horse. In 1877, Marsh delivered a very influential speech before the annual meeting of the American Association for the Advancement of Science, providing a demonstrative argument for evolution. For the first time, Marsh traced the evolution of vertebrates from fish all the way through humans. Sparing no detail, he listed a wealth of fossil examples of past life forms. The significance of this speech was immediately recognized by the scientific community, and it was printed in its entirety in several scientific journals.

Descent of Man, sexual selection, and botany

Further information: Orchids to Variation, Descent of Man to Emotions, and Insectivorous Plants to Worms

Despite repeated bouts of illness during the last twenty-two years of his life, Darwin's work continued. Having published On the Origin of Species as an abstract of his theory, he pressed on with experiments, research, and writing of his "big book". He covered human descent from earlier animals, including the evolution of society and of mental abilities, as well as explaining decorative beauty in wildlife and diversifying into innovative plant studies.

Enquiries about insect pollination led in 1861 to novel studies of wild orchids, showing adaptation of their flowers to attract specific moths to each species and ensure cross fertilisation. In 1862 Fertilisation of Orchids gave his first detailed demonstration of the power of natural selection to explain complex ecological relationships, making testable predictions. As his health declined, he lay on his sickbed in a room filled with inventive experiments to trace the movements of climbing plants. Admiring visitors included Ernst Haeckel, a zealous proponent of Darwinism incorporating Lamarckism and Goethe's idealism. Wallace remained supportive, though he increasingly turned to Spiritualism.

Darwin's book The Variation of Animals and Plants Under Domestication (1868) was the first part of his planned "big book", and included his unsuccessful hypothesis of pangenesis attempting to explain heredity. It sold briskly at first, despite its size, and was translated into many languages. He wrote most of a second part, on natural selection, but it remained unpublished in his lifetime.

Lyell had already popularised human prehistory, and Huxley had shown that anatomically humans are apes. With The Descent of Man, and Selection in Relation to Sex published in 1871, Darwin set out evidence from numerous sources that humans are animals, showing continuity of physical and mental attributes, and presented sexual selection to explain impractical animal features such as the peacock's plumage as well as human evolution of culture, differences between sexes, and physical and cultural racial classification, while emphasising that humans are all one species. According to an editorial in Nature journal: "Although Charles Darwin opposed slavery and proposed that humans have a common ancestor, he also advocated a hierarchy of races, with white people higher than others."

Letter of enquiry from Charles Darwin to the physiologist John Burdon-SandersonPunch's almanac for 1882, published shortly before Darwin's death, depicts him amidst evolution from chaos to Victorian gentleman with the title Man Is But A Worm

His research using images was expanded in his 1872 book The Expression of the Emotions in Man and Animals, one of the first books to feature printed photographs, which discussed the evolution of human psychology and its continuity with the behaviour of animals. Both books proved very popular, and Darwin was impressed by the general assent with which his views had been received, remarking that "everybody is talking about it without being shocked." His conclusion was "that man with all his noble qualities, with sympathy which feels for the most debased, with benevolence which extends not only to other men but to the humblest living creature, with his god-like intellect which has penetrated into the movements and constitution of the solar system – with all these exalted powers – Man still bears in his bodily frame the indelible stamp of his lowly origin."

His evolution-related experiments and investigations led to books on Insectivorous Plants, The Effects of Cross and Self Fertilisation in the Vegetable Kingdom, different forms of flowers on plants of the same species, and The Power of Movement in Plants. He continued to collect information and exchange views from scientific correspondents all over the world, including Mary Treat, whom he encouraged to persevere in her scientific work. He was the first person to recognise the significance of carnivory in plants. His botanical work was interpreted and popularised by various writers including Grant Allen and H. G. Wells, and helped transform plant science in the late 19th century and early 20th century.

Death and funeral

See also: Darwin from Insectivorous Plants to Worms § Death

In 1882, he was diagnosed with what was called "angina pectoris" which then meant coronary thrombosis and disease of the heart. At the time of his death, the physicians diagnosed "anginal attacks", and "heart-failure"; there has since been scholarly speculation about his life-long health issues.

He died at Down House on 19 April 1882. His last words were to his family, telling Emma, "I am not the least afraid of death—Remember what a good wife you have been to me—Tell all my children to remember how good they have been to me". While she rested, he repeatedly told Henrietta and Francis, "It's almost worthwhile to be sick to be nursed by you".

He had expected to be buried in St Mary's churchyard at Downe, but at the request of Darwin's colleagues, after public and parliamentary petitioning, William Spottiswoode (President of the Royal Society) arranged for Darwin to be honoured by burial in Westminster Abbey, close to John Herschel and Isaac Newton. The funeral, held on Wednesday 26 April, was attended by thousands of people, including family, friends, scientists, philosophers and dignitaries.

Children

Further information: Darwin–Wedgwood family § Charles Darwin

William Erasmus Darwin	27 December 1839 –	8 September 1914
Anne Elizabeth Darwin	2 March 1841 –	23 April 1851
Mary Eleanor Darwin	23 September 1842 –	16 October 1842
Henrietta Emma Darwin	25 September 1843 –	17 December 1927
George Howard Darwin	9 July 1845 –	7 December 1912
Elizabeth Darwin	8 July 1847 –	8 June 1926
Francis Darwin	16 August 1848 –	19 September 1925
Leonard Darwin	15 January 1850 –	26 March 1943
Horace Darwin	13 May 1851 –	29 September 1928
Charles Waring Darwin	6 December 1856 –	28 June 1858

The Darwins had ten children: two died in infancy, and Annie's death at the age of ten had a devastating effect on her parents. Charles was a devoted father and uncommonly attentive to his children. Whenever they fell ill, he feared that they might have inherited weaknesses from inbreeding due to the close family ties he shared with his wife and cousin, Emma Wedgwood. He examined inbreeding in his writings, contrasting it with the advantages of outcrossing in many species.

Charles Waring Darwin, born in December 1856, was the tenth and last of the children. Emma Darwin was aged 48 at the time of the birth, and the child was mentally subnormal and never learnt to walk or talk. He probably had Down syndrome, which had not then been medically described. The evidence is a photograph by William Erasmus Darwin of the infant and his mother, showing a characteristic head shape, and the family's observations of the child. Charles Waring died of scarlet fever on 28 June 1858, when Darwin wrote in his journal: "Poor dear Baby died."

Of his surviving children, George, Francis and Horace became Fellows of the Royal Society, distinguished as an astronomer, botanist and civil engineer, respectively. All three were knighted. Another son, Leonard, went on to be a soldier, politician, economist, eugenicist, and mentor of the statistician and evolutionary biologist Ronald Fisher.

Views and opinions

Religious views

Further information: Religious views of Charles Darwin

Darwin's family tradition was nonconformist Unitarianism, while his father and grandfather were freethinkers, and his baptism and boarding school were Church of England. When going to Cambridge to become an Anglican clergyman, he did not "in the least doubt the strict and literal truth of every word in the Bible". He learned John Herschel's science which, like William Paley's natural theology, sought explanations in laws of nature rather than miracles and saw adaptation of species as evidence of design. On board HMS Beagle, Darwin was quite orthodox and would quote the Bible as an authority on morality. He looked for "centres of creation" to explain distribution, and suggested that the very similar antlions found in Australia and England were evidence of a divine hand.

Upon his return, he expressed a critical view of the Bible's historical accuracy and questioned the basis for considering one religion more valid than another. In the next few years, while intensively speculating on geology and the transmutation of species, he gave much thought to religion and openly discussed this with his wife Emma, whose beliefs similarly came from intensive study and questioning.

The theodicy of Paley and Thomas Malthus vindicated evils such as starvation as a result of a benevolent creator's laws, which had an overall good effect. To Darwin, natural selection produced the good of adaptation but removed the need for design, and he could not see the work of an omnipotent deity in all the pain and suffering, such as the ichneumon wasp paralysing caterpillars as live food for its eggs. Though he thought of religion as a tribal survival strategy, Darwin was reluctant to give up the idea of God as an ultimate lawgiver. He was increasingly troubled by the problem of evil.

Darwin remained close friends with the vicar of Downe, John Brodie Innes, and continued to play a leading part in the parish work of the church, but from c. 1849 would go for a walk on Sundays while his family attended church. He considered it "absurd to doubt that a man might be an ardent theist and an evolutionist" and, though reticent about his religious views, in 1879 he wrote that "I have never been an atheist in the sense of denying the existence of a God. – I think that generally ... an agnostic would be the most correct description of my state of mind".

The "Lady Hope Story", published in 1915, claimed that Darwin had reverted to Christianity on his sickbed. The claims were repudiated by Darwin's children and have been dismissed as false by historians.

Human society

Darwin's views on social and political issues reflected his time and social position. He grew up in a family of Whig reformers who, like his uncle Josiah Wedgwood, supported electoral reform and the emancipation of slaves. Darwin was passionately opposed to slavery, while seeing no problem with the working conditions of English factory workers or servants.

Taking taxidermy lessons in 1826 from the freed slave John Edmonstone, whom Darwin long recalled as "a very pleasant and intelligent man", reinforced his belief that black people shared the same feelings, and could be as intelligent as people of other races. He took the same attitude to native people he met on the Beagle voyage. Though commonplace in Britain at the time, Silliman and Bachman noticed the contrast with slave-owning America. Around twenty years later, racism became a feature of British society, but Darwin remained strongly against slavery, against "ranking the so-called races of man as distinct species", and against ill-treatment of native people.

Darwin's interaction with Yaghans (Fuegians) such as Jemmy Button during the second voyage of HMS Beagle had a profound impact on his view of indigenous peoples. At his arrival in Tierra del Fuego he made a colourful description of "Fuegian savages". This view changed as he came to know Yaghan people more in detail. By studying the Yaghans, Darwin concluded that a number of basic emotions by different human groups were the same and that mental capabilities were roughly the same as for Europeans. While interested in Yaghan culture, Darwin failed to appreciate their deep ecological knowledge and elaborate cosmology until the 1850s when he inspected a dictionary of Yaghan detailing 32,000 words. He saw that European colonisation would often lead to the extinction of native civilisations, and "tr to integrate colonialism into an evolutionary history of civilization analogous to natural history".

Darwin's view of women was that men's eminence over them was the outcome of sexual selection, a view disputed by Antoinette Brown Blackwell in her 1875 book The Sexes Throughout Nature.

Darwin was intrigued by his half-cousin Francis Galton's argument, introduced in 1865, that statistical analysis of heredity showed that moral and mental human traits could be inherited, and principles of animal breeding could apply to humans. In The Descent of Man, Darwin noted that aiding the weak to survive and have families could lose the benefits of natural selection, but cautioned that withholding such aid would endanger the instinct of sympathy, "the noblest part of our nature", and factors such as education could be more important. When Galton suggested that publishing research could encourage intermarriage within a "caste" of "those who are naturally gifted", Darwin foresaw practical difficulties and thought it "the sole feasible, yet I fear utopian, plan of procedure in improving the human race", preferring to simply publicise the importance of inheritance and leave decisions to individuals. Francis Galton named this field of study "eugenics" in 1883, after Darwin's death, and his theories were cited to promote eugenic policies.

Evolutionary social movements

Further information: Eugenics, Social effects of evolutionary theory, and Degeneration theory

Darwin's fame and popularity led to his name being associated with ideas and movements that, at times, had only an indirect relation to his writings, and sometimes went directly against his express comments.

Thomas Malthus had argued that population growth beyond resources was ordained by God to get humans to work productively and show restraint in getting families; this was used in the 1830s to justify workhouses and laissez-faire economics. Evolution was by then seen as having social implications, and Herbert Spencer's 1851 book Social Statics based ideas of human freedom and individual liberties on his Lamarckian evolutionary theory.

Soon after the Origin was published in 1859, critics derided his description of a struggle for existence as a Malthusian justification for the English industrial capitalism of the time. The term Darwinism was used for the evolutionary ideas of others, including Spencer's "survival of the fittest" as free-market progress, and Ernst Haeckel's polygenistic ideas of human development. Writers used natural selection to argue for various, often contradictory, ideologies such as laissez-faire dog-eat-dog capitalism, colonialism and imperialism. However, Darwin's holistic view of nature included "dependence of one being on another"; thus pacifists, socialists, liberal social reformers and anarchists such as Peter Kropotkin stressed the value of cooperation over struggle within a species. Darwin himself insisted that social policy should not simply be guided by concepts of struggle and selection in nature.

After the 1880s, a eugenics movement developed on ideas of biological inheritance, and for scientific justification of their ideas appealed to some concepts of Darwinism. In Britain, most shared Darwin's cautious views on voluntary improvement and sought to encourage those with good traits in "positive eugenics". During the "Eclipse of Darwinism", a scientific foundation for eugenics was provided by Mendelian genetics. Negative eugenics to remove the "feebleminded" were popular in America, Canada and Australia, and eugenics in the United States introduced compulsory sterilisation laws, followed by several other countries. Subsequently, Nazi eugenics brought the field into disrepute.

The term "Social Darwinism" was used infrequently from around the 1890s, but became popular as a derogatory term in the 1940s when used by Richard Hofstadter to attack the laissez-faire conservatism of those like William Graham Sumner who opposed reform and socialism. Since then, it has been used as a term of abuse by those opposed to what they think are the moral consequences of evolution.

Works

Further information: Charles Darwin bibliography

Darwin was a prolific writer. Even without the publication of his works on evolution, he would have had a considerable reputation as the author of The Voyage of the Beagle, as a geologist who had published extensively on South America and had solved the puzzle of the formation of coral atolls, and as a biologist who had published the definitive work on barnacles. While On the Origin of Species dominates perceptions of his work, The Descent of Man and The Expression of the Emotions in Man and Animals had considerable impact, and his books on plants including The Power of Movement in Plants were innovative studies of great importance, as was his final work on The Formation of Vegetable Mould through the Action of Worms.

Legacy and commemoration

Further information: List of things named after Charles Darwin, List of taxa described by Charles Darwin, and Commemoration of Charles Darwin

As Alfred Russel Wallace put it, Darwin had "wrought a greater revolution in human thought within a quarter of a century than any man of our time – or perhaps any time", having "given us a new conception of the world of life, and a theory which is itself a powerful instrument of research; has shown us how to combine into one consistent whole the facts accumulated by all the separate classes of workers, and has thereby revolutionised the whole study of nature". The paleoanthropologist Trenton Holliday states that "Darwin is rightly considered to be the preeminent evolutionary scientist of all time".

By around 1880, most scientists were convinced of evolution as descent with modification, though few agreed with Darwin that natural selection "has been the main but not the exclusive means of modification". During "the eclipse of Darwinism" scientists explored alternative mechanisms. Then Ronald Fisher incorporated Mendelian genetics in The Genetical Theory of Natural Selection, leading to population genetics and the modern evolutionary synthesis, which continues to develop. Scientific discoveries have confirmed and validated Darwin's key insights.

Geographical features given his name include Darwin Sound and Mount Darwin, both named while he was on the Beagle voyage, and Darwin Harbour, named by his former shipmates on its next voyage, which eventually became the location of Darwin, the capital city of Australia's Northern Territory. Darwin's name was given, formally or informally, to numerous plants and animals, including many he had collected on the voyage. The Linnean Society of London began awards of the Darwin–Wallace Medal in 1908, to mark fifty years from the joint reading on 1 July 1858 of papers by Darwin and Wallace publishing their theory. Further awards were made in 1958 and 2008; since 2010, the awards have been annual. Darwin College, a postgraduate college at Cambridge University founded in 1964, is named after the Darwin family. From 2000 to 2017, UK £10 banknotes issued by the Bank of England featured Darwin's portrait printed on the reverse, along with a hummingbird and HMS Beagle.

See also

• 1991 Darwin
• Creation (biographical drama film)
• Creation–evolution controversy
• European and American voyages of scientific exploration
• History of biology
• History of evolutionary thought
• List of coupled cousins
• List of multiple discoveries
• Multiple discovery
• Portraits of Charles Darwin
• Tinamou egg
• Universal Darwinism

Notes

I. Robert FitzRoy was to become known after the voyage for biblical literalism, but at this time he had considerable interest in Lyell's ideas, and they met before the voyage when Lyell asked for observations to be made in South America. FitzRoy's diary during the ascent of the River Santa Cruz in Patagonia recorded his opinion that the plains were raised beaches, but on return, newly married to a very religious lady, he recanted these ideas.(Browne 1995, pp. 186, 414)

II. In the section "Morphology" of Chapter XIII of On the Origin of Species, Darwin commented on homologous bone patterns between humans and other mammals, writing: "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?" and in the concluding chapter: "The framework of bones being the same in the hand of a man, wing of a bat, fin of the porpoise, and leg of the horse … at once explain themselves on the theory of descent with slow and slight successive modifications."

III. In On the Origin of Species Darwin mentioned human origins in his concluding remark that "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."

In "Chapter VI: Difficulties on Theory" he referred to sexual selection: "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."

In The Descent of Man of 1871, Darwin discussed the first passage: "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." In a preface to the 1874 second edition, he added a reference to the second point: "it has been said by several critics, that when I found that many details of structure in man could not be explained through natural selection, I invented sexual selection; I gave, however, a tolerably clear sketch of this principle in the first edition of the 'Origin of Species,' and I there stated that it was applicable to man."

IV. See, for example, WILLA volume 4, Charlotte Perkins Gilman and the Feminization of Education by Deborah M. De Simone: "Gilman shared many basic educational ideas with the generation of thinkers who matured during the period of "intellectual chaos" caused by Darwin's Origin of the Species. Marked by the belief that individuals can direct human and social evolution, many progressives came to view education as the panacea for advancing social progress and for solving such problems as urbanisation, poverty, or immigration."

V. See, for example, the song "A lady fair of lineage high" from Gilbert and Sullivan's Princess Ida, which describes the descent of man (but not woman!) from apes.

VI. Darwin's belief that black people had the same essential humanity as Europeans, and had many mental similarities, was reinforced by the lessons he had from John Edmonstone in 1826. Early in the Beagle voyage, Darwin nearly lost his position on the ship when he criticised FitzRoy's defence and praise of slavery. (Darwin 1958, p. 74) He wrote home about "how steadily the general feeling, as shown at elections, has been rising against Slavery. What a proud thing for England if she is the first European nation which utterly abolishes it! I was told before leaving England that after living in slave countries all my opinions would be altered; the only alteration I am aware of is forming a much higher estimate of the negro character." (Darwin 1887, p. 246) Regarding Fuegians, he "could not have believed how wide was the difference between savage and civilized man: it is greater than between a wild and domesticated animal, inasmuch as in man there is a greater power of improvement", but he knew and liked civilised Fuegians like Jemmy Button: "It seems yet wonderful to me, when I think over all his many good qualities, that he should have been of the same race, and doubtless partaken of the same character, with the miserable, degraded savages whom we first met here." (Darwin 1845, pp. 205, 207–208)

In the Descent of Man, he mentioned the similarity of Fuegians' and Edmonstone's minds to Europeans' when arguing against "ranking the so-called races of man as distinct species".

He rejected the ill-treatment of native people, and for example wrote of massacres of Patagonian men, women, and children, "Every one here is fully convinced that this is the most just war, because it is against barbarians. Who would believe in this age that such atrocities could be committed in a Christian civilized country?"(Darwin 1845, p. 102)

VII. Geneticists studied human heredity as Mendelian inheritance, while eugenics movements sought to manage society, with a focus on social class in the United Kingdom, and on disability and ethnicity in the United States, leading to geneticists seeing this movement as impractical pseudoscience. A shift from voluntary arrangements to "negative" eugenics included compulsory sterilisation laws in the United States, copied by Nazi Germany as the basis for Nazi eugenics based on virulent racism and "racial hygiene".
(Thurtle, Phillip (17 December 1996). "the creation of genetic identity". SEHR. Vol. 5, no. Supplement: Cultural and Technological Incubations of Fascism. Retrieved 11 November 2008. Edwards, A. W. F. (1 April 2000). "The Genetical Theory of Natural Selection". Genetics. Vol. 154, no. April 2000. pp. 1419–1426. PMC 1461012. PMID 10747041. Retrieved 11 November 2008.Wilkins, John. "Evolving Thoughts: Darwin and the Holocaust 3: eugenics". Archived from the original on 5 December 2008. Retrieved 11 November 2008.)

VIII. David Quammen writes of his "theory that turned to these arcane botanical studies – producing more than one book that was solidly empirical, discreetly evolutionary, yet a 'horrid bore' – at least partly so that the clamorous controversialists, fighting about apes and angels and souls, would leave him... alone". David Quammen, "The Brilliant Plodder" (review of Ken Thompson, Darwin's Most Wonderful Plants: A Tour of His Botanical Legacy, University of Chicago Press, 255 pp.; Elizabeth Hennessy, On the Backs of Tortoises: Darwin, the Galápagos, and the Fate of an Evolutionary Eden, Yale University Press, 310 pp.; Bill Jenkins, Evolution Before Darwin: Theories of the Transmutation of Species in Edinburgh, 1804–1834, Edinburgh University Press, 222 pp.), The New York Review of Books, vol. LXVII, no. 7 (23 April 2020), pp. 22–24. Quammen, quoted from p. 24 of his review.

Citations

1. Freeman 2007, p. 76.
2. "Charles Darwin's personal finances revealed in new find". The Telegraph. 22 March 2009. Archived from the original on 19 October 2017.
3. ^ "Search Results: Record – Darwin; Charles Robert". The Royal Society Collections Catalogues. 20 June 2015. Archived from the original on 2 December 2021. Retrieved 2 December 2021.
4. ^ Freeman 2007, p. 106.
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By Charles Darwin

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• "The Complete Work of Charles Darwin Online". Retrieved 4 March 2024.
• Works by Charles Darwin in eBook form at Standard Ebooks
• Works by Charles Darwin at Project Gutenberg
• Works by or about Charles Robert Darwin at the Internet Archive
• Works by Charles Darwin at LibriVox (public domain audiobooks)
• The Complete Works of Charles Darwin Online – Darwin Online; Darwin's publications, private papers and bibliography, supplementary works including biographies, obituaries and reviews
• Darwin Correspondence Project Full text and notes for complete correspondence to 1867, with summaries of all the rest, and pages of commentary
• Darwin Manuscript Project
• "Archival material relating to Charles Darwin". UK National Archives.
• View books owned and annotated by Charles Darwin at the online Biodiversity Heritage Library.
• Works by Charles Darwin at the Biodiversity Heritage Library
• Digitised Darwin Manuscripts in Cambridge Digital Library
• Portraits of Charles Darwin at the National Portrait Gallery, London
• Newspaper clippings about Charles Darwin in the 20th Century Press Archives of the ZBW
• Charles Darwin in the British horticultural press – Occasional Papers from RHS Lindley Library, volume 3 July 2010
• Scientific American, 29 April 1882, pp. 256, Obituary of Charles Darwin
• "Charles Darwin". Internet Encyclopedia of Philosophy.

v t e Charles Darwin
Life	Darwin–Wedgwood family Erasmus Darwin Josiah Wedgwood Education Voyage on HMS Beagle Inception of theory Development of theory Publication of theory Reactions to On the Origin of Species Orchids to Variation Descent of Man to Emotions Insectivorous Plants to Worms Religious views Health Women Portraits of Darwin (caricatures)
Writings	Extracts from Letters to Henslow (1835) The Voyage of the Beagle (1839) The Structure and Distribution of Coral Reefs (1842) Zoology of the Voyage of H.M.S. Beagle (1838–1843) Geological Observations on the Volcanic Islands (1844) Geological Observations on South America (1846) "On the Tendency of Species to form Varieties" (1858) On the Origin of Species (1859) Fertilisation of Orchids (1862) The Variation of Animals and Plants Under Domestication (1868) Natural Selection The Descent of Man, and Selection in Relation to Sex (1871) The Expression of the Emotions in Man and Animals (1872) Insectivorous Plants (1875) On the Movements and Habits of Climbing Plants (1875) The Effects of Cross and Self Fertilisation in the Vegetable Kingdom (1876) The Different Forms of Flowers on Plants of the Same Species (1877) The Power of Movement in Plants (1880) The Formation of Vegetable Mould Through the Action of Worms (1881) The Autobiography of Charles Darwin (1887) Correspondence The Life and Letters of Charles Darwin, 1887 More Letters of Charles Darwin, 1903 List of described taxa
Related	Eugenics Dysgenics Vestiges of the Natural History of Creation The Genetical Theory of Natural Selection (1930) History of evolutionary thought Pangenesis Darwin Industry Commemoration things named for Darwin Darwinism Neural Quantum Social Universal Alternatives to Eclipse of HMS Beagle Darwin Medal Darwin Awards Huxley family

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v t e Evolutionary biology Introduction Outline Timeline of evolution History of life Index Evolution Abiogenesis Adaptation Adaptive radiation Altruism Cheating Reciprocal Baldwin effect Cladistics Coevolution Mutualism Common descent Convergence Divergence Earliest known life forms Evidence of evolution Evolutionary arms race Evolutionary pressure Exaptation Extinction Event Homology Last universal common ancestor Macroevolution Microevolution Mismatch Non-adaptive radiation Origin of life Panspermia Parallel evolution Signalling theory Handicap principle Speciation Species Species complex Taxonomy Unit of selection Gene-centered view of evolution Population genetics Artificial selection Biodiversity Evolutionarily stable strategy Fisher's principle Fitness Inclusive Gene flow Genetic drift Kin selection Parental investment Parent–offspring conflict Mutation Population Natural selection Sexual dimorphism Sexual selection Flowering plants Fungi Mate choice Social selection Trivers–Willard hypothesis Variation Development Canalisation Evolutionary developmental biology Genetic assimilation Inversion Modularity Phenotypic plasticity Of taxa Bacteria Birds origin Brachiopods Molluscs Cephalopods Dinosaurs Fish Fungi Insects butterflies Life Mammals cats canids wolves dogs hyenas dolphins and whales horses Kangaroos primates humans lemurs sea cows Plants pollinator-mediated Reptiles Spiders Tetrapods Viruses Of organs Cell DNA Flagella Eukaryotes symbiogenesis chromosome endomembrane system mitochondria nucleus plastids In animals eye hair auditory ossicle nervous system brain Of processes Aging Death Programmed cell death Avian flight Biological complexity Cooperation Color vision in primates Emotion Empathy Ethics Eusociality Immune system Metabolism Monogamy Morality Mosaic evolution Multicellularity Sexual reproduction Gamete differentiation/sexes Life cycles/nuclear phases Mating types Meiosis Sex-determination Snake venom Tempo and modes Gradualism/Punctuated equilibrium/Saltationism Micromutation/Macromutation Uniformitarianism/Catastrophism Speciation Allopatric Anagenesis Catagenesis Cladogenesis Cospeciation Ecological Hybrid Non-ecological Parapatric Peripatric Reinforcement Sympatric History Renaissance and Enlightenment Transmutation of species David Hume Dialogues Concerning Natural Religion Charles Darwin On the Origin of Species History of paleontology Transitional fossil Blending inheritance Mendelian inheritance The eclipse of Darwinism Neo-Darwinism Modern synthesis History of molecular evolution Extended evolutionary synthesis Philosophy Darwinism Alternatives Catastrophism Lamarckism Orthogenesis Mutationism Saltationism Structuralism Spandrel Theistic Vitalism Teleology in biology Related Biogeography Ecological genetics Evolutionary medicine Group selection Cultural evolution Cultural group selection Dual inheritance theory Hologenome theory of evolution Missing heritability problem Molecular evolution Astrobiology Phylogenetics Tree Polymorphism Protocell Systematics Transgenerational epigenetic inheritance Category Portal v t e Ethology Branches Animal cognition Animal communication Animal consciousness Animal culture Animal sexual behaviour Animal welfare science Anthrozoology Bee learning and communication Behavioural ecology Behavioral endocrinology Behavioural genetics Breed Cognitive ethology Comfort behaviour Grooming Comparative psychology Emotion in animals Ethogram Evolutionary neuroscience Feeding Hover Human ethology Instinct Learning Neuroethology Pain in animals Philosophical ethology Sociobiology Stereotypy Structures Hive Honeycomb Nest Instinct Swarm Tool use by non-humans Zoosemiotics Zoomusicology Ethologists Patrick Bateson Marc Bekoff Donald Broom John B. 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Wilson Solly Zuckerman Societies Association for the Study of Animal Behaviour International Society for Applied Ethology Journals Animal Behaviour Animal Cognition Animal Welfare Behavioral Ecology Behaviour Category v t e Zoology Outline Branches Anthrozoology Arthropodology (Arachnology (Acarology) Carcinology Entomology (Coleopterology Lepidopterology Melittology Myrmecology Orthopterology) Myriapodology) Biological anthropology Bryozoology Cnidariology Ethnozoology Ethology Helminthology Herpetology (Batrachology Ophiology Testudinology) Ichthyology Malacology (Conchology Teuthology) Mammalogy (Cetology Cynology Felinology Hippology Primatology) Nematology Neuroethology Ornithology Parasitology Paleozoology Planktology Zooarchaeology Animal groups Porifera Ctenophora Placozoa Cnidaria Xenacoelomorpha Ambulacraria Chordata Aschelminthes Arthropoda Platyhelminthes Mollusca Annelida Animal anatomy Animal morphology Histology Tissues Epithelial tissue Muscular tissue Nervous tissue Connective tissues Mineralized tissues Molecular anatomy Anatomy and morphology Mammalian anatomy and morphology Human Dog Elephant Cat Bird anatomy Fish anatomy Shark anatomy Spider anatomy Insect morphology Animal physiology General physiology Respiration Respiratory system Breathing Gas exchange Respiratory pigment Cellular respiration Vascular system Blood Lymph Blood vessels Arteries Veins Capillaries Heart Water vascular system By species Insect physiology Fish physiology Zoologists Karl Ernst von Baer Georges Cuvier Charles Darwin Jean-Henri Fabre William Kirby Carl Linnaeus Konrad Lorenz Thomas Say Jakob von Uexküll Alfred Russel Wallace History Pre-Darwin Post-Darwin Timeline of zoology Category v t e Evolutionary psychologists Evolutionary psychology Biologists / neuroscientists Bernard Crespi John Crook Charles Darwin Richard Dawkins Jared Diamond W. D. Hamilton Alfred Kinsey Peter Kropotkin Gordon Orians Jaak Panksepp Margie Profet Peter Richerson Giacomo Rizzolatti Randy Thornhill Robert Trivers Carel van Schaik Claus Wedekind Mary Jane West-Eberhard Wolfgang Wickler George C. Williams David Sloan Wilson E. O. Wilson Richard Wrangham Anthropologists Jerome H. Barkow Christopher Boehm Robert Boyd Donald E. Brown Napoleon Chagnon Robin Dunbar Daniel Fessler Mark Flinn John D. Hawks Joseph Henrich Ruth Mace Daniel Nettle Stephen Shennan Donald Symons John Tooby Pierre van den Berghe Psychologists / cognitive scientists Mary Ainsworth Simon Baron-Cohen Justin L. Barrett Jay Belsky Jesse Bering David F. Bjorklund Paul Bloom John Bowlby Pascal Boyer Joseph Bulbulia David Buss Josep Call Anne Campbell Donald T. Campbell Peter Carruthers Noam Chomsky Leda Cosmides Martin Daly Paul Ekman Bruce J. Ellis Anne Fernald Aurelio José Figueredo Diana Fleischman Uta Frith Gordon G. Gallup David C. Geary Gerd Gigerenzer Peter Gray Jonathan Haidt Harry Harlow Judith Rich Harris Martie Haselton Stephen Kaplan Douglas T. Kenrick Simon M. Kirby Robert Kurzban Brian MacWhinney Michael T. McGuire Geoffrey Miller Darcia Narvaez Katherine Nelson Randolph M. Nesse Steven Neuberg David Perrett Henry Plotkin Steven Pinker Paul Rozin Mark Schaller David P. Schmitt Nancy Segal Todd K. Shackelford Roger Shepard Irwin Silverman Peter K. Smith Dan Sperber Anthony Stevens Frank Sulloway Michael Tomasello Joshua Tybur Mark van Vugt Andrew Whiten Glenn Wilson Margo Wilson Other social scientists Christopher Badcock Samuel Bowles Ernst Fehr Herbert Gintis Dominic D. P. Johnson Gad Saad Literary theorists / philosophers Edmund Burke Joseph Carroll Daniel Dennett Denis Dutton Thomas Hobbes David Hume Research centers/ organizations Center for Evolutionary Psychology Human Behavior and Evolution Society Max Planck Institute for Evolutionary Anthropology Max Planck Institute for Human Cognitive and Brain Sciences New England Complex Systems Institute Publications The Adapted Mind Evolution and Human Behavior The Evolution of Human Sexuality Evolution, Mind and Behaviour Evolutionary Behavioral Sciences Evolutionary Psychology  Evolutionary psychology  Psychology portal  Evolutionary biology portal v t e Copley Medallists (1851–1900) Richard Owen (1851) Alexander von Humboldt (1852) Heinrich Wilhelm Dove (1853) Johannes Peter Müller (1854) Léon Foucault (1855) Henri Milne-Edwards (1856) Michel Eugène Chevreul (1857) Charles Lyell (1858) Wilhelm Eduard Weber (1859) Robert Bunsen (1860) Louis Agassiz (1861) Thomas Graham (1862) Adam Sedgwick (1863) Charles Darwin (1864) Michel Chasles (1865) Julius Plücker (1866) Karl Ernst von Baer (1867) Charles Wheatstone (1868) Henri Victor Regnault (1869) James Prescott Joule (1870) Julius Robert von Mayer (1871) Friedrich Wöhler (1872) Hermann von Helmholtz (1873) Louis Pasteur (1874) August Wilhelm von Hofmann (1875) Claude Bernard (1876) James Dwight Dana (1877) Jean-Baptiste Boussingault (1878) Rudolf Clausius (1879) James Joseph Sylvester (1880) Charles Adolphe Wurtz (1881) Arthur Cayley (1882) William Thomson (1883) Carl Ludwig (1884) Friedrich August Kekulé von Stradonitz (1885) Franz Ernst Neumann (1886) Joseph Dalton Hooker (1887) Thomas Henry Huxley (1888) George Salmon (1889) Simon Newcomb (1890) Stanislao Cannizzaro (1891) Rudolf Virchow (1892) George Gabriel Stokes (1893) Edward Frankland (1894) Karl Weierstrass (1895) Carl Gegenbaur (1896) Albert von Kölliker (1897) William Huggins (1898) John William Strutt (1899) Marcellin Berthelot (1900) v t e Fellows of the Royal Society elected in 1839 Fellows Thomas Dyke Acland George Barker Beriah Botfield Robert Carrington Arthur Conolly Charles Darwin Edward Davies Davenport Henry Mangles Denham Richard Drew Henry Drummond Arthur Farre Thomas William Fletcher William James Frodsham Thomas Gaskin George Godwin John T. Graves Edwin Guest George Gulliver James Halliwell-Phillipps Peter Hardy James Heywood John Hilton John Hogg Gilbert Wakefield Mackmurdo Samuel Roffey Maitland Henry Moseley H. Alexander Ormsby William Reid Robert Rigg John Rogers George Leith Roupell William Sharpey Clement Tudway Swanston James Joseph Sylvester Charles Thorp Charles Turnor John Wesley Williams James Yates Foreign Christopher Hansteen Macedonio Melloni Adolphe Quetelet Félix Savart v t e Nonverbal communication Modalities Physical Blushing Body language / Kinesics Body-to-body communication Facial expression Facial Action Coding System Microexpression Subtle expression Gesture List Speech-independent gestures Haptic communication Imitation Interpersonal synchrony Laughter Oculesics Eye contact Pupil dilation Olfaction Posture Proxemics Speech Affect Emotional prosody Paralanguage Intonation Loudness Prosody Rhythm Stress Tone Voice quality Social context Chronemics Conventions Display rules Habitus High-context and low-context cultures Interpersonal relationship Social norm Other Emoticon / Smiley One-bit message Missed call Silent service code Unconscious Microexpression Non-verbal leakage Multi-faceted Affect display Deception Emotion recognition First impression Intimacy Broader concepts Basic interpersonal communicative skills Communication Emotional intelligence Nunchi People skills Semiotics Social behavior Social cue Social competence Social skills Unsaid Further information Disorders Aprosodia Asperger syndrome Autism Fragile X Pervasive developmental disorder not otherwise specified Childhood disintegrative disorder Rett syndrome Dyssemia Nonverbal learning disorder Social (pragmatic) communication disorder Neuroanatomy Limbic system / Limbic lobe Mirror neuron Applications Cold reading Lie detection Freudian slip Poker tell Targeted advertising Technology Computer processing of body language Emotion recognition in conversation Gesture recognition List of facial expression databases Sentiment analysis Key people Ray Birdwhistell Charles Darwin Paul Ekman Related Animal communication Behavioral communication Aggressive Assertive Passive Passive-aggressive Impression management Meta-communication Monastic sign lexicons Verbal communication Non-verbal language Sign language Tactile signing Tadoma Art and literature Mime Mimoplastic art Subtext

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