Charles Robert Darwin FRS (12 February 1809 – 19 April 1882) was an English naturalist who realised that all species of life have descended over time from common ancestors, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection. He published his theory with compelling evidence for evolution in his 1859 book On the Origin of Species. 

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. Studies at the University of Cambridge encouraged his passion for natural science. His five-year voyage on HMS Beagle established him as an eminent geologist whose observations and theories supported Charles Lyell's uniformitarian ideas, and publication of his journal of the voyage made him famous as a popular author. Puzzled by the geographical distribution of wildlife and fossils he collected on the voyage, Darwin investigated the transmutation of species and conceived his theory of natural selection in 1838. 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 which described the same idea, prompting immediate joint publication of both of their theories. Darwin's work established evolutionary descent with modification as the dominant scientific explanation of diversification in nature. 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. His research on plants was published in a series of books, and in his final book, he examined earthworms and their effect on soil. 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 state funeral, and was buried in Westminster Abbey, close to John Herschel and Isaac Newton.

Charles Robert Darwin was born in Shrewsbury, Shropshire, England on 12 February 1809 at his family home, the Mount. He was the fifth of six children of wealthy society doctor and financier Robert Darwin, and Susannah Darwin (née Wedgwood). Robert Darwin had baby Charles baptised 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 Erasmus 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 University of Edinburgh Medical School with his brother Erasmus in October 1825. He found lectures dull and surgery distressing, so neglected his studies. He learned taxidermy from John Edmonstone, a freed black slave who had accompanied Charles Waterton in the South American rainforest. In Darwin's second year he joined the Plinian Society, a student natural history group whose debates strayed into radical materialism. He assisted Robert Edmund Grant's investigations of the anatomy and life cycle of marine invertebrates in the Firth of Forth, and in March 1827 presented at the Plinian his own discovery that black spores found in oyster shells were the eggs of a skate leech. Darwin was rather bored by Robert Jameson's natural history course which covered geology including the debate between Neptunism and Plutonism. He learned classification of plants, and assisted with work on the collections of the University Museum, one of the largest museums in Europe at the time.

This neglect of medical studies annoyed his father, who shrewdly sent him to Christ's College, Cambridge, for a Bachelor of Arts degree as the first step towards becoming an Anglican parson. Darwin began there in January 1828, but preferred riding and shooting to studying. His cousin William Darwin Fox introduced him to the popular craze for beetle collecting which he pursued zealously, getting some of his finds published in Stevens' Illustrations of British entomology. He became a close friend and follower of botany professor John Stevens Henslow and met other leading naturalists who saw scientific work as religious natural theology, becoming known to these dons 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 William Paley's Evidences of Christianity. In his final examination in January 1831 Darwin did well, coming tenth out of a pass list of 178. Darwin had to stay at Cambridge until June. He studied Paley's Natural Theology 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 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. 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 went with him in the summer for a fortnight to map strata in Wales. After a week with student friends at Barmouth, 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 Robert FitzRoy, 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. His father objected to the planned two-year voyage, regarding it as a waste of time, but was persuaded by his brother-in-law, Josiah Wedgwood, to agree to his son's participation.

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 surveyed and charted coasts. 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 his journal for his family. 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 natural history. In Cape Town Darwin and FitzRoy met John Herschel.

When the Beagle reached Falmouth, Cornwall, on 2 October 1836, Darwin was already a celebrity in scientific circles as in December 1835 Henslow had fostered his former pupil's reputation by giving selected naturalists a pamphlet of Darwin's geological letters. Darwin visited his home in Shrewsbury and saw relatives, then hurried to Cambridge 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 gentleman scientist, and an excited Darwin went round the London institutions being fêted and seeking experts to describe the collections. 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, 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 organise work on his collections and rewrite his Journal. He wrote his first paper, showing that the South American landmass was slowly rising, and with Lyell's enthusiastic backing 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 Galapagos 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. John Herschel's letter on the "mystery of mysteries" of new species was widely discussed, with explanations sought in laws of nature, not ad hoc miracles. Darwin stayed with his freethinking brother Erasmus, part of this Whig circle and close friend of writer Harriet Martineau who promoted Malthusianism underlying 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. By mid-March, Darwin was speculating in his Red Notebook on the possibility that "one species does change into another". He sketched branching descent, then agenealogical branching of a single evolutionary tree, in which "It is absurd to talk of one animal being higher than another", discarding Lamarck's independent lineages progressing to higher forms.

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 £73,000 in present day terms. He stretched the funding to include his planned books on geology, and agreed 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.
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 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 Jos 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.

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. He included mankind in his speculations from the outset, and on seeing an orangutan in the zoo on 28 March 1838 noted its child-like 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. On 23 June he took a break and went "geologising" in Scotland. 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 ... better than a dog anyhow", against points such as "less money for books" and "terrible loss of time." 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. 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 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.

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.

Darwin now had the framework of his theory of natural selection "by which to work", as his "prime hobby". His research included animal husbandry and extensive experiments with plants, finding evidence that species were not fixed and investigating many detailed ideas to refine and substantiate his theory. When FitzRoy's Narrative was published in May 1839, Darwin's Journal and Remarks was such a success as the third volume that later that year it was published on its own. 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 September. On 11 January 1844 Darwin mentioned his theorising to the botanist Joseph Dalton Hooker. 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 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, and after a long series of crises she died.

In eight years of work on barnacles (Cirripedia), 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. 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".

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 firmly against evolution. Lyell was intrigued by Darwin's speculations without realising their extent. When he read a paper by Alfred Russel Wallace 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, obtaining information and specimens from naturalists worldwide including Wallace who was working in Borneo. The American botanist Asa Gray 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 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 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 scarlet fever, and he put matters in the hands of Lyell and Hooker. They 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; however, Darwin's baby son died of the scarlet fever and he was too distraught to attend.

His On the Origin of Species proved unexpectedly popular went on sale to booksellers on 22 November 1859. The book aroused international interest, with less controversy than had greeted the popular Vestiges of Creation. 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". 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. 

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 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 Darwinismus incorporating Lamarckism and Goethe's idealism. Wallace remained supportive, though he increasingly turned to Spiritualism.

The Variation of Animals and Plants under Domestication of 1868 was the first part of Darwin's 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 characteristics, while emphasising that humans are all one species. 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. 

He died at Down House on 19 April 1882. He had expected to be buried in St Mary's churchyard at Downe, but at the request of Darwin's colleagues, William Spottiswoode (President of the Royal Society) arranged for Darwin to be given a state funeral and buried in Westminster Abbey, close to John Herschel and Isaac Newton. Only five non-royal personages were granted that honour of a UK state funeral during the 19th century.