Darwin

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Charles Darwin Finch Darwin's finches Galpagos Islands HMS Beagle Species Natural selection Genetics Gregor Mendel 1 24 30 37 55 62 74 88 104

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Charles Darwin

Charles Darwin
Charles Darwin

Darwin, aged 45 in 1854, by then working towards publication of On the Origin of Species Born Charles Robert Darwin 12 February 1809 The Mount, Shrewsbury, Shropshire, United Kingdom 19 April 1882 (aged73) Down House, Downe, Kent, United Kingdom England British British Naturalist tertiary education: University of Edinburgh Medical School (medicine) University of Cambridge (ordinary Bachelor of Arts) professional institution: Geological Society of London John Stevens Henslow Adam Sedgwick The Voyage of the Beagle On the Origin of Species evolution by natural selection, common descent Alexander von Humboldt John Herschel Charles Lyell Joseph Dalton Hooker Thomas Henry Huxley George Romanes Ernst Haeckel Sir John Lubbock

Died

Residence Citizenship Nationality Fields Institutions

Academic advisors

Knownfor

Influences

Influenced

Charles Darwin

2
Notable awards Royal Medal (1853) Wollaston Medal (1859) Copley Medal (1864) Emma Darwin (married 1839) 10 children (see list) Signature

Spouse Children

Charles Robert Darwin, FRS (12February 1809 19April 1882) was an English naturalist.[I] He established that all species of life have descended over time from common ancestors,[1] and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection, in which the struggle for existence has a similar effect to the artificial selection involved in selective breeding.[] Darwin published his theory of evolution with compelling evidence in his 1859 book On the Origin of Species, overcoming scientific rejection of earlier concepts of transmutation of species.[2][3] By the 1870s the scientific community and much of the general public had accepted evolution as a fact. However, many favoured competing explanations 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.[][] In modified form, Darwin's scientific discovery is the unifying theory of the life sciences, explaining the diversity of life.[4][5] 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 HMSBeagle 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.[6] Puzzled by the geographical distribution of wildlife and fossils he collected on the voyage, Darwin began detailed investigations and in 1838 conceived his theory of natural selection.[7] Although he discussed his ideas with several naturalists, he needed time for extensive research and his geological work had priority.[8] 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.[9] 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.[10] In recognition of Darwin's pre-eminence as a scientist, he was honoured with a major ceremonial funeral and buried in Westminster Abbey, close to John Herschel and Isaac Newton.[11] Darwin has been described as one of the most influential figures in human history.[12][13]

Charles Darwin

Biography
Early life and education
Charles Robert Darwin was born in Shrewsbury, Shropshire, England on 12 February 1809 at his family home, The Mount.[14] He was the fifth of six children of wealthy society doctor and financier Robert Darwin, and Susannah Darwin (ne Wedgwood). He was the grandson of two prominent abolitionists: Erasmus Darwin on his father's side, and of Josiah Wedgwood on his mother's side. Both families were largely Unitarian, though the Wedgwoods were adopting Anglicanism. Robert Darwin, himself quietly a freethinker, had baby Charles baptised in November 1809 in the Anglican St Chad's Church, Shrewsbury, 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, at the time the best medical school in the UK, 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, and often sat with this "very pleasant and intelligent man".[]

The seven-year-old Charles Darwin in 1816.

In Darwin's second year he joined the Plinian Society, a student natural history group whose debates strayed into radical materialism. 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.[15] 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.[16] 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. As Darwin was unqualified for the Tripos, he joined the ordinary degree course in January 1828.[17] He preferred riding and shooting to studying. His cousin William Darwin Fox introduced him to the popular craze for beetle collecting; Darwin pursued this 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 his own exams drew near, Darwin focused on his studies and was delighted by the language and logic of William Paley's Evidences of Christianity.[18] In his final examination in January 1831 Darwin did well, coming tenth out of 178 candidates for the ordinary degree.[19] 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 travelled with him in the

Charles Darwin summer for a fortnight, in order to map strata in Wales.[]

Voyage of the Beagle

After a week with student friends at Barmouth, Darwin returned home on 29 August to find a letter from Henslow proposing him as a suitable (if unfinished) gentleman naturalist for a self-funded supernumerary place on HMSBeagle with captain Robert FitzRoy, more as a companion than a mere collector. The ship was to leave The voyage of the Beagle in four weeks on an expedition to chart [20] 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, to agree to (and fund) his son's participation.[21] 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 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.[22] 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 in a calm spell.[][] On their first stop ashore at St. Jago, 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,[II] and Darwin saw things Lyell's way, theorising and thinking of writing a book on geology.[23] When they reached Brazil Darwin was delighted by the tropical forest,[24] but detested the sight of slavery.[25] The survey continued to the south in Patagonia. They stopped at Baha 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 identified the little known Megatherium by a tooth and its association with bony armour which had at first seemed to him like a giant version of the armour on local armadillos. The finds brought great interest when they reached England.[26][] 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.[27][28] Further south he saw stepped plains of shingle and seashells as raised beaches 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.[29][]

Charles Darwin

5 Three Fuegians on board, who had been seized during the first Beagle voyage and had spent a year in England, were taken back to Tierra del Fuego as missionaries. Darwin found them friendly and civilised, yet their relatives seemed "miserable, degraded savages", as different as wild from domesticated animals.[30] 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.[31] 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.[32]

As HMS Beagle surveyed the coasts of South America, Darwin theorised about geology and extinction of giant mammals.

Darwin experienced an earthquake in Chile 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.[33][] On the geologically new Galpagos 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.[][34] 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", and noted their depletion by European settlement.[35] The Beagle 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 natural history.[36] In Cape Town 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".[37] He later wrote that such facts "seemed to me to throw some light on the origin of species".[38]

Charles Darwin

Inception of Darwin's evolutionary theory

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.[39] 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 fted 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.[40] 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 While still a young man, Charles Darwin joined the scientific elite 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.[41][] These extinct creatures were related to living species in South America.[42] In mid-December Darwin took lodgings in Cambridge to organise work on his collections and rewrite his Journal.[43] 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.[44] 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,[45] who described God as a programmer of laws. 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. Transmutation was anathema to Anglicans defending social order,[46] 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 Galpagos mockingbirds from different islands were separate species, not just varieties, and what Darwin had thought was a "wren" was also in the finch group. Darwin had not labelled the finches by island, but from the notes of others on the Beagle, including FitzRoy, he allocated species to islands.[47] The two rheas were also distinct species, and on 14 March Darwin announced how their distribution changed going southwards.[48]

Charles Darwin

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 Macrauchenia which resembled a giant guanaco. His thoughts on lifespan, asexual reproduction and sexual reproduction developed in his "B" notebook around mid-July on to variation in offspring "to adapt & alter the race to changing world" explaining the Galpagos tortoises, mockingbirds and rheas. He sketched branching descent, then a genealogical 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.[49]

Overwork, illness, and marriage

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 77,000 in 2011.[50] He stretched the funding to include his planned books on geology, and agreed unrealistic dates with the publisher.[51] As the Victorian era began, Darwin pressed on with writing his Journal, and in August 1837 began correcting printer's proofs.[52]

In mid-July 1837 Darwin started his "B" notebook on Transmutation of Species, and on page 36 wrote "I think" above his first evolutionary tree.

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.[53] 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.[54] 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 pigeon fanciers.[][55] Over time his research drew on information from his relatives and children, the family butler, neighbours, colonists and former shipmates.[56] 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.[57]

Charles Darwin

8 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 little success.[58] 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.[59]

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."[60] 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.[61] 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, and 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 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.[][62] 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",[63] thinking this comparison "a beautiful part of my theory".[64] 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 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

Darwin chose to marry his cousin, Emma Wedgwood.

Charles Darwin 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.[65] 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.[66]

Geology books, Barnacles, evolutionary research

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.[] 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.[67] 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".[68] 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.[69] To escape the pressures of London, the Darwin in 1842 with his eldest son, William family moved to rural Down House in September.[70] On 11 January Erasmus Darwin 1844 Darwin mentioned his theorising to the botanist Joseph Dalton Hooker, writing with melodramatic humour "it is like confessing a murder".[71][72] 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."[73] By July, Darwin had expanded his "sketch" into a 230-page "Essay", to be expanded with his research results if he died prematurely.[75] 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.[76][77] Darwin completed his third geological book in 1846. He now renewed Darwin's "sandwalk" at Down House was his [74] a fascination and expertise in marine invertebrates, dating back to his usual "Thinking Path". 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.[78] 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.[79] 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.[80] 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.[81]

Charles Darwin 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.[82] 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".[83]

10

Publication of the theory of natural selection

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 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, 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."[85]

Charles Darwin, aged 46 in 1855, by then working towards publication of his theory of natural selection. He wrote to Hooker about this portrait, "if I really have as bad an expression, as my photograph gives me, how I can have one single friend is [84] surprising."

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,[86][87] 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 Lyell and Hooker. After some discussion, 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.[88] 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.[89] 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."[90] 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.[91] 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.[92] In the book, Darwin set out "one long argument" of detailed observations, inferences and consideration of anticipated objections.[93] His only allusion to human evolution was the understatement that "light will be thrown on the origin of man and his history".[94] His theory is simply stated in the introduction:

Charles Darwin 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.[95] He put a strong case for common descent, and at the end of the book 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.[96] 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, and 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.[97]

11

Responses to publication
The book aroused international interest, with less controversy than had greeted the popular Vestiges of the Natural History of Creation.[98] 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.[99] Darwin had only said "Light will be thrown on the origin of man",[100] but the first review claimed it made a creed of the "men from monkeys" idea from Vestiges.[101] Amongst early favourable responses, Huxley's reviews swiped at Richard Owen, leader of the scientific establishment Huxley was trying to overthrow.[102] In April, Owen's review attacked Darwin's friends and condescendingly dismissed his ideas, angering Darwin,[103] but Owen and others began to promote ideas of supernaturally guided evolution.[104]

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, with its ideas including higher criticism 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".[105] 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.[][106]

During the Darwin family's 1868 holiday in her Isle of Wight cottage, Julia Margaret Cameron took portraits showing the bushy beard Darwin grew between 1862 and 1866.

Charles Darwin

12

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.[107]

An 1871 caricature following publication of The Descent of Man was typical of many showing Darwin with an ape body, identifying him in popular culture as the leading author [] of evolutionary theory.

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.[108] 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.[109] 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.[110] Darwin's theory also resonated with various movements at the time[III] and became a key fixture of popular culture.[IV] 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.[]

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Descent of Man, sexual selection, and botany

More detailed articles cover Darwin's life from Orchids to Variation, from Descent of Man to Emotions and from 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 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 By 1878, an increasingly famous Darwin had suffered years of illness. 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.[111] Admiring visitors included Ernst Haeckel, a zealous proponent of Darwinismus incorporating Lamarckism and Goethe's idealism.[112] Wallace remained supportive, though he increasingly turned to Spiritualism.[113] 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.[114] 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.[115] 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 Punch's almanac for 1882, published evolution of human psychology and its continuity with the behaviour of animals. shortly before Darwin's death, Both books proved very popular, and Darwin was impressed by the general depicts him amidst evolution from chaos to Victorian gentleman with assent with which his views had been received, remarking that "everybody is [116] the title Man Is But A Worm. 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 systemwith all these exalted powersMan still bears in his bodily frame the indelible stamp of his lowly origin."[117] 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. In his last book he returned to The Formation of Vegetable Mould through the Action of Worms.

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Death and legacy

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".[118] 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", then while she rested, he repeatedly told Henrietta and Francis "It's almost worth while to be sick to be nursed by you".[119] 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 buried in Westminster Abbey, close to John Herschel and Isaac Newton.[11][120] Darwin had convinced most scientists that evolution as descent with modification was correct, and he was regarded as a great scientist who had revolutionised ideas. Though few agreed with his view that "natural selection has been the main but not the exclusive means of modification", he was honoured in June 1909 by more than 400 officials and scientists from across the world who met in Cambridge to commemorate his centenary and the fiftieth anniversary of On the Origin of Species.[121] During this period, which has been called "the eclipse of Darwinism", scientists proposed various alternative evolutionary mechanisms which eventually proved untenable. The development of the modern evolutionary synthesis from the 1930s to the 1950s, incorporating natural selection with population genetics and Mendelian genetics, brought broad scientific consensus that natural selection was the basic mechanism of evolution. This synthesis set the frame of reference for modern debates and refinements of the theory.[]

Man dressed as Charles Darwin during Lyme Regis Fossil Festival.

Children
Darwin's children William Erasmus Darwin Anne Elizabeth Darwin Mary Eleanor Darwin (27 December 1839 1914) (2 March 1841 23 April 1851) (23 September 1842 16 October 1842)

Henrietta Emma "Etty" Darwin (25 September 1843 1929) George Howard Darwin Elizabeth "Bessy" Darwin Francis Darwin Leonard Darwin Horace Darwin Charles Waring Darwin (9 July 1845 7 December 1912) (8 July 1847 1926) (16 August 1848 19 September 1925) (15 January 1850 26 March 1943) (13 May 1851 29 September 1928) (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 this topic in his writings, contrasting it with the advantages of crossing

Charles Darwin amongst many organisms.[122] Despite his fears, most of the surviving children and many of their descendants went on to have distinguished careers (see Darwin-Wedgwood family).[123] Of his surviving children, George, Francis and Horace became Fellows of the Royal Society,[124] distinguished as astronomer,[125] botanist and civil engineer, respectively. His son Leonard went on to be a soldier, politician, economist, eugenicist and mentor of the statistician and evolutionary biologist Ronald Fisher.[126]

15

Views and opinions

Religious views
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 doubt the literal truth of the Bible.[18] 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 the 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 related the antlion found near kangaroos to distinct "periods of Creation".[] By his return he was critical of the Bible as history, and wondered why all religions should not be equally valid.[] In the next few years, while intensively speculating on geology and transmutation of species, he gave much thought to religion and openly discussed this with his wife Emma, whose beliefs also 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,[127] 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.[] He still viewed organisms as perfectly adapted, and On the Origin of Species reflects theological views. 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.[128][129]
In 1851 Darwin was devastated when his daughter Annie died. By then his faith in Christianity had dwindled, and he had stopped going to [] church.

Darwin remained close friends with the vicar of Downe, John Innes, and continued to play a leading part in the parish work of the church,[130] but from around 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"[131][132] 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."[][131] 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.[133]

Human society
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 Antoinette Brown Blackwell in The Sexes Throughout Nature.[] 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.[134] Darwin was

Charles Darwin strongly against slavery, against "ranking the so-called races of man as distinct species", and against ill-treatment of native people.[135][VI] 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.[136] Francis Galton named this field of study "eugenics" in 1883.[V]

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Evolutionary social movements

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. 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.[137] 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.[138] 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 racist ideas of human development. Writers used natural Caricature from 1871 Vanity Fair selection to argue for various, often contradictory, ideologies such as laissez-faire dog-eat dog capitalism, racism, warfare, 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 co-operation over struggle within a species.[139] Darwin himself insisted that social policy should not simply be guided by concepts of struggle and selection in nature.[140] 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 sterilization laws, followed by several other countries. Subsequently, Nazi eugenics brought the field into disrepute.[V] 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.[141][137]

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Commemoration
During Darwin's lifetime, many geographical features were given his name. An expanse of water adjoining the Beagle Channel was named Darwin Sound by Robert FitzRoy 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 glacier caused a large wave that would have swept away their boats,[142] and the nearby Mount Darwin in the Andes was named in celebration of Darwin's 25th birthday.[143] When the Beagle was surveying Australia in 1839, Darwin's friend John Lort Stokes sighted a natural harbour which the ship's captain Wickham named Port Darwin: a nearby settlement was renamed Darwin in 1911, and it became the capital city of Australia's Northern Territory.[] More than 120 species and nine genera have been named after Darwin.[144] In one example, the group of tanagers related to those Darwin found in the Galpagos Islands became popularly known as "Darwin's finches" in 1947, fostering inaccurate legends about their significance to his work.[145]

Darwin's work has continued to be celebrated by numerous publications and events. The Linnean Society of London has commemorated Darwin's achievements by the award of the DarwinWallace Medal since 1908. Darwin Day has become an annual celebration, and in 2009 worldwide events were arranged for the bicentenary of Darwin's birth and the 150th anniversary of the publication of On the Origin of Species.[146] Darwin has been commemorated in the UK, with his portrait printed on the reverse of 10 banknotes printed along with a hummingbird and HMS Beagle, issued by the Bank of England.[147] A life size seated statue of Darwin can be seen in the main hall of the Natural History Museum in London. .[148] A seated statue of Darwin stands in front of Shrewsbury Library, the building that used to house Shrewsbury School, which Darwin attended as a boy. Darwin College, a postgraduate college at Cambridge University, is named after the Darwin family.

In 1881 Darwin was an eminent figure, still working on his contributions to evolutionary thought that had an enormous effect on many fields of science.

Works
Darwin was a prolific writer. Even without 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.[149][150]

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Notes
I.

Darwin was eminent as a naturalist, geologist, biologist, and author; after working as a physician's assistant and two years as a medical student was educated as a clergyman; and was trained in taxidermy.[]
II.

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) See, for example, WILLA volume 4, Charlotte Perkins Gilman and the Feminization of Education [151] 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."
III. IV. ^

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.
V.

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 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 (Updated 17 December 1996). "the creation of genetic identity" [152]. SEHR 5 (Supplement: Cultural and Technological Incubations of Fascism). Retrieved 11 November 2008 Edwards, A. W. F. (1 April 2000). "The Genetical Theory of Natural Selection" [153]. Genetics 154 (April 2000). pp.14191426. PMC1461012 [154]. PMID10747041 [155]. Retrieved 11 November 2008 Wilkins, John. "Evolving Thoughts: Darwin and the Holocaust 3: eugenics" [156]. Retrieved 11 November 2008.)
VI.

Darwin did not share the then common view that other races are inferior, and said of his taxidermy tutor John Edmonstone, a freed black slave, "I used often to sit with him, for he was a very pleasant and intelligent man".[] Early in the Beagle voyage he nearly lost his position on the ship when he criticised FitzRoy's defence and praise of slavery. (Darwin 1958, p.74 [157]) 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 [158] ) 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, 207208 [159]) In the Descent of Man he mentioned the Fuegians and Edmonstone when arguing against "ranking the so-called races of man as distinct species".[160] 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 [161])

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Citations
[4] The Complete Works of Darwin Online Biography. (http:/ / darwin-online. org. uk/ biography. html) darwin-online.org.uk. Retrieved 2006-12-15 [5] As Darwinian scholar Joseph Carroll of the University of MissouriSt. 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 timeone 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." [45] proposes a move on Friday 3 March 1837, Darwin's Journal () backdated from August 1838 gives a date of 6 March 1837 [50] UK CPI inflation numbers based on data available from Gregory Clark (2013), " What Were the British Earnings and Prices Then? (New Series) (http:/ / www. measuringworth. org/ ukearncpi/ )" MeasuringWorth. [54] . [84] Darwin Online: Photograph of Charles Darwin by Maull and Polyblank for the Literary and Scientific Portrait Club (1855) (http:/ / darwin-online. org. uk/ EditorialIntroductions/ vanWyhe_MaullandPolyblankPhoto. html), John van Wyhe, December 2006 [86] Ball, P. (2011). Shipping timetables debunk Darwin plagiarism accusations: Evidence challenges claims that Charles Darwin stole ideas from Alfred Russel Wallace. Nature. online (http:/ / www. nature. com/ news/ shipping-timetables-debunk-darwin-plagiarism-accusations-1. 9613) [87] J. van Wyhe and K. Rookmaaker. (2012). A new theory to explain the receipt of Wallace's Ternate Essay by Darwin in 1858. Biological Journal of the Linnean Society10.1111/j.1095-8312.2011.01808.x [97] , [112] Darwin Correspondence Project: Introduction to the Correspondence of Charles Darwin, Volume 14. (http:/ / www. darwinproject. ac. uk/ correspondence-volume-14) Cambridge University Press. Retrieved 25 June 2012 [113] . [122] . [126] Edwards, A. W. F. 2004. Darwin, Leonard (18501943). In: Oxford Dictionary of National Biography, Oxford University Press. [131] Letter 12041 (http:/ / www. darwinproject. ac. uk/ darwinletters/ calendar/ entry-12041. html) Darwin, C. R. to Fordyce, John, 7 May 1879 [132] Darwin's Complex loss of Faith (http:/ / www. guardian. co. uk/ commentisfree/ belief/ 2009/ sep/ 17/ darwin-evolution-religion) The Guardian 17 September 2009 [151] http:/ / scholar. lib. vt. edu/ ejournals/ old-WILLA/ fall95/ DeSimone. html [152] http:/ / www. stanford. edu/ group/ SHR/ 5-supp/ text/ thurtle. html [153] http:/ / www. genetics. org/ cgi/ content/ full/ 154/ 4/ 1419#The_Eclipse_of_Darwinism [154] http:/ / www. ncbi. nlm. nih. gov/ pmc/ articles/ PMC1461012 [155] http:/ / www. ncbi. nlm. nih. gov/ pubmed/ 10747041 [156] http:/ / scienceblogs. com/ evolvingthoughts/ 2006/ 09/ darwin_and_the_holocaust_3_eug_1. php [157] http:/ / darwin-online. org. uk/ content/ frameset?viewtype=text& itemID=F1497& pageseq=76 [158] http:/ / darwin-online. org. uk/ content/ frameset?viewtype=text& itemID=F1452. 1& pageseq=264 [159] http:/ / darwin-online. org. uk/ content/ frameset?itemID=F14& viewtype=text& pageseq=218 [161] http:/ / darwin-online. org. uk/ content/ frameset?viewtype=text& itemID=F14& pageseq=115

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Charles Darwin Darwin, Charles (1837). Notebook B: (Transmutation of species) (http://darwin-online.org.uk/content/ frameset?viewtype=side&itemID=CUL-DAR121.-&pageseq=1). Darwin Online. CUL-DAR121. Retrieved 20 December 2008. Darwin, Charles (1839). Narrative of the surveying voyages of His Majesty's Ships Adventure and Beagle between the years 1826 and 1836, describing their examination of the southern shores of South America, and the Beagle's circumnavigation of the globe. Journal and remarks. 18321836. (http://darwin-online.org.uk/content/ frameset?itemID=F10.3&viewtype=text&pageseq=1) III. London: Henry Colburn. Retrieved 24 October 2008. Darwin, Charles (1842). "Pencil Sketch of 1842" (http://darwin-online.org.uk/content/ frameset?viewtype=text&itemID=F1556&pageseq=33). In Darwin, Francis. The foundations of The origin of species: Two essays written in 1842 and 1844. (http://darwin-online.org.uk/content/frameset?itemID=F1556& viewtype=text&pageseq=1). Cambridge University Press (published 1909). ISBN0-548-79998-9. Darwin, Charles (1845). Journal of researches into the natural history and geology of the countries visited during the voyage of H.M.S. Beagle round the world, under the Command of Capt. Fitz Roy, R.N. 2d edition (http:// darwin-online.org.uk/content/frameset?itemID=F20&viewtype=text&pageseq=1). London: John Murray. Retrieved 24 October 2008. Darwin, Charles; Wallace, Alfred Russel (1858). "On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection". Journal of the Proceedings of the Linnean Society of London. Zoology 3 3 (9): 4650. doi: 10.1111/j.1096-3642.1858.tb02500.x (http://dx.doi.org/10. 1111/j.1096-3642.1858.tb02500.x). Darwin, Charles (1859). On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (http://darwin-online.org.uk/content/frameset?itemID=F373&viewtype=text& pageseq=1) (1st ed.). London: John Murray. ISBN1-4353-9386-4. Retrieved 24 October 2008. Darwin, Charles (1868). The variation of animals and plants under domestication (http://darwin-online.org.uk/ content/frameset?itemID=F880.1&viewtype=text&pageseq=1). London: John Murray. ISBN1-4191-8660-4. Retrieved 1 November 2008. Darwin, Charles (1871). The Descent of Man, and Selection in Relation to Sex (http://darwin-online.org.uk/ EditorialIntroductions/Freeman_TheDescentofMan.html) (1st ed.). London: John Murray. ISBN0-8014-2085-7. Retrieved 24 October 2008. Darwin, Charles (1872). The Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (http://darwin-online.org.uk/content/frameset?itemID=F391&viewtype=text& pageseq=1) (6th ed.). London: John Murray. ISBN1-4353-9386-4. Retrieved 1 November 2009. Darwin, Charles (1887). Darwin, Francis, ed. The life and letters of Charles Darwin, including an autobiographical chapter (http://darwin-online.org.uk/EditorialIntroductions/ Freeman_LifeandLettersandAutobiography.html). London: John Murray. ISBN0-404-08417-6. Retrieved 4 November 2008. Darwin, Charles (1958). Barlow, Nora, ed. The Autobiography of Charles Darwin 18091882. With the original omissions restored. Edited and with appendix and notes by his granddaughter Nora Barlow. London: Collins. |accessdate= requires |url= (help) Darwin, Charles (2006). "Journal" (http://darwin-online.org.uk/content/frameset?viewtype=side& itemID=CUL-DAR158.1-76&pageseq=1). In van Wyhe, John. Darwin's personal 'Journal' (18091881) (http:// darwin-online.org.uk/EditorialIntroductions/vanWyhe_JournalDAR158.html). Darwin Online. CUL-DAR158.176. Retrieved 20 December 2008. Desmond, Adrian; Moore, James (1991). Darwin. London: Michael Joseph, Penguin Group. ISBN0-7181-3430-3. Desmond, Adrian; Moore, James; Browne, Janet (2004). Oxford Dictionary of National Biography. Oxford, England: Oxford University Press. doi: 10.1093/ref:odnb/7176 (http://dx.doi.org/10.1093/ref:odnb/7176).

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Charles Darwin Dobzhansky, Theodosius (March 1973). "Nothing in Biology Makes Sense Except in the Light of Evolution" (http://www.2think.org/dobzhansky.shtml). The American Biology Teacher 35 (3): 125129. doi: 10.2307/4444260 (http://dx.doi.org/10.2307/4444260). Retrieved 4 November 2008. Eldredge, Niles (2006). "Confessions of a Darwinist" (http://www.vqronline.org/articles/2006/spring/ eldredge-confessions-darwinist/). The Virginia Quarterly Review (Spring 2006): 3253. Retrieved 4 November 2008. FitzRoy, Robert (1839). Voyages of the Adventure and Beagle, Volume II (http://darwin-online.org.uk/content/ frameset?itemID=F10.2&viewtype=text&pageseq=1). London: Henry Colburn. Retrieved 4 November 2008. Freeman, R. B. (1977). The Works of Charles Darwin: An Annotated Bibliographical Handlist (http:// darwin-online.org.uk/content/frameset?itemID=A1&viewtype=text&pageseq=1). Folkestone: Wm Dawson & Sons Ltd. ISBN0-208-01658-9. Retrieved 4 November 2008. Hart, Michael H. (2000). The 100: A Ranking of the Most Influential Persons in History. New York: Citadel. ISBN0-89104-175-3. Herbert, Sandra (1980). "The red notebook of Charles Darwin" (http://darwin-online.org.uk/content/ frameset?viewtype=text&itemID=F1583e&pageseq=1). Bulletin of the British Museum (Natural History). Historical Series (7 (24 April)): 1164. Retrieved 11 January 2009. Herbert, Sandra (1991). "Charles Darwin as a prospective geological author" (http://darwin-online.org.uk/ content/frameset?viewtype=text&itemID=A342&pageseq=1). British Journal for the History of Science 24 (24): 159192. doi: 10.1017/S0007087400027060 (http://dx.doi.org/10.1017/S0007087400027060). Retrieved 24 October 2008. Keynes, Richard (2000). Charles Darwin's zoology notes & specimen lists from H.M.S. Beagle. (http:// darwin-online.org.uk/content/frameset?itemID=F1840&viewtype=text&pageseq=1). Cambridge University Press. ISBN0-521-46569-9. Retrieved 22 November 2008. Keynes, Richard (2001). Charles Darwin's Beagle Diary (http://darwin-online.org.uk/content/ frameset?itemID=F1925&viewtype=text&pageseq=1). Cambridge University Press. ISBN0-521-23503-0. Retrieved 24 October 2008. Kotzin, Daniel (2004). "Point-Counterpoint: Social Darwinism" (http://caho-test.cc.columbia.edu/pcp/14008. html). Columbia American History Online. Retrieved 22 November 2008. Larson, Edward J. (2004). Evolution: The Remarkable History of a Scientific Theory. Modern Library. ISBN0-679-64288-9. Leff, David (2000). "AboutDarwin.com" (http://www.aboutdarwin.com/index.html) (20002008 ed.). Retrieved 30 December 2008. Leifchild (19 November 1859). "Review of 'Origin'" (http://darwin-online.org.uk/content/ frameset?viewtype=image&itemID=CUL-DAR226.1.8&pageseq=1). Athenaeum (1673). Retrieved 22 November 2008. Miles, Sara Joan (2001). "Charles Darwin and Asa Gray Discuss Teleology and Design" (http://www.asa3.org/ ASA/PSCF/2001/PSCF9-01Miles.html). Perspectives on Science and Christian Faith 53: 196201. Retrieved 22 November 2008. Moore, James (2005). "Darwin A 'Devil's Chaplain'?" (http://speakingoffaith.publicradio.org/programs/ darwin/moore-devilschaplain.pdf) (PDF). American Public Media. Retrieved 22 November 2008. Moore, James (2006). "Evolution and Wonder Understanding Charles Darwin" (http://speakingoffaith. publicradio.org/programs/darwin/transcript.shtml). Speaking of Faith (Radio Program). American Public Media. Retrieved 22 November 2008. Owen, Richard (1840). Darwin, C. R., ed. Fossil Mammalia Part 1. The zoology of the voyage of H.M.S. Beagle. London: Smith Elder and Co. Paul, Diane B. (2003). "Darwin, social Darwinism and eugenics". In Hodge, Jonathan; Radick, Gregory. The Cambridge Companion to Darwin. Cambridge University Press. pp.214239. ISBN0-521-77730-5.

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Charles Darwin Smith, Charles H. (1999). "Alfred Russel Wallace on Spiritualism, Man, and Evolution: An Analytical Essay" (http://www.wku.edu/~smithch/essays/ARWPAMPH.htm). Retrieved 7 December 2008. Sulloway, Frank J. (1982). "Darwin and His Finches: The Evolution of a Legend" (http://www.sulloway.org/ Finches.pdf) (PDF). Journal of the History of Biology 15 (1): 153. doi: 10.1007/BF00132004 (http://dx.doi. org/10.1007/BF00132004). Retrieved 9 December 2008. Sweet, William (2004). "Herbert Spencer" (http://www.iep.utm.edu/spencer/). Internet Encyclopedia of Philosophy. Retrieved 16 December 2008. Wilkins, John S. (1997). "Evolution and Philosophy: Does evolution make might right?" (http://www. talkorigins.org/faqs/evolphil/social.html). TalkOrigins Archive. Retrieved 22 November 2008. Wilkins, John S. (2008). "Darwin". In Tucker, Aviezer. A Companion to the Philosophy of History and Historiography. Blackwell Companions to Philosophy. Chichester: Wiley-Blackwell. pp.405415. ISBN1-4051-4908-6. van Wyhe, John (27 March 2007). "Mind the gap: Did Darwin avoid publishing his theory for many years?" (http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=A544&pageseq=1). Notes and Records of the Royal Society 61 (2): 177205. doi: 10.1098/rsnr.2006.0171 (http://dx.doi.org/10.1098/rsnr. 2006.0171). Retrieved 7 February 2008. van Wyhe, John (2008). "Charles Darwin: gentleman naturalist: A biographical sketch" (http://darwin-online. org.uk/darwin.html). Darwin Online. Retrieved 17 November 2008. van Wyhe, John (2008b). Darwin: The Story of the Man and His Theories of Evolution. London: Andre Deutsch Ltd (published 1 September 2008). ISBN0-233-00251-0. von Sydow, Momme (2005). "Darwin A Christian Undermining Christianity? On Self-Undermining Dynamics of Ideas Between Belief and Science" (http://replay.waybackmachine.org/20090326070105/http://www. psych.uni-goettingen.de/abt/1/sydow/von_Sydow_(2005)_Darwin_A_Christian_Undermining_Christianity. pdf). In Knight, David M.; Eddy, Matthew D. Science and Beliefs: From Natural Philosophy to Natural Science, 17001900. Burlington: Ashgate. pp.141156. ISBN0-7546-3996-7. Retrieved 16 December 2008. Yates, Simon (2003). "The Lady Hope Story: A Widespread Falsehood" (http://www.talkorigins.org/faqs/ hope.html). TalkOrigins Archive. Retrieved 15 December 2006.

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External links
Library resources
About Charles Darwin

Online books (http://tools.wmflabs.org/ftl/cgi-bin/ftl?st=viaf&su=27063124&library=OLBP) Resources in your library (http://tools.wmflabs.org/ftl/cgi-bin/ftl?st=viaf&su=27063124) Resources in other libraries (http://tools.wmflabs.org/ftl/cgi-bin/ftl?st=viaf&su=27063124&library=0CHOOSE0) By Charles Darwin

Online books (http://tools.wmflabs.org/ftl/cgi-bin/ftl?at=viaf&au=27063124&library=OLBP) Resources in your library (http://tools.wmflabs.org/ftl/cgi-bin/ftl?at=viaf&au=27063124) Resources in other libraries (http://tools.wmflabs.org/ftl/cgi-bin/ftl?at=viaf&au=27063124&library=0CHOOSE0)

The Complete Works of Charles Darwin Online Darwin Online (http://darwin-online.org.uk/); Darwin's publications, private papers and bibliography, supplementary works including biographies, obituaries and reviews Darwin Correspondence Project (http://www.darwinproject.ac.uk/) Full text and notes for complete correspondence to 1867, with summaries of all the rest Darwin-Hooker letters, images and text jointly published at Cambridge Digital Library (http://cudl.lib.cam. ac.uk/collections/darwinhooker) and Darwin Correspondence Project (http://www.darwinproject.ac.uk/ darwin-hooker-letters)

Charles Darwin Works by Charles Darwin (http://www.gutenberg.org/author/Charles+Darwin) at Project Gutenberg; public domain Darwin Manuscript Project (http://darwin.amnh.org/) Works by Charles Darwin in audio format (http://librivox.org/newcatalog/search.php?title=&author=charles+ darwin&status=all&action=Search) from LibriVox Video and radio clips (http://archives.cbc.ca/science_technology/natural_science/topics/3696/) Canadian Broadcasting Corporation Charles Darwin (http://www.dmoz.org/Science/Biology/History/People/Darwin,_Charles//) at the Open Directory Project Works by or about Charles Darwin (http://worldcat.org/identities/lccn-n78-95637) in libraries (WorldCat catalog) Archival material relating to Charles Darwin (http://www.nationalarchives.gov.uk/nra/searches/subjectView. asp?ID=P7461) listed at the UK National Archives A Pictorial Biography of Charles Darwin (http://www.thesecondevolution.com/darwin_intro.html) Mis-portrayal of Darwin as a Racist (http://www.rationalrevolution.net/articles/darwin_nazism.htm) Darwin's Volcano (http://www.stanford.edu/group/microdocs/darwinvolcano.html) a short video discussing Darwin and Agassiz' coral reef formation debate Chisholm, Hugh, ed. (1911). "Darwin, Charles Robert". Encyclopdia Britannica (11th ed.). Cambridge University Press. The life and times of Charles Darwin, an audio slideshow, The Guardian, Thursday 12 February 2009, (http:// www.guardian.co.uk/science/interactive/2009/feb/12/charles-darwin) (3 min 20 sec). Darwin's Brave New World (http://www.screenaustralia.gov.au/showcases/charlesdarwin/) 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 Richard Dawkins, David Suzuki, Jared Diamond A naturalist's voyage around the world (http://www.cnrs.fr/cw/dossiers/dosdarwinE/darwin.html) Account of the Beagle voyage using animation, in English from Centre national de la recherche scientifique Anonymous (1873). Cartoon portraits and biographical sketches of men of the day (http://en.wikisource.org/ wiki/Cartoon_portraits_and_biographical_sketches_of_men_of_the_day/C._R._Darwin,_F.R.S.). Illustrated by Waddy, Frederick. London: Tinsley Brothers. pp.67. Retrieved 28 December 2010. View books owned and annotated by Charles Darwin (http://biodiversitylibrary.org/collection/darwinlibrary) at the online Biodiversity Heritage Library.

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Finch

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Finch
Finch

Adult male Common Chaffinch (Fringilla coelebs) of the Fringillinae

Scientific classification Kingdom: Phylum: Class: Order: Superfamily: Family: Animalia Chordata Aves Passeriformes Passeroidea Fringillidae Vigors, 1825 Subfamilies Carduelinae Drepanidinae Euphoniinae Fringillinae The true finches are passerine birds in the family Fringillidae. They are predominantly seed-eating songbirds. Most are native to the Northern Hemisphere, but one subfamily is endemic to the Neotropics, one to the Hawaiian Islands, and one subfamily monotypic at genus level is found only in the Palaearctic. The scientific name Fringillidae comes from the Latin word fringilla for the Common Chaffinch (Fringilla coelebs) a member of that last subfamily which is common in Europe. Many birds in other families are also commonly called "finches", including some species in the very similar-looking waxbills or estrildid finches (family Estrildidae) of the Old World tropics and Australia; several groups of the bunting and American sparrow family (Emberizidae); and Darwin's finches of the Galapagos islands, which provided evidence of natural selection and are now recognized to be peculiar tanagers (Thraupidae).[1]

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Description
The smallest "classical" true finches are the Andean Siskin (Carduelis spinescens) at as little as 9.5cm (3.8in) and the Lesser Goldfinch (C. psaltria) at as little as 8g (0.28oz). The largest species is probably the Collared Grosbeak (Mycerobas affinis) at up to 24cm (9.4in) and 83g (2.9oz), although larger lengths, to 25.5cm (10.0in) in the Pine Grosbeak (Pinicola enucleator), and weights, to 86.1g (3.04oz) in the Evening Grosbeak (Coccothraustes vespertinus), have been recorded in species which are slightly smaller on average.[2][3] They typically have strong, stubby beaks, which in some species can be quite large; however, Hawaiian honeycreepers are famous for the wide range of bill shapes and sizes brought about by adaptive radiation. All true finches have 12 remiges and 9 primary rectrices. The basic plumage colour is brownish, sometimes greenish; many have considerable amounts of black, while white plumage is generally absent except as wing-bars or other signalling marks. Bright yellow and red carotenoid pigments are commonplace in this family, and thus blue structural Beak and tongue shapes of the Drepanidinae colours are rather rare, as the yellow pigments turn the blue color into green. Many, but by no means all true finches have strong sexual dichromatism, the females typically lacking the bright carotenoid markings of males.[1] Finches are typically inhabitants of well-wooded areas, but some can be found on mountains or even in deserts. They are primarily granivorous, but euphoniines include considerable amounts of arthropods and berries in their diet, and Hawaiian honeycreepers evolved to utilize a wide range of food sources, including nectar. The diet of Fringillidae nestlings includes a varying amount of small arthropods. True finches have a bouncing flight like most small passerines, alternating bouts of flapping with gliding on closed wings. Most sing well and several are commonly seen cagebirds; foremost among these is the domesticated Canary (Serinus canaria domestica). The nests are basket-shaped and usually built in trees, more rarely in bushes, between rocks or on similar substrate.[1]

Systematics and taxonomy

The taxonomic structure of the true finch family, Fringillidae, has been fairly disputed in the past, with some upranking the Hawaiian honeycreepers (Drepanidinae) as family Drepanididae and/or uniting the cardueline and fringilline finches as tribes (Carduelini and Fringillini) in one subfamily; the euphonious finches (Euphoniinae) were thought to be tanagers due to general similarity in appearance and mode of life until their real affinities were realized. In particular, North American authors have often merged the buntings and American sparrow family (Emberizidae) and sometimes the bulk of the nine-primaried oscines with the split-up Fringillidae as subfamilies of a single massive family. But the current understanding of Passeroidea phylogeny is better reflected in keeping the fundamental nine-primaried oscine clades as distinct families. However, Przewalski's "Rosefinch" (Urocynchramus pylzowi) is now classified as a distinct family, monotypic as to genus and species, and with no particularly close relatives among the Passeroidea.[4] Fossil remains of true finches are rare, and those that are known can mostly be assigned to extant genera at least. Like the other Passeroidea families, the true finches seem to be of roughly Middle Miocene origin, around 20-10 million years ago (Ma). An unidentifable finch fossil from the Messinian age, around 12 to 7.3 million years ago (Ma) during the Late Miocene subepoch, has been found at Polgrdi in Hungary.[5][6][7]

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Hawfinch (Coccothraustes coccothraustes), one of the Holarctic grosbeaks

Cassin's Finch (Haemorhous cassinii), an American rosefinch

Pallas' Rosefinch (Carpodacus roseus), a true rosefinch

Hooded Siskin (Carduelis (Spinus) magellanica)

Iiwi (Vestiaria coccinea), a Hawaiian honeycreeper

Male Violaceous Euphonia (Euphonia violacea)

Subfamilies and genera

The systematics of the cardueline finches are contentious. The layout presented here follows the recent decades' molecular phylogenetic studies, and takes into account the traditional splitting of the genus Carduelis. The exact position of several genera in the cardueline sequence is tentative.[8] Subfamily Fringillinae fringilline finches. Three species, which feed their young on insects and few if any seed Genus Fringilla Subfamily Carduelinae cardueline finches. A much larger group that contains several genera which feed their young mainly on seeds. This subfamily consists of several well-defined clades. Typical grosbeaks Genus Eophona Oriental grosbeaks Genus Coccothraustes Holarctic grosbeaks (polyphyletic?) Genus Mycerobas Mycerobas grosbeaks Bullfinches Genus Pinicola Pine Grosbeak Genus Pyrrhula bullfinches Arid-zone clade Genus Leucosticte mountain finches Genus N.N. Dark-breasted Rosefinch, "Carpodacus" nipalensis Genus Rhodopechys Crimson-winged Finch Genus Bucanetes Trumpeter Finch and Mongolian Finch Asian rosefinches Genus Carpodacus rosefinches (Genus in need of substantial revision. May consist of two or more genera; alternatively may include Haematospiza and Uragus) Genus Haematospiza Scarlet Finch Genus Uragus streaked rosefinches American rosefinches

Finch Genus Haemorhous Goldfinch-canary-crossbill clade (Carduelis and Serinus are polyphyletic and are probably best regarded as a collection of independent sub-groups or genera.) Genus Serinus sensu lato canaries, African seedeaters, serins and African siskins (Sub)Genus Serinus sensu stricto European Serin and relatives (possibly related to Spinus) (Sub)Genus Crithagra Afrotropical canaries, African seedeaters, citrils (probably only distantly related to Serinus) Genus Linurgus Oriole Finch (possibly basal to Crithagra) Genus Rhynchostruthus golden-winged grosbeaks (tentatively placed here) Genus "Serinus" thibetanus Tibetan Serin (probably related to Spinus and Serinus sensu stricto) Genus Carduelis sensu lato (Sub)Genus Carduelis sensu stricto European Goldfinch, Citril Finch and Corsican Finch (probably only distantly related to Spinus) (Sub)Genus Spinus siskins and American goldfinches (Sub)Genus Linaria linnets and twite (related to Spinus) (Sub)Genus Chloris greenfinches and Desert Finch (Sub)Genus Acanthis redpolls (related to Loxia) (Sub)Genus Loxia crossbills Carduelinae incertae sedis Genus Pyrrhoplectes Gold-naped Finch Genus Chaunoproctus Bonin Grosbeak (extinct: 1830s) Genus Callacanthis Spectacled Finch Genus Neospiza Sao Tom Grosbeak

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Subfamily Drepanidinae Hawaiian honeycreepers. Endemic to the Hawaiian Islands; formerly often treated as a separate family. Some 10-12 living genera, about 7 recently extinct Subfamily Euphoniinae Euphonious finches. Endemic to the Neotropics; formerly placed in Thraupidae. Genus Euphonia euphonias Genus Chlorophonia chlorophonias

Footnotes
[1] [2] [3] [4] [5] [8] Newton (1973), Clement et al. (1993) Finches and Sparrows by Peter Clement. Princeton University Press (1999). ISBN 978-0691048789. CRC Handbook of Avian Body Masses by John B. Dunning Jr. (Editor). CRC Press (1992), ISBN 978-0-8493-4258-5. Clement et al. (1993), Groth (2000), Jnsson & Fjelds (2006), Arnaiz-Villena et al. (2007) Hr et al. (2001), Mlkovsk (2002) Marten & Johnson (1986), Arnaiz-Villena et al. (1998, 2001, 2007, 2008)

References
Arnaiz-Villena, A.; lvarez-Tejado, M.; Ruiz-del-Valle, V.; Garca-de-la-Torre, C.; Varela, P.; Recio, M. J.; Ferre. S. & Martnez-Laso, J (1998). "Phylogeny and rapid Northern and Southern Hemisphere speciation of goldfinches during the Miocene and Pliocene Epochs". Cellular and Molecular Life Sciences 54 (9): 10311041. doi: 10.1007/s000180050230 (http://dx.doi.org/10.1007/s000180050230). PMID 9791543 (http://www. ncbi.nlm.nih.gov/pubmed/9791543).. Erratum: "Announcement/Addendum/Erratum". Cellular and Molecular Life Sciences (CMLS) 55: 148. 1999. doi: 10.1007/s000180050280 (http://dx.doi.org/10.1007/ s000180050280).

Finch Arnaiz-Villena, A.; Guilln, J.; Ruiz-del-Valle, V.; Lowy, E.; Zamora, J.; Varela, P.; Stefani, D.; Allende, L.M. (2001). "Phylogeography of crossbills, bullfinches, grosbeaks, and rosefinches". Cellular and Molecular Life Sciences 58 (8): 115966. doi: 10.1007/PL00000930 (http://dx.doi.org/10.1007/PL00000930). PMID 11529508 (http://www.ncbi.nlm.nih.gov/pubmed/11529508). Arnaiz-Villena, A.; Moscoso, J.; Ruiz-del-Valle, V.; Gonzalez, J.; Reguera, R.; Wink, M.; Serrano-Vela, J.I. (2007). "Bayesian phylogeny of Fringillinae birds: status of the singular African oriole finch Linurgus olivaceus and evolution and heterogeneity of the genus Carpodacus" (http://www.actazool.org/temp/ {C6BDA075-F92E-48AF-815F-98BED7C65FE5}.pdf). Acta Zoologica Sinica 53 (5): 826834. Arnaiz-Villena, A.; Moscoso, J.; Ruiz-del-Valle, V.; Gonzalez, J.; Reguera, R.; Ferri, A.; Wink, M. & Serrano-Vela, J.I. (2008). "Mitochondrial DNA Phylogenetic Definition of a Group of 'Arid-Zone' Carduelini Finches" (http://www.benthamscience.com/open/tooenij/articles/V001/1TOOENIJ.pdf). Open Ornithology Journal 1: 17. doi: 10.2174/1874453200801010001 (http://dx.doi.org/10.2174/1874453200801010001). Clement, Peter; Harris, Alan & Davis, John (1993): Finches and Sparrows: an identification guide. Christopher Helm, London. ISBN 0-7136-8017-2 Groth, J. (2000). "Molecular evidence for the systematic position of Urocynchramus pylzowi" (http://sora.unm. edu/node/26265). Auk 117 (3): 787792. doi: 10.1642/0004-8038(2000)117[0787:MEFTSP]2.0.CO;2 (http:// dx.doi.org/10.1642/0004-8038(2000)117[0787:MEFTSP]2.0.CO;2). JSTOR 4089604 (http://www.jstor. org/stable/4089604). Hr, Jnos; Kkay, Jzsef; Venczel, Mrton; Gl, Erika & Kessler, Eugn (2001). "Elzetes beszmol a felstrknyi "Gdr-kert" n. slnytani lelhelykomplex jravizsglatrl [A preliminary report on the revised investigation of the paleontological locality-complex "Gdr-kert" at Felstrkny, Northern Hungary)]" (http:// www.matramuzeum.hu/e107_plugins/docrep_menu/docrep.php?0.getdoc.178.7.). Folia Historico Naturalia Musei Matraensis (in Hungarian with English abstract) 25: 4164. Jnsson, Knud A. & Fjelds, Jon (2006). "A phylogenetic supertree of oscine passerine birds (Aves: Passeri)". Zool. Scripta 35 (2): 149186. doi: 10.1111/j.1463-6409.2006.00221.x (http://dx.doi.org/10.1111/j. 1463-6409.2006.00221.x). Marten, Jill A. & Johnson, Ned K. (1986). "Genetic relationships of North American cardueline finches" (http:// sora.unm.edu/sites/default/files/journals/condor/v088n04/p0409-p0420.pdf). Condor 88 (4): 409420. doi: 10.2307/1368266 (http://dx.doi.org/10.2307/1368266). Mlkovsk, Jir (2002): Cenozoic Birds of the World (Part 1: Europe) (http://www.nm.cz/download/ JML-18-2002-CBE.pdf). Ninox Press, Prague. Newton, Ian (1973): Finches (New Naturalist series). Taplinger Publishing. ISBN 0-8008-2720-1

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Further reading
Groth, J. G. 1994. A mitochondrial cytochrome b phylogeny of cardueline finches. Journal fr Ornithologie, 135: 31. Groth, J. G. 1998. Molecular phylogeny of the cardueline finches and Hawaiian honeycreepers. Ostrich, 69: 401. Klicka, J., K.P. Johnson, and S.M. Lanyon. 2000. New World nine-primaried oscine relationships: Constructing a mitochondrial DNA framework. Auk 117:321-336. Ryan, P.G., Wright, D., Oatley, G., Wakeling, J., Cohen, C., Nowell, T.L., Bowie, R.C.K., Ward, V. & Crowe, T.M. 2004. Systematics of Serinus canaries and the status of Cape and Yellow-crowned Canaries inferred from mtDNA and morphology. Ostrich 75:288-294. Treplin, S. 2006. Inference of phylogenetic relationships in passerine birds (Aves: Passeriformes) using new molecular markers. (Dissertation - available online) http://opus.kobv.de/ubp/volltexte/2006/1123/pdf/ treplin_diss.pdf.

Finch Yuri, T., and D. P. Mindell. 2002. Molecular phylogenetic analysis of Fringillidae, "New World nine-primaried oscines" (Aves: Passeriformes). Mol. Phylogen. Evol. 23:229-243.

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External links
Finch videos, photos and sounds (http://ibc.lynxeds.com/family/finches-fringillidae) on the Internet Bird Collection National Finch and Softbill Society (http://www.nfss.org) A organization promoting breeding

Darwin's finches

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Darwin's finches
Darwin's finches

Large Ground Finch, Medium Ground Finch Small Tree Finch, Green Warbler-Finch

Scientific classification Kingdom: Phylum: Class: Order: Family: Animalia Chordata Aves Passeriformes Thraupidae Genera Geospiza Camarhynchus/Platyspiza Certhidea Pinaroloxias Darwin's finches (also known as the Galpagos finches) are a group of about fifteen species of passerine birds.[1] They often are classified as the subfamily Geospizinae or tribe Geospizini. It is still not clear which bird family they belong to, but they are not related to the true finches. They were first collected by Charles Darwin on the Galpagos Islands during the second voyage of the Beagle. All are found only on the Galpagos Islands, except the Cocos Finch from Cocos Island. The term Darwin's Finches was first applied by Percy Lowe in 1936, and popularised in 1947 by David Lack in his book Darwin's Finches.[2][3] The birds vary in size from 10 to 20cm and weigh between 8 and 38grams. The smallest are the warbler-finches and the largest is the Vegetarian Finch. The most important differences between species are in the size and shape of their beaks, and the beaks are highly adapted to different food sources. The birds are all dull-coloured.

Darwin's theory
During the survey voyage of HMS Beagle, Darwin had no idea of the significance of the birds of the Galpagos. He had learned how to preserve bird specimens while at the University of Edinburgh and had been keen on shooting, but he had no expertise in ornithology and by this stage of the voyage concentrated mainly on geology.[4] In Galpagos he mostly left bird shooting to his servant Syms Covington.[5] Nonetheless, these birds were to play an important part in the inception of Darwin's theory of evolution by natural selection.

Darwin's finches On the Galpagos Islands and afterward, Darwin thought in terms of "centres of creation" and rejected ideas of transmutation of species.[6] From Henslow's teaching he was interested in geographical distribution of species, particularly links between species on oceanic islands and on nearby continents. On Chatham Island he recorded that a mockingbird was similar to those he had seen in Chile, and after finding a different one on Charles Island he carefully noted where mockingbirds had been caught.[4] In contrast he paid little attention to the finches. When examining his specimens on the way to Tahiti Darwin noted that all the mockingbirds on Charles Island were of one species, those from Albemarle of another, and those from James and Chatham Islands of a third species. As they sailed home about nine months later this, together with other facts including what he'd heard about Galpagos tortoises, made him wonder about the stability of species.[][7] Following his return from the voyage, Darwin presented the finches to the Geological Society of London at their meeting on 4 January 1837, along with other mammal and bird specimens he had collected. The bird specimens, including the finches, were given to John Gould, the famous English ornithologist, for identification. Gould set aside his paying work and at the next meeting on 10 January reported that birds from the Galpagos Islands which Darwin had thought were blackbirds, "gross-beaks" and finches were in fact "a series of ground Finches which are so peculiar [as to form] an entirely new group, containing 12 species." This story made the newspapers.[8][] Darwin had been in Cambridge at that time. In early March he met Gould again and for the first time got a full report on the findings, including the point that his Galpagos "wren" was another closely allied species of finch. The mockingbirds Darwin had labelled by island were separate species rather than just varieties. Gould found more species than Darwin had anticipated,[9] and concluded that 25 of the 26 land birds were new and distinct forms, found nowhere else in the world but closely allied to those found on the South American continent.[] Darwin now saw that if the finch species were confined to individual islands, like the mockingbirds, this would help to account for the number of species on the islands, and he sought information from others on the expedition. Specimens had also been collected by Captain Robert FitzRoy, FitzRoys steward Harry Fuller and Darwin's servant Covington, who had labelled them by island.[] From these, Darwin tried to reconstruct the locations where he had collected his own specimens. The conclusions supported his idea of the transmutation of species.[]

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Text from the Voyage of the Beagle

At the time he rewrote his diary for publication as Journal and Remarks (later The Voyage of the Beagle), he described Gould's findings on the number of birds, noting that "Although the species are thus peculiar to the archipelago, yet nearly all in their general structure, habits, colour of feathers, and even tone of voice, are strictly American".[10] In the first edition of The Voyage of the Beagle Darwin said that "It is very remarkable that a nearly perfect gradation of structure in this one group can be traced in the form of the beak, from one exceeding in dimensions that of the largest gros-beak, to another differing but little from that of a warbler".[11] In 1839 Darwin conceived of his theory of natural selection, and by the time of the second edition in 1845 Darwin had brought together his theory. He now added two closing sentences: "Seeing this gradation and diversity of structure in one small, intimately related group of birds, one might really fancy that from an original paucity of birds in this archipelago, one species had been taken and modified for different ends".[12][13] The remaining land-birds form a most singular group of finches, related to each other in the structure of their beaks, short tails, form of body and plumage: there are thirteen species, which Mr. Gould has divided into four subgroups. All these species are peculiar to this archipelago; and so is the whole group, with the exception of one species of the sub-group Cactornis, lately brought from Bow Island, in the Low Archipelago. Of Cactornis, the two species may be often seen climbing about the flowers of the great cactus-trees; but all the other species of this group of finches, mingled together in flocks, feed on the dry and sterile ground of the lower districts. The males of all, or certainly of the greater number, are jet black; and the females (with perhaps one or two exceptions) are brown. The most curious fact is the perfect gradation in the size of the beaks in the different species of Geospiza, from one as large as that of a hawfinch to that of a chaffinch, and (if

Darwin's finches Mr. Gould is right in including his sub-group, Certhidea, in the main group) even to that of a warbler. The largest beak in the genus Geospiza is shown in Fig. 1, and the smallest in Fig. 3; but instead of there being only one intermediate species, with a beak of the size shown in Fig. 2, there are no less than six species with insensibly graduated beaks. The beak of the sub-group Certhidea, is shown in Fig. 4. The beak of Cactornis is somewhat like that of a starling, and that of the fourth subgroup, Camarhynchus, is slightly parrot-shaped. Seeing this gradation and diversity of structure in one small, intimately related group of birds, one might really fancy that from an original paucity of birds in this archipelago, one species had been taken and modified for different ends. In a like manner it might be fancied that a bird originally a buzzard, had been induced here to undertake the office of the carrion-feeding Polybori of the American continent.[14]

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Text from the Origin of Species

Darwin discussed the divergence of species of birds in the Galpagos more explicitly in his chapter on geographical distribution in On the Origin of Species: The most striking and important fact for us in regard to the inhabitants of islands, is their affinity to those of the nearest mainland, without being actually the same species. [In] the Galapagos Archipelago... almost every product of the land and water bears the unmistakable stamp of the American continent. There are twenty-six land birds, and twenty-five of these are ranked by Mr. Gould as distinct species, supposed to have been created here; yet the close affinity of most of these birds to American species in every character, in their habits, gestures, and tones of voice, was manifest.... The naturalist, looking at the inhabitants of these volcanic islands in the Pacific, distant several hundred miles from the continent, yet feels that he is standing on American land. Why should this be so? why should the species which are supposed to have been created in the Galapagos Archipelago, and nowhere else, bear so plain a stamp of affinity to those created in America? There is nothing in the conditions of life, in the geological nature of the islands, in their height or climate, or in the proportions in which the several classes are associated together, which resembles closely the conditions of the South American coast: in fact there is a considerable dissimilarity in all these respects. On the other hand, there is a considerable degree of resemblance in the volcanic nature of the soil, in climate, height, and size of the islands, between the Galapagos and Cape de Verde Archipelagos: but what an entire and absolute difference in their inhabitants! The inhabitants of the Cape de Verde Islands are related to those of Africa, like those of the Galapagos to America. I believe this grand fact can receive no sort of explanation on the ordinary view of independent creation; whereas on the view here maintained, it is obvious that the Galapagos Islands would be likely to receive colonists, whether by occasional means of transport or by formerly continuous land, from America; and the Cape de Verde Islands from Africa; and that such colonists would be liable to modification;the principle of inheritance still betraying their original birthplace.[15]

Polymorphism in Darwin's finches

Whereas Darwin spent just five weeks in the Galpagos, and David Lack spent three months, Peter and Rosemary Grant and their colleagues have made research trips to the Galpagos for about thirty years, particularly studying Darwin's finches. Here we look briefly at the case of the large cactus finch Geospiza conirostris on Isla Genovesa (formerly Tower Island) which is formed from a shield volcano, and is home to a variety of birds. These birds, like all well-studied groups,[16] show various kinds of morphism. Males are dimorphic in song type: songs A and B are quite distinct. Also, males with song A have shorter bills than B males. This is also a clear difference. With these beaks males are able to feed differently on their favourite cactus, the prickly pear Opuntia. Those with long beaks are able to punch holes in the cactus fruit and eat the fleshy aril pulp which surrounds the seeds, whereas those with shorter beaks tear apart the cactus base and eat the pulp and any insect larvae and pupae (both groups eat flowers and buds). This dimorphism clearly maximises their feeding opportunities during the non-breeding season when food is scarce.

Darwin's finches If the population is panmixic,[17][18] then Geospiza conirostris exhibits a balanced genetic polymorphism and not, as originally supposed, a case of nascent sympatric speciation. The selection maintaining the polymorphism maximises the species' niche by expanding its feeding opportunity. The genetics of this situation cannot be clarified in the absence of a detailed breeding program, but two loci with linkage disequilibrium[19] is a possibility. Another interesting dimorphism is for the bills of young finches, which are either 'pink' or 'yellow'. All species of Darwin's finches exhibit this morphism, which lasts for two months. No interpretation of this phenomenon is known.[20]

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Taxonomy
Family
For some decades taxonomists have placed these birds in the family Emberizidae with the New World sparrows and Old World buntings (Sulloway 1982). However, the Sibley-Ahlquist taxonomy puts Darwin's finches with the tanagers (Monroe and Sibley 1993), and at least one recent work follows that example (Burns and Skutch 2003). The American Ornithologists' Union, in its North American check-list, places the Cocos Finch in the Emberizidae but with an asterisk indicating that the placement is probably wrong (AOU 19982006); in its tentative South American check-list, the Galpagos species are incertae sedis, of uncertain place (Remsen et al. 2007).

Species
Genus Geospiza Large Cactus Finch (Geospiza conirostris) Sharp-beaked Ground Finch (Geospiza difficilis) Vampire Finch (Geospiza difficilis septentrionalis) Medium Ground Finch (Geospiza fortis) Small Ground Finch (Geospiza fuliginosa) Large Ground Finch (Geospiza magnirostris) Darwin's Large Ground Finch (Geospiza magnirostris magnirostris) possibly extinct (1957?) Common Cactus Finch (Geospiza scandens) Genus Camarhynchus Large Tree Finch (Camarhynchus psittacula) Medium Tree Finch (Camarhynchus pauper) Small Tree Finch (Camarhynchus parvulus) Woodpecker Finch (Camarhynchus pallidus) sometimes separated in Cactospiza Mangrove Finch (Camarhynchus heliobates) Genus Certhidea Green Warbler-Finch (Certhidea olivacea) Grey Warbler-Finch (Certhidea fusca) Genus Pinaroloxias Cocos Finch (Pinaroloxias inornata) Genus Platyspiza Vegetarian Finch (Platyspiza crassirostris)

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Molecular basis of beak evolution

Developmental research in 2004 found that bone morphogenetic protein 4 (BMP4), and its differential expression during development, resulted in variation of beak size and shape among finches. BMP4 acts in the developing embryo to lay down skeletal features, including the beak.[21] The same group showed that the different beak shapes of Darwin's finches develop are also influenced by slightly different timing and spatial expression of a gene called calmodulin (CaM).[22] Calmodulin acts in a similar way to BMP4, affecting some of the features of beak growth. The authors suggest that changes in the temporal and spatial expression of these two factors are possible developmental controls of beak morphology.

Notes
[2] Lack, David. 1947. Darwin's Finches. Cambridge University Press (reissued in 1961 by Harper, New York, with a new preface by Lack; reissued in 1983 by Cambridge University Press with an introduction and notes by Laurene M. Ratcliffe and Peter T. Boag). ISBN 0-521-25243-1 [4] Grant, K. Thalia and Estes, Gregory B. 2009 "Darwin in Galapagos: Footsteps to a New World" Princeton University Press, Princeton. [16] Huxley J. 1955. Morphism in birds. 11th Int Ornith Cong (Basel 1954, p309-328) touches on this theme. [17] B. Rosemary Grant and Peter R. Grant 1989. Evolutionary dynamics of a natural population: the large cactus finch of the Galpagos. Chicago, p241 first para. [18] Grant, Peter R. 1999. Ecology and evolution of Darwin's finches. Princeton NJ, p428 in Afterword. [19] Maynard Smith J. 1998. Evolutionary genetics. 2nd ed, Chapter 5, Oxford. [20] Grant, Peter R. 1999. Ecology and evolution of Darwin's finches. Princeton NJ. (see plate 7)

References
Darwin, Charles (1839), Narrative of the surveying voyages of His Majesty's Ships Adventure and Beagle between the years 1826 and 1836, describing their examination of the southern shores of South America, and the Beagle's circumnavigation of the globe. Journal and remarks. 18321836 III, London: Henry Colburn Darwin, Charles (1845), Journal of researches into the natural history and geology of the countries visited during the voyage of H.M.S. Beagle round the world, under the Command of Capt. Fitz Roy, R.N (2nd. ed.), London: John Murray Darwin, Charles (1859), On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (http://darwin-online.org.uk/content/frameset?itemID=F373&viewtype=text& pageseq=1) (1st ed.), London: John Murray Darwin, Francis (1887), "Chapter 1, The Foundations of the 'Origin of Species'" (http://darwin-online.org.uk/ content/frameset?viewtype=text&itemID=F1452.2&pageseq=21), in Darwin, Francis, The life and letters of Charles Darwin, including an autobiographical chapter 2, London: John Murray Desmond, Adrian; Moore, James (1991), Darwin, London: Michael Joseph, Penguin Group, ISBN0-7181-3430-3, OCLC 185764721 (http://www.worldcat.org/oclc/185764721) Eldredge, Niles (2006), "Confessions of a Darwinist" (http://www.vqronline.org/articles/2006/spring/ eldredge-confessions-darwinist/), The Virginia Quarterly Review (Spring 2006): 3253, retrieved 2008-11-04 Grant, Peter R.; Grant, B. Rosemary (2008), How and Why Species Multiply: The Radiation of Darwin's Finches, Princeton University Press, ISBN978-0-691-13360-7 Lack, David (1940), "Evolution of the Galapagos Finches", Nature (7 September 1940) 146 (146): 324327, Bibcode: 1940Natur.146..324L (http://adsabs.harvard.edu/abs/1940Natur.146..324L), doi: 10.1038/146324a0 (http://dx.doi.org/10.1038/146324a0) |accessdate= requires |url= (help) Check-list of North American Birds (http://web.archive.org/web/20070404023350/http://www.aou.org/ checklist/index.php3), American Ornithologists' Union, 19982006, archived from the original (http://www. aou.org/checklist/index.php3) on April 4, 2007, retrieved 2007-04-09 Burns, Kevin J.; Skutch, Alexander F. (2003), "Tanagers and Tanager-Finches" (http://www.amazon.com/ Firefly-Encyclopedia-Birds-Christopher-Perrins/dp/1552977773/ref=si3_rdr_bb_product/

Darwin's finches 103-5704731-0011011), in Christopher Perrins, ed., The Firefly Encyclopedia of Birds, Firefly Books, pp.629631, ISBN1-55297-777-3, retrieved 2007-04-09 It is not clear whether this placement was made by Burns and Skutch or by Perrins. Keynes, Richard (2000), Charles Darwin's zoology notes & specimen lists from H.M.S. Beagle (http:// darwin-online.org.uk/content/frameset?itemID=F1840&viewtype=text&pageseq=1), Cambridge University Press, retrieved 2008-12-08 More than one of |author= and |last= specified (help) Sibley, Charles G. (1993), New Haven, Conn.: Yale University Press, ISBN0-300-07083-7 A World Checklist of Birds http://books.google.com/?vid=ISBN0300070837title= A World Checklist of Birds (http://books. google.com/?vid=ISBN0300070837title=), retrieved 2007-04-09 Missing or empty |title= (help) Monroe and Sibley consider the tanagers to be a tribe (Thraupini) of a big family Fringillidae rather than a family of their own (Thraupidae). Zimmer, J. (2007-04-05), A classification of the bird species of South America (http://www.museum.lsu.edu/ ~Remsen/SACCBaseline.html), American Ornithologists' Union, retrieved 2007-04-09 Steinheimer, F. D. (2004), "Charles Darwin's bird collection and ornithological knowledge during the voyage of H.M.S. Beagle, 18311836" (http://darwin-online.org.uk/content/frameset?viewtype=text&itemID=A161& pageseq=1), Journal of Ornithology 145 (4): 300320, doi: 10.1007/s10336-004-0043-8 (http://dx.doi.org/10. 1007/s10336-004-0043-8), retrieved 2008-12-08

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Sulloway, Frank J. (1982), "The Beagle collections of Darwin's finches (Geospizinae)" (http://darwin-online. org.uk/content/frameset?itemID=A86&viewtype=image&pageseq=1), Bulletin of the British Museum (Natural History), Historical Series 43 (2): 4994, retrieved 2008-12-08 Sulloway, Frank J. (Spring 1982), "Darwin and His Finches: The Evolution of a Legend" (http://www.sulloway. org/Finches.pdf) (PDF), Journal of the History of Biology 15 (1): 153, doi: 10.1007/BF00132004 (http://dx. doi.org/10.1007/BF00132004), retrieved 2008-12-09 Sulloway, Frank J. (2006), "Why Darwin Rejected Intelligent Design" (http://www.sulloway.org/Why Darwin Rejected Intelligent Design (2006).pdf), in Brockman, John, Intelligent Thought: Science versus the Intelligent Design Movement, New York: Vintage, pp.107126

External links
Grant, K.T. and Estes, G. B. 2009. "Darwin in Galapagos: Footsteps to a New World" Princeton University Press, Princeton. (http://www.darwiningalapagos.com) Sulloway, F.J. (1982): "Darwin and his finches: the evolution of a legend". J. Hist. Biol. 15: p.153 Different bills and song melodies (http://www.nature.com/nature/journal/v409/n6817/fig_tab/409185a0_F1. html) Genetics and the Origin of Birds Species, Grant and Grant in PNAS (http://www.pnas.org/cgi/reprint/94/15/ 7768) Sato et al. Phylogeny of Darwin's finches as revealed by mtDNA sequences in PNAS (http://www.pnas.org/ cgi/content/full/96/9/5101?maxtoshow=&HITS=10&hits=10&RESULTFORMAT=&author1=Sato& andorexactfulltext=and&searchid=1081449482400_6822&stored_search=&FIRSTINDEX=0& sortspec=relevance&volume=96&firstpage=5101&resourcetype=1) Sato A, Tichy H, O'hUigin C, Grant PR, Grant BR, Klein J (March 2001), "On the origin of Darwin's finches" (http://mbe.oxfordjournals.org/cgi/content/full/18/3/299), Mol. Biol. Evol. 18 (3): 299311, PMID 11230531 (http://www.ncbi.nlm.nih.gov/pubmed/11230531) Galpagos Online. Darwin's Finches. (http://www.galapagosonline.com/Galapagos_Natural_History/ Birds_and_Animals/Birds/Darwins_Finches.html) Galapagos Online. List of birds of the Galapagos Islands. (http://www.galapagosonline.com/ Galapagos_Natural_History/Birds_and_Animals/Birds/Bird_List.html#Mimidae)

Darwin's finches Darwin's Finches Evolve Before Scientists' Eyes (http://www.livescience.com/ 4147-darwin-finches-evolve-scientists-eyes.html): new developments reported 13 July 2006 Fink F.A.Q. (http://fink.sourceforge.net/faq/general.php?phpLang=en#what) Darwin's finches inspired the naming of the Fink project, a collaborative initiative for porting open source software to the Darwin platform to enable its use and evolution in the Apple Mac OS X environment. "Fink" is the German name for "finch." Aug 2006 Nature Article (http://www.nature.com/nature/journal/v442/n7102/abs/nature04843.html) that shows how modulation of a certain gene during development can account for the differences seen in beak shape. Speciation (http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/S/Speciation.html) Kimball's Biology Pages

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Galpagos Islands

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Galpagos Islands
Galpagos Islands

Native name: Archipilago de Galpagos

Geography Location Coordinates Major islands Area Pacific Ocean 040S 9033W 18 8,010km2 (3,093sqmi) [1]

Coordinates: 040S 9033W [1]

Highestelevation 1,707m (5,600ft) Highestpoint Volcn Wolf Country Ecuador Province Capital city Galpagos Puerto Baquerizo Moreno Demographics Population 26,640 (as of 2012) Additional information

UNESCO World Heritage Site

Official name: Galpagos Islands Type: Criteria: Designated: ReferenceNo. Region: Extension: Natural vii, viii, ix, x 1978 (2nd session) 1 [2]

Latin America and the Caribbean 2001 and 2003

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Endangered: 20072010

The Galpagos Islands (official name: Archipilago de Col;n other Spanish names: Islas Galpagos) are an archipelago of volcanic islands distributed on either side of the Equator in the Pacific Ocean, 926km (575mi) west of continental Ecuador, of which they are a part. The Galpagos Islands and their surrounding waters form an Ecuadorian province, a national park, and a biological marine reserve. The principal language on the islands is Spanish. The islands have a population of slightly over 25,000.[3] The islands are famed for their vast number of endemic species and were studied by Charles Darwin during the voyage of the Beagle. His observations and collections contributed to the inception of Darwin's theory of evolution by natural selection. The first recorded visit to the islands happened by chance in 1535, when the Dominican friar Fray Tomas de Berlanga went to Peru to arbitrate in a dispute between Francisco Pizarro and his subordinates. De Berlanga was blown off course, though he eventually returned to the Spanish Empire and described the conditions of the islands and the animals that inhabited them. The first navigation chart of the islands was made by the buccaneer Ambrose Cowley in 1684.Wikipedia:Disputed statement He named the individual islands after some of his fellow pirates or after the British noblemen who helped the privateer's cause. More recently, the Ecuadorian Government gave most of the islands Spanish names. While the Spanish names are official, many users (especially ecological researchers) continue to use the older English names, principally because those were the names used when Charles Darwin visited.

Physical geography
The islands are located in the eastern Pacific Ocean, 973km (525nmi; 605mi) off the west coast of South America. The closest land mass is that of mainland Ecuador, the country to which they belong, 926km/500nmi to the east. The islands are found at the coordinates 140'N136'S, 8916'9201'W. Straddling the equator, islands in the chain are located in both the northern and southern hemispheres, with Volcn Wolf and Volcn Ecuador on Isla Isabela being directly on the equator. Espaola Island, the southernmost islet of the archipelago, and Darwin Island, the northernmost one, are spread out over a distance of 220km (137mi). The International Hydrographic Organization (IHO) considers them wholly within the South Pacific Ocean, however.[4] The Galpagos Archipelago consists of 7,880km2 (3,040sqmi) of land spread over 45,000km2 (17,000sqmi) of ocean. The largest of the islands, Isabela, measures 2,250 sq mi/5,827km2 [5] and makes up close to three-quarters of the total land area of the Galpagos. Volcn Wolf on Isabela is the highest point, with an elevation of 1,707m (5,600ft) above sea level.

Orthographic projection centred over the Galpagos.

The group consists of 18 main islands, 3 smaller islands, and 107 rocks and islets. The islands are located at the Galapagos Triple Junction. The archipelago is located on the Nazca Plate (a tectonic plate), which is moving east/southeast, diving under the South American Plate at a rate of about 2.5 inches (6.4cm) per year.[6] It is also atop the Galapagos hotspot, a place where the Earth's crust is being melted from

Galpagos Islands

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below by a mantle plume, creating volcanoes. The first islands formed here at least 8million and possibly up to 90million years ago.[] While the older islands have disappeared below the sea as they moved away from the mantle plume, the youngest islands, Isabela and Fernandina, are still being formed, with the most recent volcanic eruption in April 2009 where lava from the volcanic island Fernandina started flowing both towards the island's shoreline and into the centre caldera.

School of scalloped hammerheads, Wolf Island, Galapagos Islands

Main islands
The 18 [7] main islands (with a land area larger than 1km2) of the archipelago (with their English names) shown alphabetically: Baltra (South Seymour) Island Also known as South Seymour, Baltra is a small flat island located near the centre of the Galpagos. It was created by geological uplift. The island is very arid, and vegetation consists of salt bushes, prickly pear cacti and palo santo trees. Until 1986, Baltra (Seymour) Airport was the only airport serving the Galpagos. Now, there are two airports which receive flights from the continent; the other is located on San Cristbal Island. Private planes flying to Galpagos must fly to Baltra, as it is the only airport with facilities for planes overnight. On arriving in Baltra, all visitors are immediately transported by bus to one of two docks. The first dock is located in a small bay, where the boats cruising Galpagos await passengers. The second is a ferry dock, which connects Baltra to the island of Santa Cruz. In 2007 and 2008, the Baltra airport was remodelled to include additional restaurants, shops and an improved visitor area. During the 1940s, scientists decided to move 70 of Baltra's land iguanas to the neighboring North Seymour Island as part of an experiment. This move had unexpected results during the military occupation of Baltra in World War II; the native iguanas became extinct on the island. During the 1980s, iguanas from North Seymour were brought to the Charles Darwin Research Station as part of a breeding and repopulation project, and in the 1990s, land iguanas were reintroduced to Baltra. As of 1997, scientists counted 97 iguanas living on Baltra; 13 of which were born on the islands. Bartolom (Bartholomew) Island Bartolom Island is a volcanic islet just off the east coast of Santiago Island in the Galpagos
Another school of scalloped hammerheads at Wolf Island, Galapagos

Graspus graspus enjoying the weather

Satellite photo of the Galpagos islands overlayed with the names of the visible main islands.

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Islands group. it is one of the "younger" islands in the Galpagos archipelago. This island, and neighbouring Sulivan Bay on Santiago (James) island, are named after lifelong friend of Charles Darwin, Sir Bartholomew James Sulivan, who was a Lieutenant aboard HMS Beagle.[8] Today Sulivan Bay is often misspelled Sullivan Bay. This island is one of the few that is home to the Galapagos penguin which is the only wild penguin species to live on the equator. The green turtle is another animal that resides on the island. Darwin (Culpepper) Island This island is named after Charles Darwin. It has an area of 1.1 square km (0.4 sq mi) and a maximum altitude of 168m (551ft). Here fur seals, frigates, marine iguanas, swallow-tailed gulls, sea lions, whales, marine turtles, and red-footed and Nazca boobies can be seen. Espaola (Hood) Island Its name was given in honor of Spain. It also is known as Hood, after Viscount Samuel Hood. It has an area of 60 square km (23 sq mi) and a maximum altitude of 206m (676ft). Espaola is the oldest island at around 3.5million years, and the southernmost in the group. Due to its remote location, Espaola has a large number of endemic species. It has its own species of lava lizard, mockingbird, and tortoise. Espaola's marine iguanas exhibit a distinctive red coloration change between the breeding season. Espaola is the only place where the waved albatross nests. Some of the birds have attempted to breed on Genovesa (Tower) Island, but unsuccessfully. Espaola's steep cliffs serve as the perfect runways for these birds, which take off for their ocean feeding grounds near the mainland of Ecuador and Peru. Espaola has two visitor sites. Gardner Bay is a swimming and snorkelling site, and offers a great beach. Punta Suarez has migrant, resident, and endemic wildlife, including brightly colored marine iguanas, Espaola lava lizards, hood mockingbirds, swallow-tailed gulls, blue-footed boobies, Nazca boobies, red-billed tropicbirds, Galpagos hawks, 3 species of Darwin's finches, and the waved albatross. Fernandina (Narborough) Island The name was given in honor of King Ferdinand II of Aragon, who sponsored the voyage of Columbus. Fernandina has an area of 642 square km (248 sq mi) and a maximum altitude of 1,494 m (4,902ft). This is the youngest and westernmost island. On 13 May 2005, a new, very eruptive process began on this island, when an ash and water vapor cloud

Isabela seen from Spot Satellite.

Waved Albatrosses on Espaola.

Galpagos marine iguana.

Main Street on San Cristbal Island.

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rose to a height of 7km (23,000ft) and lava flows descended the slopes of the volcano on the way to the sea. Punta Espinosa is a narrow stretch of land where hundreds of marine iguanas gather, largely on black lava rocks. The famous flightless cormorants inhabit this island, as do Galpagos penguins, pelicans, Galpagos sea lions and Galpagos fur seals. Different types of lava flows can be compared, and the mangrove forests can be observed.
An animated tour of the Galpagos.

Floreana (Charles or Santa Mara) Island It was named after Juan Jos Flores, the first President of Ecuador, during whose administration the government of Ecuador took possession of the archipelago. It is also called Santa Maria, after one of the caravels of Columbus. It has an area of 173 square km (67 sq mi) and a maximum elevation of 640 m (2,100ft). It is one of the islands with the most interesting human history, and one of the earliest to be inhabited. Flamingos and green sea turtles nest (December to May) on this island. The patapegada or Galpagos petrel, a sea bird NASA oceanographer Gene Carl Feldman which spends most of its life away from land, is found here. At Post reflects on his unique perspective on this region. Office Bay, since the 18th century whalers kept a wooden barrel that served as post office so that mail could be picked up and delivered to their destinations, mainly Europe and the United States, by ships on their way home. At the "Devil's Crown", an underwater volcanic cone and coral formations are found. Genovesa (Tower) Island The name is derived from Genoa, Italy. It has an area of 14 square km (5.4 sq mi) and a maximum altitude of 76m (249ft). This island is formed by the remaining edge of a large caldera that is submerged. Its nickname of "the bird island" is clearly justified. At Darwin Bay, frigatebirds and swallow-tailed gulls, the only nocturnal species of gull in the world, can be seen. Red-footed boobies, noddy terns, lava gulls, tropic birds, doves, storm petrels and Darwin finches are also in sight. Prince Philip's Steps is a bird-watching plateau with Nazca and red-footed boobies. There is a large Palo Santo forest. Isabela (Albemarle) Island (Ecuador) This island was named in honor of Queen Isabela. With an area of 4,640 square km (1,792 sq mi), it is the largest island of the Galpagos. Its highest point is Volcn Wolf, with an altitude of 1,707 m (5,600ft). The island's seahorse shape is the product of the merging of six large volcanoes into a single land mass. On this island, Galpagos penguins, flightless cormorants, marine iguanas, pelicans and Sally Lightfoot crabs abound. At the skirts and calderas of the volcanoes of Isabela, land iguanas and Galpagos tortoises can be observed, as well as Darwin finches, Galpagos hawks, Galpagos doves and very interesting lowland vegetation. The third-largest human settlement of the archipelago, Puerto Villamil, is located at the southeastern tip of the island. It is the only island to have the equator run across it. It is also the only place in the world where a penguin can be in its natural habitat in the Northern Hemisphere.

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Marchena (Bindloe) Island: Named after Fray Antonio Marchena, it has an area of 130 square km (50 sq mi) and a maximum altitude of 343 m (1,125ft). Galpagos hawks and sea lions inhabit this island, and it is home to the Marchena lava lizard, an endemic animal. North Seymour Island Its name was given after an English nobleman, Lord Hugh Seymour. It has an area of 1.9 square km (0.7 sq mi) and a maximum altitude of 28m (92ft). This island is home to a large population of blue-footed boobies and swallow-tailed gulls. It hosts one of the largest populations of frigate birds. It was formed from geological uplift.

North Seymour Island in the Galapagos about to land on shore

Pinzn (Duncan) Island Named after the Pinzn brothers, captains of the Pinta and Nia caravels, it has an area of 18 square km (7 sq mi) and a maximum altitude of 458 m (1,503ft). Pinta (Abingdon) Island Named after the Pinta caravel, it has an area of 60km2 (23sqmi) and a maximum altitude of 777m (2,549ft). Sea lions, Galpagos hawks, giant tortoises, marine iguanas, and dolphins can be seen here. Pinta Island was home to the last remaining Pinta tortoise, called Lonesome George. He was moved from Pinta Island to the Charles Darwin Research Station on Santa Cruz Island, where scientists were attempting to breed from him. However, Lonesome George died in June 2012 without producing any offspring. Rbida (Jervis) Island It bears the name of the convent of Rbida, where Columbus left his son during his voyage to the Americas. It has an area of 4.9 square km (1.9 sq mi) and a maximum altitude of 367 m (1,204ft). The high amount of iron contained in the lava at Rbida gives it a distinctive red colour. White-cheeked pintail ducks live in a saltwater lagoon close to the beach, where brown pelicans and boobies have built their nests. Until recently, flamingos were also found in the lagoon, but they have since moved on to other islands, likely due to a lack of food on Rbida. Nine species of finches have been reported in this island. San Cristbal (Chatham) Island It bears the name of the patron saint of seafarers, "St. Christopher". Its English name was given after William Pitt, 1st Earl of Chatham. It has an area of 558 square km (215 sq mi) and its highest point rises to 730m (2395ft). This is the first island in the Galapagos Archipelago Charles Darwin visited during his voyage on the Beagle. This islands hosts frigate birds, sea lions, giant tortoises, blue- and red-footed boobies, tropicbirds, marine iguanas, dolphins and swallow-tailed gulls. Its vegetation includes Calandrinia galapagos, Lecocarpus darwinii, and trees such as Lignum vitae. The largest freshwater lake in the archipelago, Laguna El Junco, is located in the highlands of San Cristbal. The capital of the province of Galpagos, Puerto Baquerizo Moreno, lies at the southern tip of the island.

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Santa Cruz (Indefatigable) Island (Galpagos) Given the name of the Holy Cross in Spanish, its English name derives from the British vessel HMS Indefatigable. It has an area of 986 square km (381 sq mi) and a maximum altitude of 864m (2834ft). Santa Cruz hosts the largest human population in the archipelago, the town of Puerto Ayora. The Charles Darwin Research Station and the headquarters of the Galpagos National Park Service are located here. The GNPS and CDRS operate a tortoise breeding centre here, where young tortoises are hatched, reared, and prepared to be reintroduced to their natural habitat. The Highlands of Santa Cruz offer exuberant flora, and are famous for the lava tunnels. Large tortoise populations are found here. Black Turtle Cove is a site surrounded by mangroves, which sea turtles, rays and small sharks sometimes use as a mating area. Cerro Dragn, known for its flamingo lagoon, is also located here, and along the trail one may see land iguanas foraging.

From an aircraft flying out of Baltra Island (on the right) and the Santa Cruz(on the left), the Itabaca Channel is the waterway in the middle where water taxis take people in between and to waiting boats off shore waiting for multi day cruises.

Santa F (Barrington) Island Named after a city in Spain, it has an area of 24 square km (9 sq mi) and a maximum altitude of 259m (850ft). Santa Fe hosts a forest of Opuntia cactus, which are the largest of the archipelago, and Palo Santo. Weathered cliffs provide a haven for swallow-tailed gulls, red-billed tropic birds and shear-waters petrels. Santa Fe species of land iguanas are often seen, as well as lava lizards. Santiago (San Salvador, James) Island (Galpagos) Its name is equivalent to Saint James in English; it is also known as San Salvador, after the first island discovered by Columbus in the Caribbean Sea. This island has an area of 585 square km (226 sq mi) and a maximum altitude of 907m (2976ft). Marine iguanas, sea lions, fur seals, land and sea turtles, flamingos, dolphins and sharks are found here. Pigs and goats, which were introduced by humans to the islands and have caused great harm to the endemic species, have been eradicated (pigs by 2002; goats by the end of 2006). Darwin finches and Galpagos hawks are usually seen, as well as a colony of fur seals. At Sulivan Bay, a recent (around 100years ago) pahoehoe lava flow can be observed. Wolf (Wenman) Island This island was named after the German geologist Theodor Wolf. It has an area of 1.3 square km (0.5 sq mi) and a maximum altitude of 253m (830ft). Here, fur seals, frigatebirds, Nazca and red-footed boobies, marine iguanas, sharks, whales, dolphins and swallow-tailed gulls can be seen. The most famous resident is the vampire finch, which feeds partly on blood pecked from other birds, and is only found on this island.

Minor islands
Daphne Major A small island directly north of Santa Cruz and directly west of Baltra, this very inaccessible island appears, though unnamed, on Ambrose Cowley's 1684 chart. It is important as the location of multidecade finch population studies by Peter and Rosemary Grant. South Plaza Island (Plaza Sur) It is named in honor of a former president of Ecuador, General Leonidas Plaza. It has an area of 0.13 square km (0.05 sq mi) and a maximum altitude of 23m (75ft). The flora of South Plaza includes Opuntia cactus and Sesuvium plants, which form a reddish carpet on top of the lava formations. Iguanas (land, marine and some hybrids of both species) are abundant, and large numbers of birds can be observed from the cliffs at the southern part of the island, including tropic birds and swallow-tailed gulls. Nameless Island A small islet used mostly for scuba diving.

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Weather
Although located on the Equator, the Humboldt Current brings cold water to the islands, causing frequent drizzles during most of the year. The weather is periodically influenced by the El Nio events, which occur about every 37 years and are characterized by warm sea surface temperatures, a rise in sea level, greater wave action, and a depletion of nutrients in the water.[9] During the season known as the garua (June to November), the temperature by the sea is 22 C (72F), a steady and cold wind blows from south and southeast, frequent drizzles (garuas) last most of the day, and dense fog conceals the islands. During the warm season (December to May), the average sea and air temperature rises to 25 C (77F), there is no wind at all, there are sporadic, though strong, rains and the sun shines. Weather changes as altitude increases in the large islands. Temperature decreases gradually with altitude, while precipitation increases due to the condensation of moisture in clouds on the slopes. There is a large range in precipitation from one place to another, not only with altitude, but also depending on the location of the islands, and also with the seasons. The following table corresponding to the wet 1969 shows the variation of precipitation in different places of Santa Cruz Island:

These satellite maps show chlorophyll concentration (which corresponds with the abundance of phytoplankton) during El Nio (top) and La Nia (lower). Blue represents low concentrations, yellow, orange and red indicate high concentrations. Currents that normally fertilize the phytoplankton reverse during El Nio, resulting in barren oceans. These same currents are strengthened by La Nia, resulting in an explosion of ocean life.

The bottom image shows sea surface temperature, cool upwelling waters are coloured purple. Thriving phytoplankton populations are indicated by high chlorophyll concentrations (top image), coloured green and yellow. Images acquired on 2 March 2009.

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Location

Charles Darwin Station 6m 23.0mm 16.8mm 249.0mm 68.5mm 31.4mm 16.8mm 12.0mm 3.8mm 18.5mm 3.2mm 11.0mm 15.7mm 469.7mm

Devine Farm Media Luna

Altitude January February March April May June July August September October November December TOTALS

320 m 78.0mm 155.2mm 920.8mm 79.5mm 214.6mm 147.3mm 42.2mm 13.7mm 90.9mm 22.6mm 52.8mm 84.1mm 1901.7mm

620 m 172.6mm 117.0mm 666.7mm 166.4mm 309.8mm 271.8mm 135.6mm 89.5mm 282.6mm 96.5mm 172.7mm 175.3mm 2656.4mm

The precipitation also depends on the geographical location. During March 1969, the precipitation over Charles Darwin Station, on the southern coast of Santa Cruz was 249.0mm (9.80in), while on Baltra Island, the precipitation during the same month was only 137.6mm (5.42in). This is because Baltra is located behind Santa Cruz with respect to the prevailing southerly winds, so most of the moisture gets precipitated in the Santa Cruz highlands. There are significant changes in precipitation from one year to another, too. At Charles Darwin Station, the precipitation during March 1969 was 249.0mm (9.80in), but during March 1970, it was only 1.2mm (0.05in).

History
European discovery of the Galpagos Islands occurred when Spaniard Fray Toms de Berlanga, the fourth Bishop of Panama, sailed to Peru to settle a dispute between Francisco Pizarro and his lieutenants. De Berlanga's vessel drifted off course when the winds diminished, and his party reached the islands on 10 March 1535. According to a 1952 study by Thor Heyerdahl and Arne Skjlsvold, remains of potsherds and other artifacts from several sites on the islands suggest visitation by South American peoples prior to the arrival of the Spanish.[10] However, no remains of graves, ceremonial vessels and constructions have ever been found, suggesting no permanent settlement occurred at the time.[11]

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The Galpagos Islands first appeared on the maps, of Gerardus Mercator and Abraham Ortelius, in about 1570.[12] The islands were named "Insulae de los Galopegos" (Islands of the Tortoises) in reference to the giant tortoises found there.[13] The first English captain to visit the Galpagos Islands was Richard Hawkins, in 1593. Until the early 19th century, the archipelago was often used as a hideout by mostly English pirates who pilfered Spanish galleons carrying gold and silver from South America to Spain.

The Galpagos tortoise or Galpagos giant tortoise (Chelonoidis nigra) is the largest living species of tortoise, this is from the Island of Santa Cruz.

In 1793, James Colnett described the flora and fauna of Galpagos, and suggested the islands could be used as base for the whalers operating in the Pacific Ocean. He drew the first accurate navigation charts of the islands. Whalers and maritime fur traders killed and captured thousands of the Galpagos tortoises to extract their fat. The tortoises could be kept on board ship as a means of providing of fresh protein, as these animals could survive for several months on board without any food or water. The hunting of the tortoises was responsible for greatly diminishing, and in some cases eliminating, certain species. Along with whalers came the fur-seal hunters, who brought the population of this animal close to extinction. The first known permanent human resident on Galapagos was Patrick Watkins, an Irish sailor who was marooned on the Island Floreana from 18071809. According to later accounts,[14][15] Watkins managed to survive by hunting, growing vegetables and trading with visiting whalers, before finally stealing an open boat and navigating to Guayaquil. In 1818 the Nantucket whaleship Globe, Captain George Washington Gardner, had discovered a "mother lode" of sperm whales some thousand miles west of the South American coast approximately at the equator. He returned to Nantucket in 1820 with more than 2000 barrels of sperm whale oil and the news of his discovery. This led to an influx of whaleships to exploit the new whaling ground and the Galapagos Islands became a frequent stop for the whalers both before and after visiting what came to be known as the Offshore Grounds. This led to the establishment in the Galapagos Islands of a kind of unofficial "post office" where whaleships stopped to pick up and drop off letters as well as for purposes of reprovisioning and repairs.[16]
As described in 1684.

Galpagos Islands In October 1820, the whaleship Essex, out of Nantucket, stopped at the Galapagos for these purposes on its way to the Offshore Grounds. On what was then known as Charles Island, while most of the crew were hunting tortoises one crewmember, English boatsteerer Thomas Chappel, for reasons still unclear, lit a fire which quickly burned out of control. Some of the tortoise hunters had a narrow escape and had to run a gauntlet of fire to get back to the ship. Soon almost the entire island was in flames. Crewmembers reported that after a day of sailing away they could still see the flames against the horizon. One crewmember who returned to the Galapagos several years afterward described the entire island as still a blackened wasteland.[17] Ecuador annexed the Galpagos Islands on 12 February 1832, naming them the Archipelago of Ecuador. This new name added to several names that had been, and are still, used to refer to the archipelago. The first governor of Galpagos, General Jos de Villamil, brought a group of convicts to populate the island of Floreana, and in October 1832, some artisans and farmers joined them. The voyage of the Beagle brought the survey ship HMS Beagle, under captain Robert FitzRoy, to the Galpagos on 15 September 1835 to survey approaches to harbours. The captain and others on board, including his companion, the young naturalist Charles Darwin, made observations on the geology and biology on Chatham, Charles, Albemarle and James islands before they left on 20 October to continue on their round-the-world expedition. Primarily a geologist at the time, Darwin was impressed by the quantity of volcanic craters they saw, later referring to the archipelago as "that land of craters." His study of several volcanic formations over the 5 weeks he stayed in the islands, led to several important geological discoveries, including the first, correct explanation for how volcanic tuff is formed.[18] Darwin noticed the mockingbirds differed between islands, though he thought the birds now known as Darwin's finches were unrelated to each other, and did not bother labelling them by island.[] The Englishman Nicolas Lawson, acting Governor of Galpagos for the Republic of the Equator, met them on Charles Island, and as they walked to the prison colony, told Darwin the tortoises differed from island to island. Towards the end of the voyage, Darwin speculated that the distribution of the mockingbirds and the tortoises might "undermine the stability of Species".[19] When specimens of birds were analysed on his return to England, it was found that many apparently different kinds of birds were species of finches, which were unique to islands. These facts were crucial in Darwin's development of his theory of natural selection explaining evolution, which was presented in The Origin of Species.[] Jos Valdizn and Manuel Julin Cobos tried a new colonization, beginning the exploitation of a type of lichen found in the islands (Roccella portentosa) used as a coloring agent. After the assassination of Valdizn by some of his workers, Cobos brought from the continent a group of more than a hundred workers to San Cristbal Island, and tried his luck at planting sugar cane. He ruled his plantation with an iron hand, which led to his assassination in 1904. In 1897, Antonio Gil began another plantation on Isabela Island. Over the course of a whole year, from September 1904, an expedition of the Academy of Sciences of California, led by Rollo Beck, stayed in the Galpagos collecting scientific material on geology, entomology, ornithology, botany, zoology and herpetology. Another expedition from that Academy was done in 1932 (Templeton Crocker Expedition) to collect insects, fish, shells, fossils, birds and plants. For a long time during the early 1900s and at least through 1929, a cash strapped Ecuador had reached out for potential buyers of the islands to alleviate financial troubles at home. The US had repeatedly expressed its interest in buying the islands for military use as they were positioned strategically guarding the Panama Canal.[20] In 1920s and 1930s, a small wave of European settlers arrived in the islands. Ecuadorian laws provided all colonists with the possibility of receiving twenty hectares each of free land, the right to maintain their citizenship, freedom from taxation for the first ten years in Galapagos, and the right to hunt and fish freely on all uninhabited islands where they might settle.[21] The first European colonists to arrive were Norwegians who settled briefly on Floreana, before moving on to San Cristobal and Santa Cruz. A few years later, other colonists from Europe, America and Ecuador started arriving on the islands, seeking a simpler life[22] Descendants of the Norwegian Kastdalen family and the German Angermeyer still live on the islands.

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Galpagos Islands During World War II, Ecuador authorized the United States to establish a naval base in Baltra Island, and radar stations in other strategic locations. Baltra was established as a United States Army Air Force base. Crews stationed at Baltra patrolled the Pacific for enemy submarines, as well as provided protection for the Panama Canal. After the war, the facilities were given to the government of Ecuador. Today, the island continues as an official Ecuadorian military base. The foundations and other remains of the US base can still be seen as one crosses the island. In 1946, a penal colony was established in Isabela Island, but it was suspended in 1959. The Galpagos became a national park in 1959,[] and tourism started in the 1960s, imposing several restrictions upon the human population already living on the island. However, opportunities in the tourism, fishing, and farming industries attracted a mass of poor fishermen and farmers from mainland Ecuador. In the 1990s and 2000s, violent confrontations between parts of the local population and the Galapagos National Park Service occurred, including capturing and killing giant tortoises and holding staff of the Galapagos National Park Service hostage to obtain higher annual sea cucumber quotas.[23]
The marine iguana (Amblyrhynchus cristatus) Galpagos Islands Santa Cruz - swimming in Puerto Ayora

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Politics
The islands are administered by a provincial government. It was made a province by presidential decree by President Guillermo Rodrguez Lara on 18 February 1973. The province is divided into cantons, each covering certain islands. The capital is Puerto Baquerizo Moreno.

Demographics
The Galpagos Islands is one of the few places in the world without an indigenous population. The largest ethnic group is composed of Ecuadorian Mestizos, the mixed descendants of Spanish colonists and indigenous Native Americans, who arrived mainly in the last century from the continental part of Ecuador. There is also a large number of whites, mostly of Spanish descent. Some descendants of the early European and American colonists on the islands also still remain on the islands. In 1959, approximately 1,000 to 2,000 people called the islands their home. In 1972 a census in the archipelago recorded a population of 3,488. By the 1980s, this number had risen to more than 15,000 people, and 2006 estimates place the population around 25,000 people.

Flag of the Galpagos Province.

Water taxi on Puerto Ayora, Galapagos.

Five of the islands are inhabited: Baltra, Floreana, Isabela, San Cristobal and Santa Cruz.

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Travel
Options for flying into the Galapagos are limited to two islands; San Cristobal and Baltra. Private aircraft must use Baltra as it is the airport equipped with overnight plane accommodations. Seymour Airport on Baltra was recently renovated (2012-2013) to accommodate larger planes. Until 1969 the only way to visit was on a private or chartered vessel. There was no regular air service until Forrest Nelson's Hotel Galapagos began the first organized tours in April 1969. Soon other travel companies brought in tour ships and yachts, and local fishermen began converting their wooden boats for rudimentary cruising with guests. These vessels were the main source of overnight accommodations in the Galapagos. Today there are about 85 yachts and ships equipped for overnight guests, but are limited by the government to 16 up to 100 passengers. In 2006 the Baltra military governed island, was opened up to limited overnight camping. Baltra also requires permits by the military government for overnight stays on the beach. Other inhabited islands also allow camping on the beaches designated as "recreational" use to the locals. All of these camping permits are limited to number of people and nights, with most nights not to exceed 3. Land based hotels are opening on the inhabited islands of San Cristobal, Santa Cruz, Floreana and Isabela. By 2012, more than half the visitors to Galapagos made their tours using day boats and these small hotels. Restaurants,easy access and economy make this an attractive travel option. The cruise tours are still the best way to see all the complex environment and wildlife of the islands. There are only 116 visitor sites in the Galapagos: 54 land sites and 62 scuba-diving or snorkeling sites. Small groups are allowed to visit in 2-4 hour shifts only, to limit impact on the area. All groups are accompanied by licensed guides. In May 2013, Google announced it would be adding the Galapagos Islands to its Street View mapping service.[24]

Conservation
Though the first protective legislation for the Galpagos was enacted in 1930 and supplemented in 1936, it was not until the late 1950s that positive action was taken to control what was happening to the native flora and fauna. In 1955, the International Union for the Conservation of Nature organized a fact-finding mission to the Galpagos. Two years later, in 1957, UNESCO, in cooperation with the government of Ecuador, sent another expedition to study the conservation situation and choose a site for a research station. In 1959, the centenary year of Charles Darwin's publication of The Marine iguana Origin of Species, the Ecuadorian government declared 97.5% of the archipelago's land area a national park, excepting areas already colonised. The Charles Darwin Foundation (CDF) was founded the same year. The core responsibility of CDF, an international nongovernmental organization (NGO) constituted in Belgium, is to conduct research and provide the research findings to the government for effective management of Galpagos. CDF's research efforts began with the establishment of the Charles Darwin Research Station on Santa Cruz Island in 1964. During the early years, conservation programs, such as eradication of introduced species and protection of native species, were carried out by research station personnel. Now

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much of that work is accomplished by the Galapagos National Park Service using the research findings and methodologies developed by CDF. In 1986, the 70,000 square kilometers (27,000 sq mi.) of ocean surrounding the islands was declared a marine reserve, second in size only to Australia's Great Barrier Reef. In 1990, the archipelago became a whale sanctuary. UNESCO recognised the islands in 1978 as a World Heritage Site[25] and in 1985, as a biosphere reserve. This was later extended in December 2001 to include the marine reserve. In July 2010, the World Heritage Committee agreed to remove the Galapagos Islands from its list of precious sites endangered by environmental threats or overuse.[26] Noteworthy species include: Galpagos land iguanas, Conolophus spp. Marine iguana, Amblyrhynchus cristatus, the only iguana feeding in the sea Galpagos tortoise (Galpagos giant tortoise), Chelonoidis nigra, known as galpago in Spanish, it gave the name to the islands Galpagos green turtle, Chelonia mydas agassisi, a subspecies of the green turtle Sea cucumbers, the cause of environmental battles with fishermen over quotas of this expensive Asian delicacy Flightless cormorant, Phalacrocorax harrisi Great frigatebird and magnificent frigatebird Blue-footed booby, Sula nebouxii, popular among visitors for their large blue feet which they show off in courtship Galpagos penguin, Spheniscus mendiculus, the only living tropical penguin Waved albatross, Phoebastria irrorata, the only living tropical albatross Galpagos hawk, Buteo galapagoensis, the islands' main scavenger and "environmental police" Four endemic species of Galpagos mockingbirds, the first species Darwin noticed to vary from island to island Thirteen endemic species of tanagers, popularly called Darwin's finches. Among them is the sharp-beaked ground finch Geospiza difficilis septentrionalis which is sometimes called the "vampire finch" for its blood-sucking habits, and the tool-using woodpecker finch, Camarhynchus pallidus Galpagos sea lions, Zalophus wollebaeki, closely related to the California sea lion, but smaller
The Great Frigatebird Tortuga Bay on Santa Cruz Island, Galpagos Blue-footed booby

Galpagos land iguana basking on a rock

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Environmental threats
Introduced plants and animals, such as feral goats, cats, and cattle, brought accidentally or willingly to the islands by humans, represent the main threat to Galpagos. Quick to reproduce and with no natural predators, these alien species decimated the habitats of native species. The native animals, lacking natural predators on the islands, are defenseless to introduced predators. Some of the most harmful introduced plants are the guayaba or guava (Psidium guajava), avocado (Persea americana), cascarilla (Cinchona pubescens), balsa (Ochroma pyramidale), hill raspberry (Rubus niveus), various citrus (orange, grapefruit, lemon), floripondio, higuerilla (Ricinus communis) trees and the elephant grass, Pennisetum purpureum. These plants have invaded large areas and eliminated endemic species in the humid zones of San Cristobal, Floreana, Isabela and Santa Cruz. Also, these harmful plants are just a few of introduced species on the Galpagos Islands. There are over 700 introduced plant species today. There are only 500 native and endemic species. This difference is creating a major problem for the islands and the natural species that inhabit them. Many species were introduced to the Galpagos by pirates. Thor Heyerdahl quoted documents that mention the Viceroy of Peru, knowing that British pirates ate the goats that they themselves had released in the islands, ordered dogs to be freed there to eliminate the goats[27] Also, when colonization of Floreana by Jos de Villamil failed, he ordered the goats, donkeys, cattle and other animals from the farms in Floreana be transferred to other islands for the purpose of later colonization.
Galpagos tortoise on Santa Cruz Island (Galpagos)

Two Ducks on the Island of Santa Cruz

Non-native goats, pigs, dogs, rats, cats, mice, sheep, horses, donkeys, Clouds, Tortuga Bay. cows, poultry, ants, cockroaches, and some parasites inhabit the islands today. Dogs and cats attack the tame birds and destroy the nests of birds, land tortoises, and marine turtles. They sometimes kill small Galpagos tortoises and iguanas. Pigs are even more harmful, covering larger areas and destroying the nests of tortoises, turtles and iguanas, as well as eating the animals' native food. Pigs also knock down vegetation in their search for roots and insects. This problem abounds in Cerro Azul volcano and Isabela, and in Santiago, pigs may be the cause of the disappearance of the land iguanas that were so abundant when Darwin visited. The black rat (Rattus rattus) attacks small Galpagos tortoises when they leave the nest, so in Pinzn they Grapsus grapsus in the Galapagos. stopped the reproduction for a period of more than 50years; only adults were found on that island. Also, where the black rat is found, the endemic rat has disappeared. Cattle and donkeys eat all the available vegetation and compete with native species for the scarce water. In

Galpagos Islands

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1959, fishermen introduced one male and two female goats to Pinta island; by 1973, the National Park service estimated the population of goats to be over 30,000 individuals. Goats were also introduced to Marchena in 1967 and to Rabida in 1971. A recent goat eradication program, however, has cleared most of the goat population from Isabela. The fast-growing poultry industry on the inhabited islands has been cause for concern from local conservationists, who fear domestic birds could introduce disease into the endemic wild bird populations. The Galpagos marine sanctuary is under threat from a host of illegal fishing activities, in addition to other problems of development.[] The most pressing threat to the Marine Reserve comes from local, mainland and foreign fishing targeting marine life illegally within the Reserve, such as sharks (hammerheads and other species) for their fins,[] and the harvest of sea cucumbers out of season. Development threatens both land and sea species. The growth of both the tourism industry and local populations fuelled by high birth rates and illegal immigration threaten the wildlife of the Archipelago. The grounding of the oil tanker Jessica in 2001 and the subsequent oil spill brought this threat to world attention.

Spheniscus mendiculus - a juvenile Galapagos Penguin before it has banding markings.

Currently, the rapidly growing problems, including tourism and a human population explosion, are further destroying habitats. Travel Companies, conservation organizations, and other groups have formed IGTOA[28] (International Galapagos Tour Operators Association) that is dedicated to preserve and protect the Galapagos Islands for future generations of travellers. Membership is only open to organizations, but welcomes the involvement of individuals. In 2007, UNESCO put the Galpagos Islands on their List of World Heritage in Danger because of threats posed by invasive species, unbridled tourism and overfishing.[29] On 29 July 2010, the World Heritage Committee decided to remove the Galapagos Islands from the list because the Committee found significant progress had been made by Ecuador in addressing these problems.[30] On 28 January 2008, Galapagos National Park official Victor Carrion announced 53 sea lions (13 pups, 25 youngsters, 9 males and 6 females) were killed at Pinta, Galapagos Islands nature reserve, with their heads caved in. In 2001, poachers killed 35 male sea lions.[31] The Galpagos Islands were short-listed as a candidate to be one of the New7Wonders of Nature by the New7Wonders of Nature Foundation. As of February 2009, the archipelago was ranked first in Group B, the category for islands.[32] The islands' biodiversity is under threat from several sources. The human population is growing at an unsustainable rate of 8% per year (1995). Introduced species have caused damage, and in 1996 a US$5 million, five-year eradication plan commenced in an attempt to rid the islands of introduced species such as goats, rats, deer, and donkeys. El Nio has adversely affected the marine ecosystem. And in January 2001, an oil slick from a stranded tanker threatened the islands, but winds and shifting ocean currents helped disperse the oil before much damage was done. The 1997-98 El Nio adversely affected wildlife in the waters surrounding the islands, as the waters were 5 C (9F) warmer than normal. Corals and barnacles suffered, hammerhead sharks were driven away, and most of the island's seabirds failed to breed 1997-98. The mortality rate of marine iguanas rose as the green algae they feed on was replaced by inedible red algae. During the 1982-83 El Nio, 70% of the marine iguanas starved to death because of this.[33]

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References
[1] [2] [5] [7] http:/ / tools. wmflabs. org/ geohack/ geohack. php?pagename=Gal%C3%A1pagos_Islands& params=0_40_S_90_33_W_scale:1400000 http:/ / whc. unesco. org/ en/ list/ 1 "Galpagos Islands." The Columbia Encyclopedia. New York: Columbia University Press, 2008. Credo Reference. Web. 17 September 2012 Miller, B., Breckheimer, I., McCleary, A., Guzmn-Ramirez, L., Caplow, S., Jones-Smith, J., & Walsh, S. (2010). Using stylized agent-based models for populationenvironment research: a case study from the Galpagos Islands. Population & Environment, 31(6), 401-426. [8] Grant, K. Thalia and Estes, Gregory B. 2009. Darwin in Galapagos: Footsteps to a New World. Princeton: Princeton University Press. (http:/ / darwiningalapagos. com/ reviews) [10] Heyerdahl, Thor; & Skjolsvold, Arne (1956). Archaeological Evidence of Pre-Spanish Visits to the Galpagos Islands, Memoirs 12, Society for American Archaeology. [11] Lundh, Jacob, 1995. "A brief account of some early inhabitants of Santa Cruz Island." In Noticias de Galpagos No. 55. Charles Darwin Foundation for the Galpagos Islands. [14] Porter, D. (1815/2007) Journal of a Cruise Made to the Pacific Ocean. Kessinger Publishing. [15] Tarnmoor, Salvator R. (Herman Melville) (1854) Encantadas or the Enchanted Isles. Putnam's Monthly Magazine of American Literature, Science, and Art. MarchMay 1854. [16] Perry, Roger; The Galapagos Islands; 1972; Dodd, Mead & Company, New York p. 44 ISBN 978-0396065760 [17] Nickerson, T. (ca. 1876) Account of the Ship Essex Sinking, 18191821. Holograph ms. in the Thomas Nickerson Collection, 18191876, Folder 1. Nantucket, Massachusetts: Nantucket Historical Society. [18] Grant, K. Thalia and Estes, Gregory B. 2009. Darwin in Galapagos: Footsteps to a New World. Princeton: Princeton University Press. (http:/ / darwiningalapagos. com) [19] Keynes, Richard ed. 2000. Charles Darwin's zoology notes & specimen lists from H.M.S. Beagle. Cambridge: Cambridge University Press. June August 1836 (http:/ / darwin-online. org. uk/ content/ frameset?itemID=F1840& viewtype=text& pageseq=23), 291293 (http:/ / darwin-online. org. uk/ content/ frameset?viewtype=text& itemID=F1840& pageseq=328) [21] Lund, J.P. Galapagos: A Brief History. http:/ / www. galapagos. to/ TEXTS/ LUNDH-3. HTM#chap12 [22] Hoff, S. (1985) Drmmen om Galapagos. Oslo: Grndahl & Snn [23] Stutz, Bruce D., 1995, "The sea cucumber war," Audubon 97. [25] Grant, K. Thalia. 2009 "Darwin and the Galapagos: Evolution of a Legacy" World Heritage No. 54 (http:/ / whc. unesco. org/ en/ review/ 54/ ) [27] Heyerdahl & Skjlsvold, op. cit. [28] igtoa.org (http:/ / www. igtoa. org) [32] New 7 Wonders of the World: Live Ranking (http:/ / www. new7wonders. com/ nature/ en/ liveranking/ ) [33] "Galapagos Islands." The Hutchinson Unabridged Encyclopedia with Atlas and Weather guide. Abington: Helicon, 2010. Credo Reference. Web. 21 September 2012

This articleincorporates text from a publication now in the public domain:Chisholm, Hugh, ed. (1911). Encyclopdia Britannica (11th ed.). Cambridge University Press.

Further reading
Black, Juan (1973). Galpagos, Archipilago del Ecuador. (Quito, Ecuador). Comprehensive monograph by a former officer of Galpagos National Park, financed by the World Wildlife Fund and the Charles Darwin Foundation for the Galpagos Islands Grant, K. Thalia and Estes, Gregory B. (2009). Darwin in Galapagos: Footsteps to a New World. Princeton University Press, Princeton. (http://www.darwiningalapagos.com). Heyerdahl, Thor; & Skjolsvold, Arne (1956). Archaeological Evidence of Pre-Spanish Visits to the Galpagos Islands, Memoirs 12, Society for American Archaeology. Mller, Bodo; & Stolt, Matthias (2003). Galpagos Die verwunschenen Inseln. (BLV). ISBN 3-86108-909-2. Quammen, David (1996). The Song of the Dodo. Touchstone, New York. Romero, Simon (4 October 2009). "To Protect Galpagos, Ecuador Limits a Two-Legged Species" (http://www. nytimes.com/2009/10/05/world/americas/05galapagos.html). The New York Times. Perry, Roger (1972). The Galapagos Islands. (Dodd, Mead & Company). Short history illustrated with photographs and a map by a former director of Charles Darwin Research Station.

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External links
Galapagos National Park (http://www.galapagospark.org) "Galpagos Islands xeric scrub" (http://worldwildlife.org/ecoregions/nt1307). Terrestrial Ecoregions. World Wildlife Fund. Islas Galapagos (http://islasgalapagos.org) Galpagos geology, with general information on the Galpagos Islands (http://www.geo.cornell.edu/geology/ Galapagos.html)

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HMS Beagle

HMS Beagle in the Straits of Magellan, pencil sketch circa 1900

Career (UK)
Ordered: Laid down: Launched: Commissioned: 16 February 1817 June 1818 11 May 1820 1820

Decommissioned: 1845, transferred to Coastguard Fate: Sold and broken up 1870

General characteristics
Class & type: Tons burthen: Length: Beam: Draught: Sail plan: Complement: Armament: Cherokee-class brig-sloop 235 bm; 242 for second voyage 90.3ft (27.5m) 24.5ft (7.5m) 12.5ft (3.8m) Brig (barque from 1825) 120 as a ship-of-war, 65 plus 9 supernumeraries on second voyage 10 guns, reduced to 6 guns for survey voyages
[]

HMS Beagle was a Cherokee-class 10-gun brig-sloop of the Royal Navy. The vessel, constructed at a cost of 7,803, was launched on 11 May 1820 from the Woolwich Dockyard on the River Thames. In July of that year she took part in a fleet review celebrating the coronation of King George IV of the United Kingdom: allegedly, to salute at the coronation she became the first full rigged man-of-war to sail under the old London Bridge. After that there was no immediate need for Beagle so she "lay in ordinary", moored afloat but without masts or rigging. She was then adapted as a survey barque and took part in three expeditions. On the second survey voyage the young naturalist Charles Darwin was on board, and his work made Beagle one of the most famous ships in history.[1]

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First voyage (18261830)

On 27 September 1825 Beagle docked at Woolwich for repairs and fitted out for her new duties at a total cost of 5,913. Her guns were reduced from ten cannons to six and a mizzen mast was added to improve her maneuverability, thereby changing her from a brig to a bark (or barque). Beagle set sail from Plymouth on 22 May 1826 on her first voyage, under the command of Captain Pringle Stokes. The mission was to accompany the larger ship HMS Adventure (380tons) on a hydrographic survey of Patagonia and Tierra del Fuego, under the overall command of the Australian Captain Phillip Parker King, Commander and Surveyor.[2][3] Faced with the more difficult part of the survey in the desolate waters of Tierra del Fuego, Captain Pringle Stokes fell into a deep depression. At Port Famine on the Strait of Magellan he locked himself in his cabin for 14 days, then after getting over-excited and talking of preparing for the next cruise, shot himself on 2 August 1828. Following four days of delirium Stokes recovered slightly, but then his condition deteriorated and he died on 12 August 1828.[4] Captain Parker King then replaced Stokes with the First Lieutenant of the Beagle, Lieutenant W.G. Skyring as commander, and both ships sailed to Montevideo. On 13 October King sailed the Adventure to Rio de Janeiro for refitting and provisions. During this work Rear Admiral Sir Robert Otway, commander in chief of the South American station, arrived aboard HMSGanges and announced his decision that the Beagle was also to be brought to Montevideo for repairs, and that he intended to supersede Skyring. When the Beagle arrived, Otway put the ship under the command of his aide, Flag Lieutenant Robert FitzRoy.[5] The 23-year-old aristocrat FitzRoy proved an able commander and meticulous surveyor. In one incident a group of Fuegians stole a ship's boat, and FitzRoy took their families on board as hostages. Eventually he held two men, a girl and a boy, who was given the name of Jemmy Button, and these four native Fuegians were taken back with them when the Beagle returned to England on 14 October 1830. During this survey, the Beagle Channel was identified and named after the ship.[]

Second voyage (18311836)

FitzRoy had been given reason to hope that the South American Survey would be continued under his command, but when the Lords of the Admiralty appeared to abandon the plan, he made alternative arrangements to return the Fuegians. A kind uncle heard of this and contacted the Admiralty. Soon afterwards FitzRoy heard that he was to be appointed commander of HMS Chanticleer to go to Tierra del Fuego, but due to her poor condition Beagle was substituted for the voyage. FitzRoy was re-appointed as commander on 27 June 1831 and the Beagle was commissioned on 4 July 1831 under his command, with Lieutenants John Clements Wickham and Bartholomew James Sulivan.[]

Beagle being hailed by native Fuegians during the survey of Tierra del Fuego, painted by Conrad Martens who became ship's artist in 1833.

Longitudinal section of HMS Beagle as of 1832

Beagle was immediately taken into dock at Devonport for extensive rebuilding and refitting. As she required a new deck, FitzRoy had the upper-deck raised considerably, by 8inches (200mm) aft and 12inches (300mm) forward. The Cherokee-class ships had the reputation of being "coffin brigs," which handled badly and were prone to sinking; the raised deck gave the Beagle better handling and made

HMS Beagle her less liable to become top-heavy and capsize by helping the decks to drain more quickly so that less water would collect in the gunwales. Additional sheathing added to the hull added about seven tons to her burthen and perhaps fifteen to her displacement,[] and the ship was one of the first to be fitted with the lightning conductor invented by William Snow Harris. FitzRoy spared no expense in her fitting out, which included 22 chronometers,[][] and five examples of the Sympiesometer, a kind of mercury-free barometer patented by Alexander Adie which was favoured by FitzRoy as giving the accurate readings required by the Admiralty.[6] FitzRoy had found a need for expert advice on geology during the first voyage, and had resolved that if on a similar expedition, he would "endeavour to carry out a person qualified to examine the land; while the officers, and myself, would attend to hydrography."[] Command in that era could involve stress and loneliness, as shown by the suicide of Captain Stokes, and FitzRoy's own uncle Viscount Castlereagh had committed suicide under stress of overwork.[7] His attempts to get a friend to accompany him fell through, and he asked his friend and superior, Captain Francis Beaufort, to seek a gentleman naturalist as a self-financing passenger who would give him company during the voyage. A sequence of inquiries led to Charles Darwin, a young gentleman on his way to becoming a rural clergyman, joining the voyage.[8] Beagle was originally scheduled to leave on 24 October 1831 but because of delays in her preparations the departure was delayed until December. She attempted to depart on 10 December but ran into bad weather. Finally, on the morning of 27 December, the Beagle left her anchorage in the Barn Pool, under Mount Edgecumbe on the west side of Plymouth Sound,[9] on what was to become a ground breaking scientific expedition. After completing extensive surveys in South America she returned via New Zealand, Sydney, Hobart Town (6 February 1836), to Falmouth, Cornwall, England on 2 October 1836.[10]

57

The Beagle Laid Ashore drawn by Conrad Martens (1834) and engraved by Thomas Landseer (1838)

Darwin had kept a diary of his experiences, and rewrote this as the book titled Journal and Remarks, published in 1839 as the third volume of the official account of the expedition. This travelogue and scientific journal was widely popular, and was reprinted many times with various titles, becoming known as The Voyage of the Beagle.[11]

Third voyage (18371843)

Six months later, Beagle set off in 1837 to survey large parts of the coast of Australia under the command of Commander John Clements Wickham, who had been a Lieutenant on the second voyage, with assistant surveyor Lieutenant John Lort Stokes who had been a Midshipman on the first voyage of Beagle, then mate and assistant surveyor on the second voyage (no relation to Pringle Stokes). They started with the western coast between the Swan River (modern Perth, Australia) and the Fitzroy River, Western Australia, then surveyed both In 1837 HMS Beagle set off on a survey of shores of the Bass Strait at the southeast corner of the continent. To aid Australia, shown here in an 1841 watercolour by Owen Stanley. Beagle in her surveying operations in Bass Strait, the Colonial cutter Vansittart, of Van Diemens Land, was most liberally lent by His Excellency Sir John Franklin, and placed under the command of Mr Charles Codrington Forsyth, the Senior Mate, assisted by Mr Pasco, another of her Mates. In May 1839 they sailed north to survey the

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shores of the Arafura Sea opposite Timor. When Wickham fell ill and resigned, the command was taken over in March 1841 by Lieutenant John Lort Stokes who continued the survey. The third voyage was completed in 1843. Numerous places around the coast were named by Wickham, and subsequently by Stokes when he became captain, often honouring eminent people or the members of the crew. On 9 October 1839 Wickham named Port Darwin, which was first sighted by Stokes, in honour of their former shipmate Charles Darwin. They were reminded of him (and his "geologising") by the discovery there of a new fine-grained sandstone.[] A settlement there became the town of Palmerston in 1869, and was renamed Darwin in 1911.[]

"General Chart of Australia", showing coasts examined by Beagle during the third voyage in red, from John Lort Stokes' Discoveries in Australia

Final years
In 1845 Beagle was refitted as a static coastguard watch vessel like many similar watch ships stationed in rivers and harbours throughout the nation. She was transferred to HM Customs and Excise to control smuggling on the Essex coast in the navigable waterways beyond the north bank of the Thames Estuary. She was moored mid-river in the River Roach which forms part of an extensive maze of waterways and marshes known as The River Crouch and River Roach Tidal River System, located around and to the south and west of Burnham-on-Crouch. This large maritime area has a tidal coastline of 243km (151 miles), part of Essex's 565km (351 miles) of coastline - the largest coastline in the United Kingdom. In 1851, oyster companies and traders who cultivated and harvested the "Walflete" or "Walfleet" oyster Ostrea edulis, petitioned for the Customs and Excise watch vessel WV-7 (ex HMS Beagle) to be removed as she was obstructing the river and its oyster-beds. In the 1851 Navy List dated 25 May, it showed her renamed as Southend "W.V. No. 7" at Paglesham. In 1870, she was sold to "Messrs Murray and Trainer" for breaking up.

Possible resting place

Investigations started in 2000 by a team led by Dr Robert Prescott of the University of St Andrews found documents confirming that "W.V. 7" was Beagle, and noted a vessel matching her size shown midstream on the River Roach (in Paglesham Reach) on the 1847 hydrographic survey chart. A later chart showed a nearby indentation to the north bank of Paglesham Reach near the Eastend Wharf and near Waterside Farm. This could have been a dock for W.V. 7 - Beagle. Site investigations found an area of marshy ground some 15ft (5m) deep on the tidal river-bank, about 150 metres west of the boat-house. This discovery matched the chart position and many fragments of pottery of the correct period were found in the same area.[12] An atomic dielectric resonance survey carried out in November 2003 found traces of timbers forming the size and shape of the lower hull, indicating a substantial amount of timbers from below the waterline still in place. An old anchor of 1841 pattern was excavated. It was also found that the 1871 census recorded a new farmhouse in the name of William Murray and Thomas Rainer, leading to speculation that the merchant's name was a misprint for T. Rainer. The farmhouse was demolished in the 1940s, but a nearby boathouse incorporated timbers matching knee timbers used in Beagle. Two more large anchors similar to the one excavated from the ship's present location are known to have been found in neighbouring villages. It is believed that there were four anchors in the ship. Their investigations featured in a BBC Television programme which showed how each watch ship would have accommodated seven coastguard officers, drawn from other areas to minimise collusion with the locals. Each officer had about three rooms to house his family, forming a small community. They would use small boats to intercept smugglers, and the investigators found a causeway giving access at low tide across the soft mud of the river bank.

HMS Beagle Apparently the next coastguard station along was Kangaroo, a sister ship of Beagle.

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Replica
On 31 December 2011 the Nao Victoria Museum in Punta Arenas announced the building of the first full-size replica of HMS Beagle.[13][14] Construction began on 1 November 2012, using Nothofagus dombeyi timber from the local rainforest.[15][16] In 2013 the Chilean national press started to get interest in the work in progress.[17]
HMS Beagle replica 1:1

The first frames of the HMS Beagle replica under construction at the Nao Victoria Museum

Starboard view of the HMS Beagle replica from Nao Victoria's Aftcastle (20 March 2013)

Portsideview of the HMS Beagle replica (20 March 2013)

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Sources and references

HMS Beagle: Survey Ship "Extraordinary" / Karl Heinz Marquardt (2007) ISBN 0-85177-703-1 The Voyage of the Beagle, Charles Darwin (including FitzRoy's commentary on refitting the Beagle from his account of the voyage), Penguin Books, London 1989 ISBN 0-14-043268-X. Abridged version of Darwin's Journal and Remarks, with introduction by Janet Browne and Michael Neve. FitzRoy, Robert (1836). "Sketch of the Surveying Voyages of his Majesty's Ships Adventure and Beagle, 1825-1836. Commanded by Captains P. P. King, P. Stokes, and R. Fitz-Roy, Royal Navy. (Communicated by John Barrow)" [18]. Journal of the Geological Society of London 6: 311343. Retrieved 2012-05-14. King, P. P. (1839). FitzRoy, Robert, ed. Narrative of the surveying voyages of His Majesty's Ships Adventure and Beagle between the years 1826 and 1836, describing their examination of the southern shores of South America, and the Beagle's circumnavigation of the globe. Proceedings of the first expedition, 1826-30, under the command of Captain P. Parker King, R.N., F.R.S. [19] I. London: Henry Colburn FitzRoy, Robert (1839). Narrative of the surveying voyages of His Majesty's Ships Adventure and Beagle between the years 1826 and 1836, describing their examination of the southern shores of South America, and the Beagle's circumnavigation of the globe. Proceedings of the second expedition, 1831-36, under the command of Captain Robert Fitz-Roy, R.N. [20] II. London: Henry Colburn FitzRoy, Robert (1839). Narrative of the surveying voyages of His Majesty's Ships Adventure and Beagle between the years 1826 and 1836, describing their examination of the southern shores of South America, and the Beagle's circumnavigation of the globe. Appendix to Volume II [21]. London: Henry Colburn Darwin, Charles (1839). Narrative of the surveying voyages of His Majesty's Ships Adventure and Beagle between the years 1826 and 1836, describing their examination of the southern shores of South America, and the Beagle's circumnavigation of the globe. Journal and remarks. 1832-1836. [20] III. London: Henry Colburn John Lort Stokes (1846) Discoveries in Australia, With an Account of the Coasts and Rivers Explored and Surveyed During The Voyage of H.M.S. Beagle, in the Years 1837-38-39-40-41-42-43. By Command of the Lords Commissioners of the Admiralty. Also a Narrative of Captain Owen Stanley's Visits to the Islands in the Arafura Sea. Vol. 1 [22] and Vol. 2 [23] London: T. and W. Boone
[1] HMS Beagle - Port Cities, London (http:/ / www. portcities. org. uk/ london/ server/ show/ ConFactFile. 64/ HMS-Beagle. html), Royal Museum Greenwich, retrieved 3 February 2013, the story about sailing full-rigged under London Bridge appears on page 332 of William Howitt's 1865 The history of discovery in Australia, Tasmania, and New Zealand (http:/ / books. google. co. uk/ books?id=tsANAAAAQAAJ& pg=PA332& dq=beagle+ coronation+ King+ "George+ IV"& hl=en& sa=X& ei=WL0OUfD4C4jK0QWvzoHIDQ& redir_esc=y#v=onepage& q=beagle coronation King "George IV"& f=false), Longman, Green, Longman, Roberts, and Green, London. [3] King was born on Norfolk Island and left for England in 1796. Colonial Secretary Index, 1788-1825, In the New South Wales State Records (http:/ / www. records. nsw. gov. au/ indexes/ colsec/ k/ F31c_kh-ky-03. htm). [4] Guardian review: Man on a suicide mission (http:/ / books. guardian. co. uk/ reviews/ biography/ 0,,986985,00. html) [12] "Hunting the lost Beagle" BBC News 9 January 2009 (http:/ / news. bbc. co. uk/ 1/ hi/ sci/ tech/ 7819991. stm) [18] http:/ / darwin-online. org. uk/ content/ frameset?viewtype=text& itemID=A73& pageseq=1 [19] http:/ / darwin-online. org. uk/ content/ frameset?itemID=F10. 1& viewtype=text& pageseq=1 [20] http:/ / darwin-online. org. uk/ content/ frameset?itemID=F10. 2& viewtype=text& pageseq=1 [21] http:/ / darwin-online. org. uk/ content/ frameset?itemID=F10. 2a& viewtype=text& pageseq=1 [22] http:/ / www. gutenberg. org/ ebooks/ 12115 [23] http:/ / www. gutenberg. org/ ebooks/ 12146

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External links
Darwin Online - bibliography (http://darwin-online.org.uk/EditorialIntroductions/ Freeman_JournalofResearches.html): Proceedings of the first and second expeditions, and Darwin's Journal (The Voyage of the Beagle). Works by Charles Darwin (http://www.gutenberg.org/author/Charles+Darwin) at Project Gutenberg list includes The Voyage of the Beagle John Lort Stokes, Discoveries in Australia, Volume 1 (http://www.gutenberg.org/etext/12115), Volume 2 (http://www.gutenberg.org/etext/12146). Digitised copies of the original logs of HMS Beagle (http://badc.nerc.ac.uk/data/corral/ships/beagle.html), British Atmospheric Data Centre (http://badc.nerc.ac.uk)/The National Archives as part of the CORRAL project Robert FitzRoy, 1836, Sketch of the Surveying Voyages of his Majesty's Ships Adventure and Beagle, 1825-1836. Commanded by Captains P. P. King, P. Stokes, and R. Fitz-Roy, Royal Navy (http://darwin-online.org.uk/ content/frameset?itemID=A73&viewtype=image&pageseq=1). Journal of the Geological Society of London 6: 311-343 Visit and Testimony of Captain Fitz-Roy (http://www.caphorniers.cl/Fitz Roy/relato ing/testimony01.htm) HMS Beagle - Port Cities (http://www.portcities.org.uk/london/server/show/ConFactFile.64/HMS-Beagle. html) The sympiesometer of Alexander Adie (http://www.antique-horology.org/_editorial/sympiezometerfontijn/) The Journal of Syms Covington - Chapter 1. (http://www.asap.unimelb.edu.au/bsparcs/covingto/chap_1. htm) The replica HMS Beagle project (http://www.thebeagleproject.com/) BBC News - Darwin's Beagle ship 'found' (http://news.bbc.co.uk/2/hi/science/nature/3490564.stm) The Observer - Evolution of radar points to HMS Beagle's resting place. (http://observer.guardian.co.uk/ uk_news/story/0,6903,1148523,00.html) BBC News - Plans to build HMS Beagle replica for 2009 Darwin bicentenary. (http://news.bbc.co.uk/1/hi/ wales/south_west/5190734.stm) Museo Nao Victoria - Oficial blog of the building process for the full size HMS Beagle replica. (http:// construyendoelhmsbeagle.blogspot.com/)

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Species
In biology, a species (plural: species) is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, the difficulty of defining species is known as the species problem. Differing measures are often used, such as similarity of DNA, morphology, or ecological niche. Presence of specific locally adapted traits may further subdivide species into "infraspecific taxa" such as subspecies (and in botany other taxa are used, such as varieties, subvarieties, and formae). Species hypothesized to have the same ancestors are placed in one genus, based on similarities. The similarity of species is judged based on comparison of physical attributes, especially their DNA sequences, where available. All species are given a two-part name, a "binomial name". The first part of a binomial name is the generic name, the genus of the species. The second part is either called the specific name (a term used only in zoology) or the specific epithet (the term used in botany, which can also be used in zoology). For example, Boa constrictor is one of four species of the Boa genus. The first part of the name is capitalized, and the second part has a lower case. The binomial name is written in italics when printed and underlined when handwritten. A usable definition of the word "species" and reliable methods of identifying particular species are essential for stating and testing biological theories and for measuring biodiversity, though other taxonomic levels such as families may be considered in broad-scale studies.[] Extinct species known only from fossils are generally difficult to assign precise taxonomic rankings, which is why higher taxonomic levels such as families are often used for fossil-based studies.[][] The total number of non-bacterial and non-archaeal species in the world has been estimated at 8.7 million,[][] with previous estimates ranging from two million to 100 million.[]

The hierarchy of biological classification's eight major taxonomic ranks. A genus contains one or more species. Intermediate minor rankings are not shown.

Biologists' working definition

A usable definition of the word "species" and reliable methods of identifying particular species is essential for stating and testing biological theories and for measuring biodiversity. Traditionally, multiple examples of a proposed species must be studied for unifying characters before it can be regarded as a species. It is generally difficult to give precise taxonomic rankings to extinct species known only from fossils. Some biologists may view species as statistical phenomena, as opposed to the traditional idea, with a species seen as a class of organisms. In that case, a species is defined as a separately evolving lineage that forms a single gene pool. Although properties such as DNA-sequences and morphology are used to help separate closely related lineages,[1] this definition has fuzzy boundaries.[2] However, the exact definition of the term "species" is still controversial, particularly in prokaryotes,[3] and this is called the species problem.[] Biologists have proposed a range of more precise definitions, but the definition used is a pragmatic choice that depends on the particularities of the species of concern.[]

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Common names and species

The commonly used names for plant and animal taxa sometimes correspond to species: for example, "lion", "walrus", and "Camphor tree" each refers to a species. In other cases common names do not: for example, "deer" refers to a family of 34 species, including Eld's Deer, Red Deer and Elk (Wapiti). The last two species were once considered a single species, illustrating how species boundaries may change with increased scientific knowledge.

Placement within genera

Ideally, a species is given a formal, scientific name, although in practice there are very many unnamed species (which have only been described, not named). When a species is named, it is placed within a genus. From a scientific point of view this can be regarded as a hypothesis that the species is more closely related to other species within its genus (if any) than to species of other genera. Species and genus are usually defined as part of a larger taxonomic hierarchy. The best-known taxonomic ranks are, in order: life, domain, kingdom, phylum, class, order, family, genus, and species. This assignment to a genus is not immutable; later a different (or the same) taxonomist may assign it to a different genus, in which case the name will also change. In biological nomenclature, the name for a species is a two-part name (a binomial name), treated as Latin, although roots from any language can be used as well as names of locales or individuals. The generic name is listed first (with its leading letter capitalized), followed by a second term. The terminology used for the second term differs between zoological and botanical nomenclature. In zoological nomenclature, the second part of the name can be called the specific name or the specific epithet. For example, gray wolves belong to the species Canis lupus, coyotes to Canis latrans, golden jackals to Canis aureus, etc., and all of those belong to the genus Canis (which also contains many other species). For the gray wolf, the genus name is Canis, the specific name or specific epithet is lupus, and the binomen, the name of the species, is Canis lupus. In botanical nomenclature, the second part of the name can only be called the specific epithet. The 'specific name' in botany is always the combination of genus name and specific epithet. For example, the species commonly known as the longleaf pine is Pinus palustris; the genus name is Pinus, the specific epithet is palustris, the specific name is Pinus palustris. This binomial naming convention, later formalized in the biological codes of nomenclature, was first used by Leonhart Fuchs and introduced as the standard by Carolus Linnaeus in his 1753 Species Plantarum (followed by his 1758 Systema Naturae, 10th edition). At that time, the chief biological theory was that species represented independent acts of creation by God and were therefore considered objectively real and immutable, so the hypothesis of common descent did not apply.

Abbreviated names
Books and articles sometimes intentionally do not identify species fully and use the abbreviation "sp." in the singular or "spp." in the plural in place of the specific epithet: for example, Canis sp. This commonly occurs in the following types of situations: The authors are confident that some individuals belong to a particular genus but are not sure to which exact species they belong. This is particularly common in paleontology. The authors use "spp." as a short way of saying that something applies to many species within a genus, but do not wish to say that it applies to all species within that genus. If scientists mean that something applies to all species within a genus, they use the genus name without the specific epithet. In books and articles, genus and species names are usually printed in italics. Abbreviations such as "sp.", "spp.", "subsp.", etc. should not be italicized.

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Identification codes
Various codes have been devised for identifying particular species. For example: NCBI employs a numeric 'taxid' or Taxonomy identifier, a "stable unique identifier", e.g. the taxid of H. sapiens is 9606 [4] KEGG employs a three-letter code for a limited number of organisms; in this code, for example, H. sapiens is simply hsa;[5] UniProt employs an "organism mnemonic" of not more than five alphanumeric characters, e.g. HUMAN for H. sapiens [6] ITIS The Integrated Taxonomic Information System provides a unique number for each species.

Difficulty of defining "species" and identifying particular species

It is surprisingly difficult to define the word "species" in a way that applies to all naturally occurring organisms,[] and the debate among biologists about how to define "species" and how to identify actual species is called the species problem. Over two dozen distinct definitions of "species" are in use amongst biologists.[] Most textbooks follow Ernst Mayr's definition of a species as "groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups".[] It has been argued that this definition of species is not only a useful formulation, but is also a natural consequence of the effect of sexual reproduction on the dynamics of natural selection.[7][8][9][10] (Also see Speciation.) Various parts of this definition serve to exclude some unusual or artificial matings: Those that occur only in captivity (when the animal's normal mating partners may not be available) or as a result of deliberate human action Animals that may be physically and physiologically capable of mating but, for various reasons, do not normally do so in the wild The typical textbook definition above works well for most multi-celled organisms, but there are several types of situations in which it breaks down: By definition it applies only to organisms that reproduce sexually. So it does not work for asexually reproducing single-celled organisms and for the relatively few parthenogenetic or apomictic multi-celled organisms. The term "phylotype" is often applied to such organisms. Biologists frequently do not know whether two morphologically similar groups of organisms are "potentially" capable of interbreeding. There is considerable variation in the degree to which hybridization may succeed under natural conditions, or even in the degree to which some organisms use sexual reproduction between individuals to breed. In ring species, members of adjacent populations interbreed successfully but members of some non-adjacent populations do not. In a few cases it may be physically impossible for animals that are members of the same species to mate. However, these are cases, such as in breeds of dogs, in which human intervention has caused gross morphological changes, and are therefore excluded by the biological species concept.Wikipedia:Disputed statement Horizontal gene transfer makes it even more difficult to define the word "species". There is strong evidence of horizontal gene transfer between very dissimilar groups of prokaryotes, and at least occasionally between dissimilar groups of eukaryotes; and Williamson[] argues that there is evidence for it in some crustaceans and echinoderms. All definitions of the word "species" assume that an organism gets all its genes from one or two parents that are very like that organism, but horizontal gene transfer makes that assumption false.

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Definitions of species
The question of how best to define "species" is one that has occupied biologists for centuries, and the debate itself has become known as the species problem. Darwin wrote in chapter II of On the Origin of Species: No one definition has satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of a species. Generally the term includes the unknown element of a distinct act of creation.[11] But later, in The Descent of Man, when addressing "The question whether mankind consists of one or several species", Darwin revised his opinion to say: it is a hopeless endeavour to decide this point on sound grounds, until some definition of the term "species" is generally accepted; and the definition must not include an element that cannot possibly be ascertained, such as an act of creation.[12] The modern theory of evolution depends on a fundamental redefinition of "species". Prior to Darwin, naturalists viewed species as ideal or general types, which could be exemplified by an ideal specimen bearing all the traits general to the species. Darwin's theories shifted attention from uniformity to variation and from the general to the particular. According to intellectual historian Louis Menand, Once our attention is redirected to the individual, we need another way of making generalizations. We are no longer interested in the conformity of an individual to an ideal type; we are now interested in the relation of an individual to the other individuals with which it interacts. To generalize about groups of interacting individuals, we need to drop the language of types and essences, which is prescriptive (telling us what finches should be), and adopt the language of statistics and probability, which is predictive (telling us what the average finch, under specified conditions, is likely to do). Relations will be more important than categories; functions, which are variable, will be more important than purposes; transitions will be more important than boundaries; sequences will be more important than hierarchies.[] This shift results in a new approach to "species"; Darwin concluded that species are what they appear to be: ideas, which are provisionally useful for naming groups of interacting individuals. "I look at the term species", he wrote, "as one arbitrarily given for the sake of convenience to a set of individuals closely resembling each other ... It does not essentially differ from the word variety, which is given to less distinct and more fluctuating forms. The term variety, again, in comparison with mere individual differences, is also applied arbitrarily, and for convenience sake."
[]

Practically, biologists define species as populations of organisms that have a high level of genetic similarity. This may reflect an adaptation to the same niche, and the transfer of genetic material from one individual to others, through a variety of possible means. The exact level of similarity used in such a definition is arbitrary, but this is the most common definition used for organisms that reproduce asexually (asexual reproduction), such as some plants and microorganisms. This lack of any clear species concept in microbiology has led to some authors arguing that the term "species" is not useful when studying bacterial evolution. Instead they see genes as moving freely between even distantly related bacteria, with the entire bacterial domain being a single gene pool. Nevertheless, a kind of rule of thumb has been established, saying that species of Bacteria or Archaea with 16S rRNA gene sequences more similar than 97% to each other need to be checked by DNA-DNA Hybridization if they belong to the same species or not.[13] This concept has been updated recently, saying that the border of 97% was too low and can be raised to 98.7%.[14] In the study of sexually reproducing organisms, where genetic material is shared through the process of reproduction, the ability of two organisms to interbreed and produce fertile offspring of both sexes is generally accepted as a simple indicator that the organisms share enough genes to be considered members of the same species. Thus a "species" is a group of interbreeding organisms. This definition can be extended to say that a species is a group of organisms that could potentially interbreed fish could still be classed as the same species even if they live in different lakes, as long as they could still interbreed

Species were they ever to come into contact with each other. On the other hand, there are many examples of series of three or more distinct populations, where individuals of the population in the middle can interbreed with the populations to either side, but individuals of the populations on either side cannot interbreed. Thus, one could argue that these populations constitute a single species, or two distinct species. This is not a paradox; it is evidence that species are defined by gene frequencies, and thus have fuzzy boundaries. Consequently, any single, universal definition of "species" is necessarily arbitrary. Instead, biologists have proposed a range of definitions; which definition a biologists uses is a pragmatic choice, depending on the particularities of that biologist's research. In practice, these definitions often coincide, and the differences between them are more a matter of emphasis than of outright contradiction. Nevertheless, no species concept yet proposed is entirely objective, or can be applied in all cases without resorting to judgment. Given the complexity of life, some have arguedWikipedia:Avoid weasel words that such an objective definition is in all likelihood impossible, and biologists should settle for the most practical definition. For most vertebrates, this is the biological species concept (BSC), and to a lesser extent (or for different purposes) the phylogenetic species concept (PSC). Many BSC subspecies are considered species under the PSC; the difference between the BSC and the PSC can be summed up insofar as that the BSC defines a species as a consequence of manifest evolutionary history, while the PSC defines a species as a consequence of manifest evolutionary potential. Thus, a PSC species is "made" as soon as an evolutionary lineage has started to separate, while a BSC species starts to exist only when the lineage separation is complete. Accordingly, there can be considerable conflict between alternative classifications based upon the PSC versus BSC, as they differ completely in their treatment of taxa that would be considered subspecies under the latter model (e.g., the numerous subspecies of honey bees).

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Typological species
A group of organisms in which individuals are members of the species if they sufficiently conform to certain fixed properties. The clusters of variations or phenotypes within specimens (i.e. longer or shorter tails) would differentiate the species. This method was used as a "classical" method of determining species, such as with Linnaeus early in evolutionary theory. However, we now know that different phenotypes do not always constitute different species (e.g.: a 4-winged Drosophila born to a 2-winged mother is not a different species). Species named in this manner are called morphospecies.[15][16]

Evolutionary species
A single evolutionary lineage of organisms within which genes can be shared, and that maintains its integrity with respect to other lineages through both time and space. At some point in the evolution of such a group, some members may diverge from the main population and evolve into a subspecies, a process that may eventually lead to the formation of a new species if isolation (geographical or ecological) is maintained. A species that gives rise to another species is a paraphyletic species, or paraspecies.[]

Phylogenetic (cladistic) species

A group of organisms that shares an ancestor; a lineage that maintains its integrity with respect to other lineages through both time and space. At some point in the progress of such a group, members may diverge from one another: when such a divergence becomes sufficiently clear, the two populations are regarded as separate species. This differs from evolutionary species in that the parent species goes extinct taxonomically when a new species evolves, the mother and daughter populations now forming two new species. Subspecies as such are not recognized under this approach; either a population is a phylogenetic species or it is not taxonomically distinguishable.

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Other
Ecological species A set of organisms adapted to a particular set of resources, called a niche, in the environment. According to this concept, populations form the discrete phenetic clusters that we recognize as species because the ecological and evolutionary processes controlling how resources are divided up tend to produce those clusters.[] Biological / reproductive species Two organisms that are able to reproduce naturally to produce fertile offspring of both sexes. Organisms that can reproduce but almost always make infertile hybrids of at least one sex, such as a mule, hinny or F1 male cattalo are not considered to be the same species.[citation needed] Biological / isolation species A set of actually or potentially interbreeding populations. This is generally a useful formulation for scientists working with living examples of the higher taxa like mammals, fish, and birds, but more problematic for organisms that do not reproduce sexually. The results of breeding experiments done in artificial conditions may or may not reflect what would happen if the same organisms encountered each other in the wild, making it difficult to gauge whether or not the results of such experiments are meaningful in reference to natural populations.[citation needed] Genetic species Based on similarity of DNA of individuals or populations. Techniques to compare similarity of DNA include DNA-DNA hybridization, and genetic fingerprinting (or DNA barcoding).[citation needed] Cohesion species Most inclusive population of individuals having the potential for phenotypic cohesion through intrinsic cohesion mechanisms. This is an expansion of the mate-recognition species concept to allow for post-mating isolation mechanisms; no matter whether populations can hybridize successfully, they are still distinct cohesion species if the amount of hybridization is insufficient to completely mix their respective gene pools.[citation needed] Evolutionarily Significant Unit (ESU) An evolutionarily significant unit is a population of organisms that is considered distinct for purposes of conservation. Often referred to as a species or a wildlife species, an ESU also has several possible definitions, which coincide with definitions of species.[citation needed] Phenetic species Based on phenetics. Microspecies A species with very little genetic variability, usually one that reproduces by apomixis. Recognition species Based on shared reproductive systems, including mating behavior. The Recognition concept of species has been introduced by Hugh E. H. Paterson, after earlier work by Wilhelm Petersen.[citation needed] Mate-recognition species A group of organisms that are known to recognize one another as potential mates. Like the isolation species concept above, it applies only to organisms that reproduce sexually. Unlike the isolation species concept, it focuses specifically on pre-mating reproductive isolation.[citation needed]

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Numbers of species
Bearing in mind the aforementioned problems with categorizing species, the following numbers are only a guide. Based on various discussions from the first decade of the new millennium, counts can roughly be broken down as follows:[17] Number of prokaryotic Domain Bacteria species,

This number is very difficult to assess, but the discussed range varies from tens of thousands to billions;[18][19][][][20] most recent approaches and studies appear to favor the larger magnitude An estimate of the number of some undiscovered and discovered eukaryotic number.[][21][22] Smaller numbers species.Wikipedia:Verifiability[citation needed] arise from assumptions based on a plateauing of identification of new species (which has technical explanations other than that fewer species remain to be identified).[18] Larger numbers address the fact that success in culturing bacteria has only been achieved in half of identified Bacterial phyla (where lack of success in attempts to culture a bacterial isolate limits abilities to study and delineate new species),[][] and address the difficulty of applying traditional botanic and zoologic definitions of species to asexually reproducing bacteria (where more modern sequencing and molecular approaches support higher species tallies).[][] Number of prokaryotic species, Domain Archaea As a further microbial Domain, the issues and difficulties of Domain Bacteria also pertain to any counting of species of Archaea, all the more given their various extreme habitats. As the Wikipedia article on the Archaebacteria notes (Classification, Species section), the "classification of archaea into species is also controversial" as they also reproduce asexually (likewise eliminating applicability of species definitions based on interbreeding), and face the same difficulties associated with organism isolation and culturing (see citations for Bacteria, above).[][][23] Archaebacteria have been shown to exhibit high rates of horizontal gene transfer (resulting from a bacterial cognate of sex), including between organisms quite separate based on genomic analysis.[24] As the Archaea article notes, "[c]urrent knowledge on genetic diversity is fragmentary and the total number of archaean species cannot be estimated with any accuracy" ... though like Domain Bacteria, the number of cultured and studied phyla relative to the total is low (as of 2005, less than 50% of known phyla cultured).[25] Taken together, very high numbers of unique archaebacterial types are likely, as in the case of Domain Bacteria. Number of eukaryotic species This number has historically varied from a few million to about 100 millions. However these higher numbers, which were based on the potential deep marine and arthropod diversity, are now considered unlikely. The total number of eukaryotic species is likely to be 5 3 million of which about 1.5 million have been already named.[] Some older estimates for various eukaryote phyla are (including some updated numbers): As many as 1.5 million fungi;[] 3,067 brown algae

Species 17,000 lichens; 321,212 plants, including: 10,134 red and green algae 16,236 mosses, 12,000 ferns and horsetails, 1,021 gymnosperms, 281,821 angiosperms; 1,367,555 non-insect animals, including: 1,305,250 invertebrates 2,175 corals 85,000 mollusks As many as 1.1 million arachnids, including ~1 million mites[26] and ~100,000 other arachnids 47,000 crustaceans 68,827 other invertebrates; 63,649 vertebrates 31,300 fish, 7,093 amphibians,[27] 9,768 reptiles,[28] 9,998 birds, 5,490 mammals; As many as 1030 million insects;[29] At present, organisations such as the Global Taxonomy Initiative, the European Distributed Institute of Taxonomy and the Census of Marine Life[30] (the latter only for marine organisms) are trying to improve taxonomy and implement previously undiscovered species to the taxonomy system, although current knowledge covers only a portion of the organisms in the biosphere and thus does not enable a complete understanding of the workings of the environment. Humankind is also currently wiping out undiscovered species at an unprecedented rate,[31] which means that even before a new species has had the chance of being studied and classified, it may already be extinct.

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Importance in biological classification

The idea of species has a long history. It is one of the most important levels of classification, for several reasons: It often corresponds to what lay people treat as the different basic kinds of organism dogs are one species, cats another. It is the standard binomial nomenclature (or trinomial nomenclature) by which scientists typically refer to organisms. It is the highest taxonomic level that cannot be made more or less inclusionary. After years of use, the concept remains central to biology and a host of related fields, and yet also remains at times ill-defined.

Implications of assignment of species status

The naming of a particular species may be regarded as a hypothesis about the evolutionary relationships and distinguishability of that group of organisms. As further information comes to hand, the hypothesis may be confirmed or refuted. Sometimes, especially in the past when communication was more difficult, taxonomists working in isolation have given two distinct names to individual organisms later identified as the same species. When two named species are discovered to be of the same species, the older species name is usually retained, and the newer species name dropped, a process called synonymization, or colloquially, as lumping. Dividing a taxon into

Species multiple, often new, taxons is called splitting. Taxonomists are often referred to as "lumpers" or "splitters" by their colleagues, depending on their personal approach to recognizing differences or commonalities between organisms (see lumpers and splitters). Traditionally, researchers relied on observations of anatomical differences, and on observations of whether different populations were able to interbreed successfully, to distinguish species; both anatomy and breeding behavior are still important to assigning species status. As a result of the revolutionary (and still ongoing) advance in microbiological research techniques, including DNA analysis, in the last few decades, a great deal of additional knowledge about the differences and similarities between species has become available. Many populations formerly regarded as separate species are now considered a single taxon, and many formerly grouped populations have been split. Any taxonomic level (species, genus, family, etc.) can be synonymized or split, and at higher taxonomic levels, these revisions have been still more profound. From a taxonomical point of view, groups within a species can be defined as being of a taxon hierarchically lower than a species. In zoology only the subspecies is used, while in botany the variety, subvariety, and form are used as well. In conservation biology, the concept of evolutionary significant units (ESU) is used, which may define either species or smaller distinct population segments. Identifying and naming species is the providence of alpha taxonomy.

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History and development of the species concept

In the earliest works of science, a species was simply an individual organism that represented a group of similar or nearly identical organisms. No other relationships beyond that group were implied. Aristotle used the words genus and species to mean generic and specific categories. Aristotle and other pre-Darwinian scientists took the species to be distinct and unchanging, with an "essence", like the chemical elements. When early observers began to develop systems of organization for living things, they began to place formerly isolated species into a context. Many of these early delineation schemes would now be considered whimsical and these included consanguinity based on color (all plants with yellow flowers) or behavior (snakes, scorpions and certain biting ants). John Ray (1686), an English naturalist, was the first to give a biological definition of the term species.[32]

John Ray.

In the 18th century Swedish scientist Carolus Linnaeus classified organisms according to differences in the form of reproductive apparatus. Although his system of classification sorts organisms according to degrees of similarity, it made no claims about the relationship between similar species. At that time, it was still widely believed that there was no organic connection between species, no matter how similar they appeared. This approach also suggested a type of idealism: the notion that each species existed as an "ideal form". Although there are always differences (although sometimes minute) between individual organisms, Linnaeus considered such variation problematic. He strove to identify individual organisms that were exemplary of the species, and considered other non-exemplary organisms to be deviant and imperfect.

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By the 19th century most naturalists understood that species could change form over time, and that the history of the planet provided enough time for major changes. Jean-Baptiste Lamarck, in his 1809 Zoological Philosophy, offered one of the first logical arguments against creationism. The new emphasis was on determining how a species could change over time. Lamarck suggested that an organism could pass on an acquired trait to its offspring, i.e., the giraffe's long neck was attributed to generations of giraffes stretching to reach the leaves of higher treetops (this well-known and simplistic example, however, does not do justice to the breadth and subtlety of Lamarck's ideas). With the acceptance of the natural selection idea of Charles Darwin in the 1860s, however, Lamarck's view of goal-oriented evolution, also known as a teleological process, was eclipsed. Recent interest in inheritance of acquired characteristics centers around Linnaeus believed in the fixity of species. epigenetic processes, e.g. methylation, that do not affect DNA sequences, but instead alter expression in an inheritable manner. Thus, Neo-Lamarckism, as it is sometimes termed, is not a challenge to the theory of evolution by natural selection. Charles Darwin and Alfred Wallace provided what scientists now consider as the most powerful and compelling theory of evolution. Darwin argued that it was populations that evolved, not individuals. His argument relied on a radical shift in perspective from that of Linnaeus: rather than defining species in ideal terms (and searching for an ideal representative and rejecting deviations), Darwin considered variation among individuals to be natural. He further argued that variation, far from being problematic, actually provides the explanation for the existence of distinct species. Darwin's work drew on Thomas Malthus' insight that the rate of growth of a biological population will always outpace the rate of growth of the resources in the environment, such as the food supply. As a result, Darwin argued, not all the members of a population will be able to survive and reproduce. Those that did will, on average, be the ones possessing variationshowever slightthat make them slightly better adapted to the environment. If these variable traits are heritable, then the offspring of the survivors will also possess them. Thus, over many generations, adaptive variations will accumulate in the population, while counter-adaptive traits will tend to be eliminated. Whether a variation is adaptive or non-adaptive depends on the environment: different environments favor different traits. Since the environment effectively selects which organisms live to reproduce, it is the environment (the "fight for existence") that selects the traits to be passed on. This is the theory of evolution by natural selection. In this model, the length of a giraffe's neck would be explained by positing that proto-giraffes with longer necks would have had a significant reproductive advantage to those with shorter necks. Over many generations, the entire population would be a species of long-necked animals. In 1859, when Darwin published his theory of natural selection, the mechanism behind the inheritance of individual traits was unknown. Although Darwin made some speculations on how traits are inherited (pangenesis), his theory relies only on the fact that inheritable traits exist, and are variable (which makes his accomplishment even more remarkable.) Although Gregor Mendel's paper on genetics was published in 1866, its significance was not recognized. It was not until 1900 that his work was rediscovered by Hugo de Vries, Carl Correns and Erich von Tschermak, who realised that the "inheritable traits" in Darwin's theory are genes. The theory of the evolution of species through natural selection has two important implications for discussions of speciesconsequences that fundamentally challenge the assumptions behind Linnaeus' taxonomy. First, it suggests that species are not just similar, they may actually be related. Some students of Darwin argue that all species are descended from a common ancestor. Second, it supposes that "species" are not homogeneous, fixed, permanent things; members of a species are all different, and over time species change. This suggests that species do not have

Species any clear boundaries but are rather momentary statistical effects of constantly changing gene-frequencies. One may still use Linnaeus' taxonomy to identify individual plants and animals, but one can no longer think of species as independent and immutable. The rise of a new species from a parental line is called speciation. There is no clear line demarcating the ancestral species from the descendant species. Although the current scientific understanding of species suggests that there is no rigorous and comprehensive way to distinguish between different species in all cases, biologists continue to seek concrete ways to operationalize the idea. One of the most popular biological definitions of species is in terms of reproductive isolation; if two creatures cannot reproduce to produce fertile offspring of both sexes, then they are in different species. This definition captures a number of intuitive species boundaries, but it remains imperfect. It has nothing to say about species that reproduce asexually, for example, and it is very difficult to apply to extinct species. Moreover, boundaries between species are often fuzzy: there are examples where members of one population can produce fertile offspring of both sexes with a second population, and members of the second population can produce fertile offspring of both sexes with members of a third population, but members of the first and third population cannot produce fertile offspring, or can only produce fertile offspring of the homozygous sex. Consequently, some people reject this definition of a species. Richard Dawkins defines two organisms as conspecific if and only if they have the same number of chromosomes and, for each chromosome, both organisms have the same number of nucleotides (The Blind Watchmaker, p.118). However, most if not all taxonomists would strongly disagree[citation needed]. For example, in many amphibians, most notably in New Zealand's Leiopelma frogs, the genome consists of "core" chromosomes that are mostly invariable and accessory chromosomes, of which exist a number of possible combinations. Even though the chromosome numbers are highly variable between populations, these can interbreed successfully and form a single evolutionary unit. In plants, polyploidy is extremely commonplace with few restrictions on interbreeding; as individuals with an odd number of chromosome sets are usually sterile, depending on the actual number of chromosome sets present, this results in the odd situation where some individuals of the same evolutionary unit can interbreed with certain others and some cannot, with all populations being eventually linked as to form a common gene pool. The classification of species has been profoundly affected by technological advances that have allowed researchers to determine relatedness based on molecular markers, starting with the comparatively crude blood plasma precipitation assays in the mid-20th century to Charles Sibley's ground-breaking DNA-DNA hybridization studies in the 1970s leading to DNA sequencing techniques. The results of these techniques caused revolutionary changes in the higher taxonomic categories (such as phyla and classes), resulting in the reordering of many branches of the phylogenetic tree (see also: molecular phylogeny). For taxonomic categories below genera, the results have been mixed so far; the pace of evolutionary change on the molecular level is rather slow, yielding clear differences only after considerable periods of reproductive separation. DNA-DNA hybridization results have led to misleading conclusions, the Pomarine Skua Great Skua phenomenon being a famous example. Turtles have been determined to evolve with just one-eighth of the speed of other reptiles on the molecular level, and the rate of molecular evolution in albatrosses is half of what is found in the rather closely related storm-petrels. The hybridization technique is now obsolete and is replaced by more reliable computational approaches for sequence comparison. Molecular taxonomy is not directly based on the evolutionary processes, but rather on the overall change brought upon by these processes. The processes that lead to the generation and maintenance of variation such as mutation, crossover and selection are not uniform (see also molecular clock). DNA is only extremely rarely a direct target of natural selection rather than changes in the DNA sequence enduring over generations being a result of the latter; for example, silent transition-transversion combinations would alter the melting point of the DNA sequence, but not the sequence of the encoded proteins and thus are a possible example where, for example in microorganisms, a mutation confers a change in fitness all by itself.

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Notes and references

[7] Hopf FA, Hopf FW. (1985). The role of the Allee effect on species packing. Theor. Pop. Biol. 27, 27-50. [18] Counting in a bacterial world (http:/ / www. newscientist. com/ blog/ shortsharpscience/ 2008/ 06/ counting-in-bacterial-world. html). New Scientist, June 10, 2008. [27] http:/ / www. amphibiaweb. org Amphibiaweb, accessed 27 Feb 2013 [28] http:/ / www. reptile-database. org The Reptile Database, accessed 27 Feb 2013 [32] Historia plantarum generalis, in the volume published in 1686, Tome I, Libr. I, Chap. XX, page 40 (Quoted in Mayr, Ernst. 1982. The growth of biological thought: diversity, evolution, and inheritance. Cambridge, Mass.: Belknap Press: 256)

External links
Stanford Encyclopedia of Philosophy entry: Species (http://plato.stanford.edu/entries/species/) Barcoding of species (http://www.barcodinglife.org/) European Species Names in Linnaean, Czech, English, German and French (http://www.finitesite.com/ dandelion/Linnaeus.HTML) Catalogue of Life (http://www.catalogueoflife.org/) VisualTaxa (http://visualtaxa.redgolpe.com/) Speciation (http://users.rcn.com/jkimball.ma.ultranet/BiologyPages/S/Speciation.html) Articles online Other Species Concepts (http://evolution.berkeley.edu/evosite/evo101/VA2OtherSpeciesConcept.shtml) U.C. Berkeley 2003-12-31, ScienceDaily: Working On The 'Porsche Of Its Time': New Model For Species Determination Offered (http://www.sciencedaily.com/releases/2003/12/031231082553.htm) 2003-08-08, ScienceDaily: Cross-species Mating May Be Evolutionarily Important And Lead To Rapid Change (http://www.sciencedaily.com/releases/2003/08/030808081854.htm) 2004-01-09 ScienceDaily: Mayo Researchers Observe Genetic Fusion Of Human, Animal Cells; May Help Explain Origin Of AIDS (http://www.sciencedaily.com/releases/2004/01/040109064407.htm) 2000-09-18, ScienceDaily: Scientists Unravel Ancient Evolutionary History Of Photosynthesis (http://www. sciencedaily.com/releases/2000/09/000913211733.htm)

Natural selection

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Natural selection
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Evolutionary biology

Diagrammatic representation of the divergence of modern taxonomic groups from their common ancestor.

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Natural selection is the gradual natural process by which biological traits become either more or less common in a population as a function of the effect of inherited traits on the differential reproductive success of organisms interacting with their environment. It is a key mechanism of evolution. The term "natural selection" was popularized by Charles Darwin who intended it to be compared with artificial selection, which is now called selective breeding. Variation exists within all populations of organisms. This occurs partly because random mutations occur in the genome of an individual organism, and these mutations can be passed to offspring. Throughout the individuals lives, their genomes interact with their environments to cause variations in traits. (The environment of a genome includes the molecular biology in the cell, other cells, other individuals, populations, species, as well as the abiotic environment.) Individuals with certain variants of the trait may survive and reproduce more than individuals with other variants. Therefore the population evolves. Factors that affect reproductive success are also important, an issue that Charles Darwin developed in his ideas on sexual selection, for example. Natural selection acts on the phenotype, or the observable characteristics of an organism, but the genetic (heritable) basis of any phenotype that gives a reproductive advantage may become more common in a population (see allele frequency). Over time, this process can result in populations that specialize for particular ecological niches and may eventually result in the emergence of new species. In other words, natural selection is an important process (though not the only process) by which evolution takes place within a population of organisms. Natural selection can be contrasted with artificial selection, in which humans intentionally choose specific traits (although they may not always get what they want). In natural selection there is no intentional choice. In other words, artificial selection is teleological and natural selection is not teleological. Natural selection is one of the cornerstones of modern biology. The term was introduced by Darwin in his influential 1859 book On the Origin of Species,[1] in which natural selection was described as analogous to artificial selection, a process by which animals and plants with traits considered desirable by human breeders are systematically favored

Natural selection for reproduction. The concept of natural selection was originally developed in the absence of a valid theory of heredity; at the time of Darwin's writing, nothing was known of modern genetics. The union of traditional Darwinian evolution with subsequent discoveries in classical and molecular genetics is termed the modern evolutionary synthesis. Natural selection remains the primary explanation for adaptive evolution.

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General principles
Natural variation occurs among the individuals of any population of organisms. Many of these differences do not affect survival, but some differences may improve the chances of survival of a particular individual. A rabbit that runs faster than others may be more likely to escape from predators, and algae that are more efficient at extracting energy from sunlight will grow faster. Something that increases an animal's survival will often also include its reproductive rate; however, sometimes there is a trade-off between survival and current reproduction. Ultimately, what matters is total lifetime reproduction of the animal.

For example, the peppered moth exists in both light and dark colors in the United Kingdom, but during the industrial revolution, many of the trees on which the moths rested became blackened by soot, giving the dark-colored moths an advantage in hiding from predators. This gave dark-colored moths a better chance of surviving to produce dark-colored offspring, and in just fifty years from the first dark moth being caught, nearly all of the moths in industrial Manchester were dark. The balance was reversed by the effect of the Clean Air Act 1956, and the dark moths became rare again, demonstrating the influence of natural selection on peppered moth evolution.[] If the traits that give these individuals a reproductive advantage are also heritable, that is, passed from parent to child, then there will be a slightly higher proportion of fast rabbits or efficient algae in the next generation. This is known as differential reproduction. Even if the reproductive advantage is very slight, over many generations any heritable advantage will become dominant in the population. In this way the natural environment of an organism "selects" for traits that confer a reproductive advantage, causing gradual changes or evolution of life. This effect was first described and named by Charles Darwin. The concept of natural selection predates the understanding of genetics, the mechanism of heredity for all known life forms. In modern terms, selection acts on an organism's phenotype, or observable characteristics, but it is the organism's genetic make-up or genotype that is inherited. The phenotype is the result of the genotype and the environment in which the organism lives (see Genotype-phenotype distinction). This is the link between natural selection and genetics, as described in the modern evolutionary synthesis. Although a complete theory of evolution also requires an account of how genetic variation arises in the first place (such as by mutation and sexual reproduction) and includes other evolutionary mechanisms (such as genetic drift and gene flow), natural selection appears to be the most important mechanism for creating complex adaptations in nature.

Morpha typica and morpha carbonaria, morphs of the peppered moth resting on the same tree. The light-colored morpha typica (below the bark's scar) is hard to see on this pollution-free tree, camouflaging it from predators such as Great Tits.

Nomenclature and usage

The term natural selection has slightly different definitions in different contexts. It is most often defined to operate on heritable traits, because these are the traits that directly participate in evolution. However, natural selection is "blind" in the sense that changes in phenotype (physical and behavioral characteristics) can give a reproductive advantage regardless of whether or not the trait is heritable (non heritable traits can be the result of environmental factors or the life experience of the organism).

Natural selection Following Darwin's primary usage[1] the term is often used to refer to both the evolutionary consequence of blind selection and to its mechanisms.[2][3] It is sometimes helpful to explicitly distinguish between selection's mechanisms and its effects; when this distinction is important, scientists define "natural selection" specifically as "those mechanisms that contribute to the selection of individuals that reproduce", without regard to whether the basis of the selection is heritable. This is sometimes referred to as "phenotypic natural selection".[4] Traits that cause greater reproductive success of an organism are said to be selected for, whereas those that reduce success are selected against. Selection for a trait may also result in the selection of other correlated traits that do not themselves directly influence reproductive advantage. This may occur as a result of pleiotropy or gene linkage.[5]

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Fitness
The concept of fitness is central to natural selection. In broad terms, individuals that are more "fit" have better potential for survival, as in the well-known phrase "survival of the fittest". However, as with natural selection above, the precise meaning of the term is much more subtle. Modern evolutionary theory defines fitness not by how long an organism lives, but by how successful it is at reproducing. If an organism lives half as long as others of its species, but has twice as many offspring surviving to adulthood, its genes will become more common in the adult population of the next generation. Though natural selection acts on individuals, the effects of chance mean that fitness can only really be defined "on average" for the individuals within a population. The fitness of a particular genotype corresponds to the average effect on all individuals with that genotype. Very low-fitness genotypes cause their bearers to have few or no offspring on average; examples include many human genetic disorders like cystic fibrosis.

Darwin's illustrations of beak variation in the finches of the Galpagos Islands, which hold 13 closely related species that differ most markedly in the shape of their beaks. The beak of each species is suited to its preferred food, suggesting that beak shapes evolved by natural selection.

Since fitness is an averaged quantity, it is also possible that a favorable mutation arises in an individual that does not survive to adulthood for unrelated reasons. Fitness also depends crucially upon the environment. Conditions like sickle-cell anemia may have low fitness in the general human population, but because the sickle-cell trait confers immunity from malaria, it has high fitness value in populations that have high malaria infection rates.

Types of selection
Natural selection can act on any heritable phenotypic trait, and selective pressure can be produced by any aspect of the environment, including sexual selection and competition with members of the same or other species. However, this does not imply that natural selection is always directional and results in adaptive evolution; natural selection often results in the maintenance of the status quo by eliminating less fit variants. The unit of selection can be the individual or it can be another level within the hierarchy of biological organisation, such as genes, cells, and kin groups. There is still debate about whether natural selection acts at the level of groups or species to produce adaptations that benefit a larger, non-kin group. Likewise, there is debate as to whether selection at the molecular level prior to gene mutations and fertilization of the zygote should be ascribed to conventional natural selection because traditionally natural selection is an environmental and exterior force that acts upon a phenotype typically after birth. Some science journalists distinguish gene selection from natural selection by informally referencing selection of mutations as "pre-selection."[6] Selection at a different level such as the gene can result in an increase in fitness for that gene, while at the same time reducing the fitness of the individuals carrying that gene, in a process called intragenomic conflict. Overall, the

Natural selection combined effect of all selection pressures at various levels determines the overall fitness of an individual, and hence the outcome of natural selection. Natural selection occurs at every life stage of an individual. An individual organism must survive until adulthood before it can reproduce, and selection of those that reach this stage is called viability selection. In many species, adults must compete with each other for mates via sexual selection, and success in this competition determines who will parent the next generation. When individuals can reproduce more than once, a longer survival in the reproductive phase increases the number of offspring, called survival selection. The fecundity of both females and males (for example, giant sperm in certain species of Drosophila)[8] can be limited via "fecundity selection". The viability of produced gametes can differ, while intragenomic conflicts such as meiotic drive between the haploid gametes can result in gametic or "genic selection". Finally, the union of some combinations of eggs and sperm might be more compatible than others; this is termed compatibility selection.

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The life cycle of a sexually reproducing organism. Various components of natural [7] selection are indicated for each life stage.

Sexual selection
It is useful to distinguish between "ecological selection" and "sexual selection". Ecological selection covers any mechanism of selection as a result of the environment (including relatives, e.g. kin selection, competition, and infanticide), while "sexual selection" refers specifically to competition for mates.[9] Sexual selection can be intrasexual, as in cases of competition among individuals of the same sex in a population, or intersexual, as in cases where one sex controls reproductive access by choosing among a population of available mates. Most commonly, intrasexual selection involves malemale competition and intersexual selection involves female choice of suitable males, due to the generally greater investment of resources for a female than a male in a single offspring. However, some species exhibit sex-role reversed behavior in which it is males that are most selective in mate choice; the best-known examples of this pattern occur in some fishes of the family Syngnathidae, though likely examples have also been found in amphibian and bird species.[] Some features that are confined to one sex only of a particular species can be explained by selection exercised by the other sex in the choice of a mate, for example, the extravagant plumage of some male birds. Similarly, aggression between members of the same sex is sometimes associated with very distinctive features, such as the antlers of stags, which are used in combat with other stags. More generally, intrasexual selection is often associated with sexual dimorphism, including differences in body size between males and females of a species.[]

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Examples of natural selection

A well-known example of natural selection in action is the development of antibiotic resistance in microorganisms. Since the discovery of penicillin in 1928, antibiotics have been used to fight bacterial diseases. Natural populations of bacteria contain, among their vast numbers of individual members, considerable variation in their genetic material, primarily as the result of mutations. When exposed to antibiotics, most bacteria die quickly, but some may have mutations that make them slightly less susceptible. If the exposure to antibiotics is short, these individuals will survive the treatment. This selective elimination of maladapted individuals from a population is natural selection. These surviving bacteria will then reproduce again, producing the next generation. Due to the elimination of the maladapted individuals in the past generation, this population contains more bacteria that have some resistance against the antibiotic. At the same time, new mutations occur, contributing new genetic variation to the existing genetic variation. Spontaneous mutations are very rare, and advantageous mutations are even rarer. However, populations of bacteria are large enough that a few individuals will have beneficial mutations. If a new mutation reduces their susceptibility to an antibiotic, these individuals are more likely to survive when next confronted with that antibiotic.

Given enough time and repeated exposure to the antibiotic, a population of antibiotic-resistant bacteria will emerge. This new changed population of antibiotic-resistant bacteria is optimally adapted to the context it evolved in. At the same time, it is not necessarily optimally adapted any more to the old antibiotic free environment. The end result of natural selection is two populations that are both optimally adapted to their specific environment, while both perform substandard in the other environment. The widespread use and misuse of antibiotics has resulted in increased microbial resistance to antibiotics in clinical use, to the point that the methicillin-resistant Staphylococcus aureus (MRSA) has been described as a "superbug" because of the threat it poses to health and its relative invulnerability to existing drugs.[10] Response strategies typically include the use of different, stronger antibiotics; however, new strains of MRSA have recently emerged that are resistant even to these drugs.[11] This is an example of what is known as an evolutionary arms race, in which bacteria continue to develop strains that are less susceptible to antibiotics, while medical researchers continue to develop new antibiotics that can kill them. A similar situation occurs with pesticide resistance in plants and insects. Arms races are not necessarily induced by man; a well-documented example involves the spread of a gene in the butterfly Hypolimnas bolina suppressing male-killing activity by Wolbachia bacteria parasites on the island of Samoa, where the spread of the gene is known to have occurred over a period of just five years [12]

Resistance to antibiotics is increased though the survival of individuals that are immune to the effects of the antibiotic, whose offspring then inherit the resistance, creating a new population of resistant bacteria.

Evolution by means of natural selection

A prerequisite for natural selection to result in adaptive evolution, novel traits and speciation, is the presence of heritable genetic variation that results in fitness differences. Genetic variation is the result of mutations, recombinations and alterations in the karyotype (the number, shape, size and internal arrangement of the chromosomes). Any of these changes might have an effect that is highly advantageous or highly disadvantageous, but large effects are very rare. In the past, most changes in the genetic material were considered neutral or close to neutral because they occurred in noncoding DNA or resulted in a synonymous substitution. However, recent research suggests that many mutations in non-coding DNA do have slight deleterious effects.[][13] Although both mutation

Natural selection rates and average fitness effects of mutations are dependent on the organism, estimates from data in humans have found that a majority of mutations are slightly deleterious.[] By the definition of fitness, individuals with greater fitness are more likely to contribute offspring to the next generation, while individuals with lesser fitness are more likely to die early or fail to reproduce. As a result, alleles that on average result in greater fitness become more abundant in the next generation, while alleles that in general reduce fitness become rarer. If the selection forces remain the same for many generations, beneficial alleles become more and more abundant, until they dominate the population, while alleles with a lesser fitness disappear. In every generation, new mutations and re-combinations arise spontaneously, producing a new spectrum of phenotypes. Therefore, each new generation will be enriched by the increasing abundance of alleles that contribute to those traits that were favored by selection, enhancing these traits over successive generations.

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Some mutations occur in so-called regulatory genes. Changes in these can have large effects on the phenotype of the individual because they regulate the function of many other genes. Most, but not all, mutations in regulatory genes result in non-viable zygotes. Examples of nonlethal regulatory mutations occur in HOX genes in humans, which can result in a cervical rib[14] or polydactyly, an increase in the number of fingers or toes.[15] When such mutations result in a higher fitness, natural selection will favor these phenotypes and the novel trait will spread in the population. Established traits are not immutable; traits that have high fitness in one environmental context may be much less fit if environmental conditions change. In the absence of natural selection to preserve such a trait, it will become more variable and deteriorate over time, possibly resulting in a vestigial manifestation of the trait, also called evolutionary baggage. In many circumstances, the apparently vestigial structure may retain a limited functionality, or may be co-opted for other advantageous traits in a phenomenon known as preadaptation. A famous example of a vestigial structure, the eye of the blind mole rat, is believed to retain function in photoperiod perception.[]

The exuberant tail of the peacock is thought to be the result of sexual selection by females. This peacock is leucistic; selection against leucism and albinism in nature is intense because they are easily spotted by predators or are unsuccessful in competition for mates.

Speciation

Speciation requires selective mating, which result in a reduced gene flow. Selective mating can be the result of 1. Geographic isolation, 2. Behavioral isolation, or 3. Temporal isolation. For example, a change in the physical environment (geographic isolation by an extrinsic barrier) would follow number 1, a change in camouflage for number 2 or a shift in mating times (i.e., one species of deer shifts location and therefore changes its "rut") for number 3.[citation needed] Over time, these subgroups might diverge radically to become different species, either because of differences in selection pressures on the different subgroups, or because different mutations arise spontaneously in the different populations, or because of founder effects some potentially beneficial alleles may, by chance, be present in only one or other of two subgroups when they first become separated. A lesser-known mechanism of speciation occurs via hybridization, well-documented in plants and occasionally observed in species-rich groups of animals such as cichlid fishes.[] Such mechanisms of rapid speciation can reflect a mechanism of evolutionary change known as punctuated equilibrium, which suggests that evolutionary change and in particular speciation typically happens quickly after interrupting long periods of stasis.

X-ray of the left hand of a ten year old boy with polydactyly.

Natural selection Genetic changes within groups result in increasing incompatibility between the genomes of the two subgroups, thus reducing gene flow between the groups. Gene flow will effectively cease when the distinctive mutations characterizing each subgroup become fixed. As few as two mutations can result in speciation: if each mutation has a neutral or positive effect on fitness when they occur separately, but a negative effect when they occur together, then fixation of these genes in the respective subgroups will lead to two reproductively isolated populations. According to the biological species concept, these will be two different species.

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Historical development
Pre-Darwinian theories
Several ancient philosophers expressed the idea that nature produces a huge variety of creatures, randomly, and that only those creatures that manage to provide for themselves and reproduce successfully survive; well-known examples include Empedocles[16] and his intellectual successor, the Roman poet Lucretius.[17] Empedocles' idea that organisms arose entirely by the incidental workings of causes such as heat and cold was criticized by Aristotle in Book II of Physics.[18] He posited natural teleology in its place. He believed that form was achieved for a purpose, citing the regularity of heredity in species as proof.[][] Nevertheless, he acceded that new types of animals, monstrosities (), can occur in very rare instances (Generation of Animals, Book IV).[] As quoted in Darwin's Origin of Species, Aristotle considered whether different forms, e.g. of teeth, might have appeared accidentally, but only the useful forms survived:

The modern theory of natural selection derives from the work of Charles Darwin in the nineteenth century.

So what hinders the different parts (of the body) from having this merely accidental relation in nature? as the teeth, for example, grow by necessity, the front ones sharp, adapted for dividing, and the grinders flat, and serviceable for masticating the food; since they were not made for the sake of this, but it was the result of accident. And in like manner as to other parts in which there appears to exist an adaptation to an end. Wheresoever, therefore, all things together (that is all the parts of one whole) happened like as if they were made for the sake of something, these were preserved, having been appropriately constituted by an internal spontaneity; and whatsoever things were not thus constituted, perished and still perish. Aristotle, Physicae Auscultationes(lib.2, cap.8, s.2) But he rejected this possibility in the next paragraph: ...Yet it is impossible that this should be the true view. For teeth and all other natural things either invariably or normally come about in a given way; but of not one of the results of chance or spontaneity is this true. We do not ascribe to chance or mere coincidence the frequency of rain in winter, but frequent rain in summer we do; nor heat in the dog-days, but only if we have it in winter. If then, it is agreed that things are either the result of coincidence or for an end, and these cannot be the result of coincidence or spontaneity, it follows that they must be for an end; and that such things are all due to nature even the champions of the theory which is before us would agree. Therefore action for an end is present in things which come to be and are by nature. Aristotle,Aristotle, Physicae Auscultationes(lib.2, cap.8, s.2) The struggle for existence was later described by Islamic writer Al-Jahiz in the 9th century, who argued that environmental factors influence animals to develop new characteristics to ensure [19][20][21] survival. Wikipedia:Verifiability

Natural selection The classical arguments were reintroduced in the 18th century by Pierre Louis Maupertuis[22] and others, including Charles Darwin's grandfather Erasmus Darwin. While these forerunners had an influence on Darwinism, they later had little influence on the trajectory of evolutionary thought after Charles Darwin. Until the early 19th century, the prevailing view in Western societies was that differences between individuals of a species were uninteresting departures from their Platonic idealism (or typus) of created kinds. However, the theory of uniformitarianism in geology promoted the idea that simple, weak forces could act continuously over long periods of time to produce radical changes in the Earth's landscape. The success of this theory raised awareness of the vast scale of geological time and made plausible the idea that tiny, virtually imperceptible changes in successive generations could produce consequences on the scale of differences between species. Early 19th-century evolutionists such as Jean Baptiste Lamarck suggested the inheritance of acquired characteristics as a mechanism for evolutionary change; adaptive traits acquired by an organism during its lifetime could be inherited by that organism's progeny, eventually causing transmutation of species.[23] This theory has come to be known as Lamarckism and was an influence on the anti-genetic ideas of the Stalinist Soviet biologist Trofim Lysenko.[]

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Darwin's theory
In 1859, Charles Darwin set out his theory of evolution by natural selection as an explanation for adaptation and speciation. He defined natural selection as the "principle by which each slight variation [of a trait], if useful, is preserved".[24] The concept was simple but powerful: individuals best adapted to their environments are more likely to survive and reproduce. As long as there is some variation between them, there will be an inevitable selection of individuals with the most advantageous variations. If the variations are inherited, then differential reproductive success will lead to a progressive evolution of particular populations of a species, and populations that evolve to be sufficiently different eventually become different species.[25] Darwin's ideas were inspired by the observations that he had made on the Beagle voyage, and by the work of a political economist, the Reverend Thomas Malthus, who in An Essay on the Principle of Population, noted that population (if unchecked) increases exponentially, whereas the food supply grows only arithmetically; thus, inevitable limitations of resources would have demographic implications, leading to a "struggle for existence".[] When Darwin read Malthus in 1838 he was already primed by his work as a naturalist to appreciate the "struggle for existence" in nature and it struck him that as population outgrew resources, "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 is Darwin's own summary of the idea, which can be found in the fourth chapter of the Origin: If during the long course of ages and under varying conditions of life, organic beings vary at all in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the high geometrical powers of increase of each species, at some age, season, or year, a severe struggle for life, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of all organic beings to each other and to their conditions of existence, causing an infinite diversity in structure, constitution, and habits, to be advantageous to them, I think it would be a most extraordinary fact if no variation ever had occurred useful to each being's own welfare, in the same way as so many variations have occurred useful to man. But, if variations useful to any organic being do occur, assuredly individuals thus characterised will have the best chance of being preserved in the struggle for life; and from the strong principle of inheritance they will tend to produce offspring similarly characterised. This principle of preservation, I have called, for the sake of brevity, Natural Selection. Once he had his theory "by which to work", Darwin was meticulous about gathering and refining evidence as his "prime hobby" before making his idea public. He was in the process of writing his "big book" to present his researches when the naturalist Alfred Russel Wallace independently conceived of the principle and described it in an essay he sent to Darwin to forward to Charles Lyell. Lyell and Joseph Dalton Hooker decided (without Wallace's

Natural selection knowledge) to present his essay together with unpublished writings that Darwin had sent to fellow naturalists, and On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection was read to the Linnean Society announcing co-discovery of the principle in July 1858.[26] Darwin published a detailed account of his evidence and conclusions in On the Origin of Species in 1859. In the 3rd edition of 1861 Darwin acknowledged that others a notable one being William Charles Wells in 1813, and Patrick Matthew in 1831 had proposed similar ideas, but had neither developed them nor presented them in notable scientific publications.[27] Darwin thought of natural selection by analogy to how farmers select crops or livestock for breeding, which he called "artificial selection"; in his early manuscripts he referred to a Nature, which would do the selection. At the time, other mechanisms of evolution such as evolution by genetic drift were not yet explicitly formulated, and Darwin believed that selection was likely only part of the story: "I am convinced that [it] has been the main, but not exclusive means of modification."[28] In a letter to Charles Lyell in September 1860, Darwin regretted the use of the term "Natural Selection", preferring the term "Natural Preservation".[29] For Darwin and his contemporaries, natural selection was in essence synonymous with evolution by natural selection. After the publication of On the Origin of Species, educated people generally accepted that evolution had occurred in some form. However, natural selection remained controversial as a mechanism, partly because it was perceived to be too weak to explain the range of observed characteristics of living organisms, and partly because even supporters of evolution balked at its "unguided" and non-progressive nature,[30] a response that has been characterized as the single most significant impediment to the idea's acceptance.[31] However, some thinkers enthusiastically embraced natural selection; after reading Darwin, Herbert Spencer introduced the term survival of the fittest, which became a popular summary of the theory.[32] The fifth edition of On the Origin of Species published in 1869 included Spencer's phrase as an alternative to natural selection, with credit given: "But the expression often used by Mr. Herbert Spencer, of the Survival of the Fittest, is more accurate, and is sometimes equally convenient."[33] Although the phrase is still often used by non-biologists, modern biologists avoid it because it is tautological if "fittest" is read to mean "functionally superior" and is applied to individuals rather than considered as an averaged quantity over populations.[34]

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Modern evolutionary synthesis

Natural selection relies crucially on the idea of heredity, but it was developed long before the basic concepts of genetics. Although the Austrian monk Gregor Mendel, the father of modern genetics, was a contemporary of Darwin's, his work would lie in obscurity until the early 20th century. Only after the integration of Darwin's theory of evolution with a complex statistical appreciation of Gregor Mendel's 're-discovered' laws of inheritance did natural selection become generally accepted by scientists. The work of Ronald Fisher (who developed the required mathematical language and The Genetical Theory of Natural Selection),[2] J.B.S. Haldane (who introduced the concept of the "cost" of natural selection),[35] Sewall Wright (who elucidated the nature of selection and adaptation),[36] Theodosius Dobzhansky (who established the idea that mutation, by creating genetic diversity, supplied the raw material for natural selection: see Genetics and the Origin of Species),[37] William Hamilton (who conceived of kin selection), Ernst Mayr (who recognised the key importance of reproductive isolation for speciation: see Systematics and the Origin of Species)[38] and many others formed the modern evolutionary synthesis. This synthesis cemented natural selection as the foundation of evolutionary theory, where it remains today.

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Impact of the idea

Darwin's ideas, along with those of Adam Smith and Karl Marx, had a profound influence on 19th century thought. Perhaps the most radical claim of the theory of evolution through natural selection is that "elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner" evolved from the simplest forms of life by a few simple principles. This claim inspired some of Darwin's most ardent supportersand provoked the most profound opposition. The radicalism of natural selection, according to Stephen Jay Gould,[39] lay in its power to "dethrone some of the deepest and most traditional comforts of Western thought". In particular, it challenged long-standing beliefs in such concepts as a special and exalted place for humans in the natural world and a benevolent creator whose intentions were reflected in nature's order and design. In the words of the philosopher Daniel Dennett,[40] "Darwin's dangerous idea" of evolution by natural selection is a "universal acid," which cannot be kept restricted to any vessel or container, as it soon leaks out, working its way into ever-wider surroundings. Thus, in the last decades, the concept of natural selection has spread from evolutionary biology into virtually all disciplines, including evolutionary computation, quantum darwinism, evolutionary economics, evolutionary epistemology, evolutionary psychology, and cosmological natural selection. This unlimited applicability has been called Universal Darwinism.

Cell and molecular biology

In the 19th century, Wilhelm Roux, a founder of modern embryology, wrote a book entitled Der Kampf der Teile im Organismus (The struggle of parts in the organism) in which he suggested that the development of an organism results from a Darwinian competition between the parts of the embryo, occurring at all levels, from molecules to organs. In recent years, a modern version of this theory has been proposed by Jean-Jacques Kupiec. According to this cellular Darwinism [41], stochasticity at the molecular level generates diversity in cell types whereas cell interactions impose a characteristic order on the developing embryo.

Social and psychological theory

The social implications of the theory of evolution by natural selection also became the source of continuing controversy. Friedrich Engels, a German political philosopher and co-originator of the ideology of communism, wrote in 1872 that "Darwin did not know what a bitter satire he wrote on mankind when he showed that free competition, the struggle for existence, which the economists celebrate as the highest historical achievement, is the normal state of the animal kingdom".[42] Interpretation of natural selection as necessarily 'progressive', leading to increasing 'advances' in intelligence and civilisation, was used as a justification for colonialism and policies of eugenics, as well as broader sociopolitical positions now described as Social Darwinism. Konrad Lorenz won the Nobel Prize in Physiology or Medicine in 1973 for his analysis of animal behavior in terms of the role of natural selection (particularly group selection). However, in Germany in 1940, in writings that he subsequently disowned, he used the theory as a justification for policies of the Nazi state. He wrote "... selection for toughness, heroism, and social utility...must be accomplished by some human institution, if mankind, in default of selective factors, is not to be ruined by domestication-induced degeneracy. The racial idea as the basis of our state has already accomplished much in this respect."[43] Others have developed ideas that human societies and culture evolve by mechanisms that are analogous to those that apply to evolution of species.[44] More recently, work among anthropologists and psychologists has led to the development of sociobiology and later evolutionary psychology, a field that attempts to explain features of human psychology in terms of adaptation to the ancestral environment. The most prominent such example, notably advanced in the early work of Noam Chomsky and later by Steven Pinker, is the hypothesis that the human brain is adapted to acquire the grammatical rules of natural language.[45] Other aspects of human behavior and social structures, from specific cultural norms such as incest avoidance to broader patterns such as gender roles, have been hypothesized to have similar origins as adaptations to the early environment in which modern humans evolved. By analogy to the action of natural selection

Natural selection on genes, the concept of memes "units of cultural transmission", or culture's equivalents of genes undergoing selection and recombination has arisen, first described in this form by Richard Dawkins[46] and subsequently expanded upon by philosophers such as Daniel Dennett as explanations for complex cultural activities, including human consciousness.[47]

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Information and systems theory

In 1922, Alfred Lotka proposed that natural selection might be understood as a physical principle that could be described in terms of the use of energy by a system,[48] a concept that was later developed by Howard Odum as the maximum power principle whereby evolutionary systems with selective advantage maximise the rate of useful energy transformation. Such concepts are sometimes relevant in the study of applied thermodynamics. The principles of natural selection have inspired a variety of computational techniques, such as "soft" artificial life, that simulate selective processes and can be highly efficient in 'adapting' entities to an environment defined by a specified fitness function.[49] For example, a class of heuristic optimization algorithms known as genetic algorithms, pioneered by John Holland in the 1970s and expanded upon by David E. Goldberg,[50] identify optimal solutions by simulated reproduction and mutation of a population of solutions defined by an initial probability distribution.[51] Such algorithms are particularly useful when applied to problems whose solution landscape is very rough or has many local minima.

Genetic basis of natural selection

The idea of natural selection predates the understanding of genetics. We now have a much better idea of the biology underlying heritability, which is the basis of natural selection.

Genotype and phenotype

Natural selection acts on an organism's phenotype, or physical characteristics. Phenotype is determined by an organism's genetic make-up (genotype) and the environment in which the organism lives. Often, natural selection acts on specific traits of an individual, and the terms phenotype and genotype are used narrowly to indicate these specific traits. When different organisms in a population possess different versions of a gene for a certain trait, each of these versions is known as an allele. It is this genetic variation that underlies phenotypic traits. A typical example is that certain combinations of genes for eye color in humans that, for instance, give rise to the phenotype of blue eyes. (On the other hand, when all the organisms in a population share the same allele for a particular trait, and this state is stable over time, the allele is said to be fixed in that population.) Some traits are governed by only a single gene, but most traits are influenced by the interactions of many genes. A variation in one of the many genes that contributes to a trait may have only a small effect on the phenotype; together, these genes can produce a continuum of possible phenotypic values.[52]

Directionality of selection
When some component of a trait is heritable, selection will alter the frequencies of the different alleles, or variants of the gene that produces the variants of the trait. Selection can be divided into three classes, on the basis of its effect on allele frequencies.[53] Directional selection occurs when a certain allele has a greater fitness than others, resulting in an increase of its frequency. This process can continue until the allele is fixed and the entire population shares the fitter phenotype. It is directional selection that is illustrated in the antibiotic resistance example above. Far more common is stabilizing selection (which is commonly confused with purifying selection[54][55]), which lowers the frequency of alleles that have a deleterious effect on the phenotype that is, produce organisms of lower

Natural selection fitness. This process can continue until the allele is eliminated from the population. Purifying selection results in functional genetic features, such as protein-coding genes or regulatory sequences, being conserved over time due to selective pressure against deleterious variants. Finally, a number of forms of balancing selection exist, which do not result in fixation, but maintain an allele at intermediate frequencies in a population. This can occur in diploid species (that is, those that have homologous pairs of chromosomes) when heterozygote individuals, who have different alleles on each chromosome at a single genetic locus, have a higher fitness than homozygote individuals that have two of the same alleles. This is called heterozygote advantage or overdominance, of which the best-known example is the malarial resistance observed in heterozygous humans who carry only one copy of the gene for sickle cell anemia. Maintenance of allelic variation can also occur through disruptive or diversifying selection, which favors genotypes that depart from the average in either direction (that is, the opposite of overdominance), and can result in a bimodal distribution of trait values. Finally, balancing selection can occur through frequency-dependent selection, where the fitness of one particular phenotype depends on the distribution of other phenotypes in the population. The principles of game theory have been applied to understand the fitness distributions in these situations, particularly in the study of kin selection and the evolution of reciprocal altruism.[][]

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Selection and genetic variation

A portion of all genetic variation is functionally neutral in that it produces no phenotypic effect or significant difference in fitness; the hypothesis that this variation accounts for a large fraction of observed genetic diversity is known as the neutral theory of molecular evolution and was originated by Motoo Kimura. When genetic variation does not result in differences in fitness, selection cannot directly affect the frequency of such variation. As a result, the genetic variation at those sites will be higher than at sites where variation does influence fitness.[53] However, after a period with no new mutation, the genetic variation at these sites will be eliminated due to genetic drift. Mutation selection balance Natural selection results in the reduction of genetic variation through the elimination of maladapted individuals and consequently of the mutations that caused the maladaptation. At the same time, new mutations occur, resulting in a mutation-selection balance. The exact outcome of the two processes depends both on the rate at which new mutations occur and on the strength of the natural selection, which is a function of how unfavorable the mutation proves to be. Consequently, changes in the mutation rate or the selection pressure will result in a different mutation-selection balance. Genetic linkage Genetic linkage occurs when the loci of two alleles are linked, or in close proximity to each other on the chromosome. During the formation of gametes, recombination of the genetic material results in reshuffling of the alleles. However, the chance that such a reshuffle occurs between two alleles depends on the distance between those alleles; the closer the alleles are to each other, the less likely it is that such a reshuffle will occur. Consequently, when selection targets one allele, this automatically results in selection of the other allele as well; through this mechanism, selection can have a strong influence on patterns of variation in the genome. Selective sweeps occur when an allele becomes more common in a population as a result of positive selection. As the prevalence of one allele increases, linked alleles can also become more common, whether they are neutral or even slightly deleterious. This is called genetic hitchhiking. A strong selective sweep results in a region of the genome where the positively selected haplotype (the allele and its neighbors) are in essence the only ones that exist in the population. Whether a selective sweep has occurred or not can be investigated by measuring linkage disequilibrium, or whether a given haplotype is overrepresented in the population. Normally, genetic recombination results in a reshuffling of the

Natural selection different alleles within a haplotype, and none of the haplotypes will dominate the population. However, during a selective sweep, selection for a specific allele will also result in selection of neighboring alleles. Therefore, the presence of a block of strong linkage disequilibrium might indicate that there has been a 'recent' selective sweep near the center of the block, and this can be used to identify sites recently under selection. Background selection is the opposite of a selective sweep. If a specific site experiences strong and persistent purifying selection, linked variation will tend to be weeded out along with it, producing a region in the genome of low overall variability. Because background selection is a result of deleterious new mutations, which can occur randomly in any haplotype, it does not produce clear blocks of linkage disequilibrium, although with low recombination it can still lead to slightly negative linkage disequilibrium overall.[]

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References
[1] Darwin C (1859) On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life John Murray, London; modern reprint Published online at The complete work of Charles Darwin online (http:/ / darwin-online. org. uk/ ): On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life (http:/ / darwin-online. org. uk/ content/ frameset?itemID=F373& viewtype=side& pageseq=2). [2] Fisher RA (1930) The Genetical Theory of Natural Selection Clarendon Press, Oxford [3] Works employing or describing this usage: [4] Works employing or describing this usage: Lande R & Arnold SJ (1983) The measurement of selection on correlated characters. Evolution 37:1210-26 Futuyma DJ (2005) Evolution. Sinauer Associates, Inc., Sunderland, Massachusetts. ISBN 0-87893-187-2 Haldane, J.B.S. 1953. The measurement of natural selection. Proceedings of the 9th International Congress of Genetics. 1: 480-487 [5] Sober E (1984; 1993) The Nature of Selection: Evolutionary Theory in Philosophical Focus University of Chicago Press ISBN 0-226-76748-5 [6] http:/ / www. sciencedaily. com/ releases/ 2011/ 01/ 110125172418. htm [7] Modified from Christiansen FB (1984) The definition and measurement of fitness. In: Evolutionary ecology (ed. Shorrocks B) pp6579. Blackwell Scientific, Oxford by adding survival selection in the reproductive phase [8] Pitnick S & Markow TA (1994) Large-male advantage associated with the costs of sperm production in Drosophila hydei, a species with giant sperm. Proc Natl Acad Sci USA 91:9277-81; Pitnick S (1996) Investment in testes and the cost of making long sperm in Drosophila. Am Nat 148:57-80 [11] (http:/ / www. blackwell-synergy. com/ doi/ abs/ 10. 1111/ j. 1469-0691. 2006. 01343. x) [12] Sylvain Charlat, Emily A. Hornett, James H. Fullard, Neil Davies, George K. Roderick, Nina Wedell & Gregory D. D. Hurst (2007). "Extraordinary flux in sex ratio". Science 317 (5835): 214. . PMID 17626876. [13] Bejerano G, Pheasant M, Makunin I, Stephen S, Kent WJ, Mattick JS & Haussler D (2004) Ultraconserved elements in the human genome. Science 304:1321-5 [15] Zakany J, FromentalRamain C, Warot X & Duboule D (1997) Regulation of number and size of digits by posterior Hox genes: a dose-dependent mechanism with potential evolutionary implications. Proc Natl Acad Sci USA 94:13695-700 [20] Mehmet Bayrakdar (Third Quarter, 1983). "Al-Jahiz And the Rise of Biological Evolutionism", The Islamic Quarterly. London. [23] Chevalier de Lamarck J-B, de Monet PA (1809) Philosophie Zoologique [26] Wallace, Alfred Russel (1870) Contributions to the Theory of Natural Selection New York: Macmillan & Co. (http:/ / www. hti. umich. edu/ cgi/ t/ text/ text-idx?c=moa& idno=AJP5195. 0001. 001& view=toc) [30] Eisley L. (1958). Darwin's Century: Evolution and the Men Who Discovered It. Doubleday & Co: New York, USA. [31] Kuhn TS. [1962] (1996). The Structure of Scientific Revolution 3rd ed. University of Chicago Press: Chicago, Illinois, USA. ISBN 0-226-45808-3 [34] Mills SK, Beatty JH. [1979] (1994). The Propensity Interpretation of Fitness. Originally in Philosophy of Science (1979) 46: 263-286; republished in Conceptual Issues in Evolutionary Biology 2nd ed. Elliott Sober, ed. MIT Press: Cambridge, Massachusetts, USA. pp3-23. ISBN 0-262-69162-0. [35] Haldane JBS (1932) The Causes of Evolution; Haldane JBS (1957) The cost of natural selection. J Genet 55:511-24( (http:/ / www. blackwellpublishing. com/ ridley/ classictexts/ haldane2. pdf). [37] Dobzhansky Th (1937) Genetics and the Origin of Species Columbia University Press, New York. (2nd ed., 1941; 3rd edn., 1951) [38] Mayr E (1942) Systematics and the Origin of Species Columbia University Press, New York. ISBN 0-674-86250-3 [39] The New York Review of Books: Darwinian Fundamentalism (http:/ / www. nybooks. com/ articles/ 1151) (accessed May 6, 2006) [40] Dennett, D. C. (1995). Darwin's dangerous idea: evolution and the meanings of life. Simon & Schuster. [41] http:/ / www. scitopics. com/ Cellular_Darwinism_stochastic_gene_expression_in_cell_differentiation_and_embryo_development. html [42] Engels F (1873-86) Dialectics of Nature 3d ed. Moscow: Progress, 1964 (http:/ / www. marxists. org/ archive/ marx/ works/ 1883/ don/ index. htm)

Natural selection
[43] Quoted in translation in Eisenberg L (2005) Which image for Lorenz? Am J Psychiatry 162:1760 (http:/ / ajp. psychiatryonline. org/ cgi/ content/ full/ 162/ 9/ 1760) [44] e.g. Wilson, DS (2002) Darwin's Cathedral: Evolution, Religion, and the Nature of Society. University of Chicago Press, ISBN 0-226-90134-3 [45] Pinker S. [1994] (1995). The Language Instinct: How the Mind Creates Language. HarperCollins: New York, NY, USA. ISBN 0-06-097651-9 [46] Dawkins R. [1976] (1989). The Selfish Gene. Oxford University Press: New York, NY, USA, p.192. ISBN 0-19-286092-5 [47] Dennett DC. (1991). Consciousness Explained. Little, Brown, and Co: New York, NY, USA. ISBN 0-316-18066-1 [48] Lotka AJ (1922a) Contribution to the energetics of evolution (http:/ / www. pubmedcentral. nih. gov/ picrender. fcgi?artid=1085052& blobtype=pdf) [PDF] Proc Natl Acad Sci USA 8:14751 Lotka AJ (1922b) Natural selection as a physical principle (http:/ / www. pubmedcentral. nih. gov/ picrender. fcgi?artid=1085053& blobtype=pdf) [PDF] Proc Natl Acad Sci USA 8:1514 [49] Kauffman SA (1993) The Origin of order. Self-organization and selection in evolution. New York: Oxford University Press ISBN 0-19-507951-5 [50] Goldberg DE. (1989). Genetic Algorithms in Search, Optimization and Machine Learning. Addison-Wesley: Boston, MA, USA [51] Mitchell, Melanie, (1996), An Introduction to Genetic Algorithms, MIT Press, Cambridge, MA. [52] Falconer DS & Mackay TFC (1996) Introduction to Quantitative Genetics Addison Wesley Longman, Harlow, Essex, UK ISBN 0-582-24302-5 [53] Rice SH. (2004). Evolutionary Theory: Mathematical and Conceptual Foundations. Sinauer Associates: Sunderland, Massachusetts, USA. ISBN 0-87893-702-1 See esp. ch. 5 and 6 for a quantitative treatment. [55] http:/ / www. nature. com/ scitable/ topicpage/ Negative-Selection-1136

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Further reading
For technical audiences Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Harvard University Press. ISBN0-674-00613-5. Maynard Smith, John (1993). The Theory of Evolution: Canto Edition. Cambridge University Press. ISBN0-521-45128-0. Popper, Karl (1978) Natural selection and the emergence of mind. Dialectica 32:339-55. See (http:// mertsahinoglu.com/research/karl-popper-on-the-scientific-status-of-darwins-theory-of-evolution/) Sober, Elliott (1984) The Nature of Selection: Evolutionary Theory in Philosophical Focus. University of Chicago Press. Williams, George C. (1966) Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. Oxford University Press. Williams George C. (1992) Natural Selection: Domains, Levels and Challenges. Oxford University Press. For general audiences Dawkins, Richard (1996) Climbing Mount Improbable. Penguin Books, ISBN 0-670-85018-7. Dennett, Daniel (1995) Darwin's Dangerous Idea: Evolution and the Meanings of Life. Simon & Schuster ISBN 0-684-82471-X. Gould, Stephen Jay (1997) Ever Since Darwin: Reflections in Natural History. Norton, ISBN 0-393-06425-5. Jones, Steve (2001) Darwin's Ghost: The Origin of Species Updated. Ballantine Books ISBN 0-345-42277-5. Also published in Britain under the title Almost like a whale: the origin of species updated. Doubleday. ISBN 1-86230-025-9. Lewontin, Richard (1978) Adaptation. Scientific American 239:212-30 Mayr, Ernst (2001) What evolution is. Weidenfeld & Nicolson, London. ISBN 0-297-60741-3 Weiner, Jonathan (1994) The Beak of the Finch: A Story of Evolution in Our Time. Vintage Books, ISBN 0-679-73337-X. Historical Zirkle, C (1941). "Natural Selection before the "Origin of Species". Proceedings of the American Philosophical Society 84 (1): 71123.

Natural selection Kohm M (2004) A Reason for Everything: Natural Selection and the English Imagination. London: Faber and Faber. ISBN 0-571-22392-3. For review, see (http://human-nature.com/nibbs/05/wyhe.html) van Wyhe J (2005) Human Nature Review 5:1-4

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External links
On the Origin of Species by Charles Darwin (http://www.literature.org/authors/darwin-charles/ the-origin-of-species/chapter-04.html) Chapter 4, Natural Selection Natural Selection (http://www.wcer.wisc.edu/ncisla/muse/naturalselection/index.html)- Modeling for Understanding in Science Education, University of Wisconsin Natural Selection (http://evolution.berkeley.edu/evolibrary/search/topicbrowse2.php?topic_id=53) from University of Berkeley education website T. Ryan Gregory: Understanding Natural Selection: Essential Concepts and Common Misconceptions (http:// www.springerlink.com/content/2331741806807x22/fulltext.html) Evolution: Education and Outreach ENCODE threads explorer (http://www.nature.com/encode/#/threads/ impact-of-evolutionary-selection-on-functional-regions) Impact of functional information on understanding variation. Nature (journal)

Genetics
Part of a series on

Genetics
Key components

Chromosome DNA RNA Genome Heredity Mutation Nucleotide Variation Glossary Index Outline History and topics

Introduction History Evolution (molecular) Population genetics Mendelian inheritance Quantitative genetics Molecular genetics Research

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DNA sequencing Genetic engineering Genomics ( template) Medical genetics Branches of genetics Biology portal

Genetics (from Ancient Greek genetikos, "genitive" and that from genesis, "origin"),[1][2][3] a discipline of biology, is the science of genes, heredity, and variation in living organisms.[][4] Genetics concerns the process of trait inheritance from parents to offspring, including the molecular structure and function of genes, gene behavior in the context of a cell or organism (e.g. dominance and epigenetics), gene distribution, and variation and change in populations (such as through Genome-Wide Association Studies). Given that genes are universal to living organisms, genetics can be applied to the study of all living systems; including bacteria, plants, animals, and humans. The observation that living things inherit traits from their parents has been used since prehistoric times to improve crop plants and animals through selective breeding[citation needed]. The modern science of genetics, seeking to understand this process, began with the work of Gregor Mendel in the mid-19th century.[] Mendel observed that organisms inherit traits by way of discrete 'units of inheritance.' This term, still used today, is a somewhat ambiguous definition of a gene. A more modern working definition of a gene is a portion (or sequence) of DNA that codes for a known cellular function. This portion of DNA is variable, it may be small or large, have a few subregions or many subregions. The word 'Gene' refers to portions of DNA that are required for a single cellular process or single function, more than the word refers to a single tangible item. A quick idiom that is often used (but not always true) is 'one gene, one protein' meaning a singular gene codes for a singular protein type in a cell. Another analogy is that a 'gene' is like a 'sentence' and 'letters' are like 'nucleotides.' A series of nucleotides can be put together without forming a gene (non-coding regions of DNA), like a string of letters can be put together without forming a sentence (babble). Nonetheless, all sentences must have letters, like all genes must have a nucleotides. The sequence of nucleotides in a gene is read and translated by a cell to produce a chain of amino acids which in turn spontaneously fold into proteins. The order of amino acids in a protein corresponds to the order of nucleotides in the gene. This relationship between nucleotide sequence and amino acid sequence is known as the genetic code. The amino acids in a protein determine how it folds into its unique three-dimensional shape; a structure that is ultimately responsible for the proteins function. Proteins carry out many of the functions needed for cells to live. A change to the DNA in a gene can change a protein's amino acid sequence, thereby changing its shape and function, rendering the protein ineffective or even malignant (see: sickle cell anemia). When a gene change occurs, it is referred to as a mutation. Although genetics plays a large role in the appearance and behavior of organisms, it is a combination of genetics with the organisms' experiences (aka. environment) that determines the ultimate outcome. Genes may be activated or inactivated, which is determined by a cell's or organism's environment, intracellularly and/or extracellularly. For example, while genes play a role in determining an organism's size, the nutrition and health it experiences after inception also have a large effect.

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History
Although the science of genetics began with the applied and theoretical work of Gregor Mendel in the mid-19th century, other theories of inheritance preceded Mendel. A popular theory during Mendel's time was the concept of blending inheritance: the idea that individuals inherit a smooth blend of traits from their parents.[citation needed] Mendel's work provided examples where traits were definitely not blended after hybridization, showing that traits are produced by combinations of distinct genes rather than a continuous blend. Blending of traits in the progeny is now explained by the action of multiple genes with quantitative effects. Another theory that had some support at that time was the inheritance of acquired characteristics: the belief that individuals inherit traits strengthened by their parents. This theory (commonly associated with Jean-Baptiste Lamarck) is now known to be wrongthe experiences of individuals do not affect the genes they pass to their children,[5] although evidence in the field of epigenetics has revived some aspects of Lamarck's theory.[6] Other theories included the pangenesis of Charles Darwin (which had both acquired and inherited aspects) and Francis Galton's reformulation of pangenesis as both particulate and inherited.[7]

Mendelian and classical genetics

Modern genetics started with Gregor Johann Mendel, a German-Czech Augustinian monk and scientist who studied the nature of inheritance in plants. In his paper "Versuche ber Pflanzenhybriden" ("Experiments on Plant Hybridization"), presented in 1865 to the Naturforschender Verein (Society for Research in Nature) in Brnn, Mendel traced the inheritance patterns of certain traits in pea plants and described them mathematically.[] Although this pattern of inheritance could only be observed for a few traits, Mendel's work suggested that heredity was particulate, not acquired, and that the inheritance patterns of many traits could be explained through simple rules and ratios.
DNA, the molecular basis for inheritance. Each strand of DNA is a chain of nucleotides, matching each other in the center to form what look like rungs on a twisted ladder.

The importance of Mendel's work did not gain wide understanding until the 1890s, after his death, when other scientists working on similar problems re-discovered his research. William Bateson, a proponent of Mendel's work, coined the word genetics in 1905.[8][9] (The adjective genetic, derived from the Greek word genesis, "origin", predates the noun and was first used in a biological sense in 1860.)[10] Bateson popularized the usage of the word genetics to describe the study of inheritance in his inaugural address to the Third International Conference on Plant Hybridization in London, England, in 1906.[11] After the rediscovery of Mendel's work, scientists tried to determine which molecules in the cell were responsible for inheritance. In 1911, Thomas Hunt Morgan argued that genes are on chromosomes, based on observations of a sex-linked white eye mutation in fruit flies.[12] In 1913, his student Alfred Sturtevant used the phenomenon of genetic linkage to show that genes are arranged linearly on the chromosome.[13]

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Molecular genetics
Although genes were known to exist on chromosomes, chromosomes are composed of both protein and DNA, and scientists did not know which of these is responsible for inheritance. In 1928, Frederick Griffith discovered the phenomenon of transformation (see Griffith's experiment): dead bacteria could transfer genetic material to "transform" other still-living bacteria. Sixteen years later, in 1944, Oswald Theodore Avery, Colin McLeod and Maclyn McCarty identified the molecule responsible for transformation as DNA.[14] The role of the nucleus as the repository of genetic information in Morgan's observation of sex-linked inheritance of eukaryotes had been established by Hmmerling in 1943 in his work a mutation causing white eyes in Drosophila led [15] on the single celled alga Acetabularia. The Hershey-Chase him to the hypothesis that genes are located upon experiment in 1952 confirmed that DNA (rather than protein) is the chromosomes. genetic material of the viruses that infect bacteria, providing further evidence that DNA is the molecule responsible for inheritance.[16] James D. Watson and Francis Crick determined the structure of DNA in 1953, using the X-ray crystallography work of Rosalind Franklin and Maurice Wilkins that indicated DNA had a helical structure (i.e., shaped like a corkscrew).[17][] Their double-helix model had two strands of DNA with the nucleotides pointing inward, each matching a complementary nucleotide on the other strand to form what looks like rungs on a twisted ladder.[] This structure showed that genetic information exists in the sequence of nucleotides on each strand of DNA. The structure also suggested a simple method for duplication: if the strands are separated, new partner strands can be reconstructed for each based on the sequence of the old strand. Although the structure of DNA showed how inheritance works, it was still not known how DNA influences the behavior of cells. In the following years,[citation needed] scientists tried to understand how DNA controls the process of protein production. It was discovered that the cell uses DNA as a template to create matching messenger RNA, molecules with nucleotides very similar to DNA. The nucleotide sequence of a messenger RNA is used to create an amino acid sequence in protein; this translation between nucleotide sequenced and amino acid sequences is known as the genetic code[citation needed]. With the newfound molecular understanding of inheritance came an explosion of research [citation needed]. One important development was chain-termination DNA sequencing in 1977 by Frederick Sanger. This technology allows scientists to read the nucleotide sequence of a DNA molecule.[] In 1983, Kary Banks Mullis developed the polymerase chain reaction, providing a quick way to isolate and amplify a specific section of a DNA from a mixture.[] Through the pooled efforts of the Human Genome Project Department of Energy and the NIH and the parallel private effort by Celera Genomics, these and other methods culminated in the sequencing of the human genome in 2003.[]

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Features of inheritance
Discrete inheritance and Mendel's laws
At its most fundamental level, inheritance in organisms occurs by passing discrete heritable units, called genes, from parents to progeny.[] This property was first observed by Gregor Mendel, who studied the segregation of heritable traits in pea plants.[][] In his experiments studying the trait for flower color, Mendel observed that the flowers of each pea plant were either purple or whitebut never an intermediate between the two colors. These different, discrete versions of the same gene are called alleles. In the case of pea, which is a diploid species, each individual plant has two copies of each gene, one copy inherited from each parent.[] Many species, including humans, have this pattern of inheritance. Diploid organisms with two copies of the same allele of a given gene are called homozygous at that gene locus, while organisms with two different alleles of a given gene are called heterozygous.

A Punnett square depicting a cross between two pea plants heterozygous for purple (B) and white (b) blossoms

The set of alleles for a given organism is called its genotype, while the observable traits of the organism are called its phenotype. When organisms are heterozygous at a gene, often one allele is called dominant as its qualities dominate the phenotype of the organism, while the other allele is called recessive as its qualities recede and are not observed. Some alleles do not have complete dominance and instead have incomplete dominance by expressing an intermediate phenotype, or codominance by expressing both alleles at once.[] When a pair of organisms reproduce sexually, their offspring randomly inherit one of the two alleles from each parent. These observations of discrete inheritance and the segregation of alleles are collectively known as Mendel's first law or the Law of Segregation.

Notation and diagrams

Geneticists use diagrams and symbols to describe inheritance. A gene is represented by one or a few letters. Often a "+" symbol is used to mark the usual, non-mutant allele for a gene.[18] In fertilization and breeding experiments (and especially when discussing Mendel's laws) the parents are referred to as the "P" generation and the offspring as the "F1" (first filial) generation. When the F1 offspring mate with each other, the offspring are called the "F2" (second filial) generation. One of the common diagrams used to predict the result of cross-breeding is the Punnett square.
Genetic pedigree charts help track the inheritance patterns of traits.

When studying human genetic diseases, geneticists often use pedigree charts to represent the inheritance of traits.[] These charts map the inheritance of a trait in a family

tree.

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Interactions of multiple genes

Organisms have thousands of genes, and in sexually reproducing organisms these genes generally assort independently of each other. This means that the inheritance of an allele for yellow or green pea color is unrelated to the inheritance of alleles for white or purple flowers. This phenomenon, known as "Mendel's second law" or the "Law of independent assortment", means that the alleles of different genes get shuffled between parents to form offspring with many different combinations. (Some genes do not assort independently, demonstrating genetic linkage, a topic discussed later in this article.) Often different genes can interact in a way that influences the same trait. In the Blue-eyed Mary (Omphalodes verna), for example, there exists a gene with alleles that determine the color of flowers: blue or magenta. Another gene, however, controls whether the flowers have color at all or are white. When a plant has two copies of this white allele, its flowers are whiteregardless of whether the first gene has blue or magenta alleles. This interaction between genes is called epistasis, with the second gene epistatic to the first.[]

Human height is a trait with complex genetic causes. Francis Galton's data from 1889 shows the relationship between offspring height as a function of mean parent height. While correlated, remaining variation in offspring heights indicates environment is also an important factor in this trait.

Many traits are not discrete features (e.g. purple or white flowers) but are instead continuous features (e.g. human height and skin color). These complex traits are products of many genes.[19] The influence of these genes is mediated, to varying degrees, by the environment an organism has experienced. The degree to which an organism's genes contribute to a complex trait is called heritability.[] Measurement of the heritability of a trait is relativein a more variable environment, the environment has a bigger influence on the total variation of the trait. For example, human height is a trait with complex causes. It has a heritability of 89% in the United States. In Nigeria, however, where people experience a more variable access to good nutrition and health care, height has a heritability of only 62%.[20]

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Molecular basis for inheritance

DNA and Chromosomes
The molecular basis for genes is deoxyribonucleic acid (DNA). DNA is composed of a chain of nucleotides, of which there are four types: adenine (A), cytosine (C), guanine (G), and thymine (T). Genetic information exists in the sequence of these nucleotides, and genes exist as stretches of sequence along the DNA chain.[] Viruses are the only exception to this rulesometimes viruses use the very similar molecule RNA instead of DNA as their genetic material.[21] It should be noted that viruses cannot reproduce without a host and generally aren't considered 'living' organisms, thus are not subject to many principles of genetics. DNA normally exists as a double-stranded molecule, coiled into the shape of a double-helix. Each nucleotide in DNA preferentially pairs with its partner nucleotide on the opposite strand: A pairs with T, and The molecular structure of DNA. Bases pair C pairs with G. Thus, in its two-stranded form, each strand effectively through the arrangement of hydrogen bonding contains all necessary information, redundant with its partner strand. between the strands. This structure of DNA is the physical basis for inheritance: DNA replication duplicates the genetic information by splitting the strands and using each strand as a template for synthesis of a new partner strand.[] Genes are arranged linearly along long chains of DNA base-pair sequences. In bacteria, each cell usually contains a single circular genophore, while eukaryotic organisms (including plants and animals) have their DNA arranged in multiple linear chromosomes. These DNA strands are often extremely long; the largest human chromosome, for example, is about 247 million base pairs in length.[22] The DNA of a chromosome is associated with structural proteins that organize, compact, and control access to the DNA, forming a material called chromatin; in eukaryotes, chromatin is usually composed of nucleosomes, segments of DNA wound around cores of histone proteins.[23] The full set of hereditary material in an organism (usually the combined DNA sequences of all chromosomes) is called the genome. While haploid organisms have only one copy of each chromosome, most animals and many plants are diploid, containing two of each chromosome and thus two copies of every gene.[] The two alleles for a gene are located on identical loci of the two homologous chromosomes, each allele inherited from a different parent.

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95 Many species have so called sex chromosomes. They are special in that they determine the sex of the organism.[] In humans and many other animals, the Y-chromosome contains the gene that triggers the development of the specifically male characteristics. In evolution, this chromosome has lost most of its content and also most of its genes, while the X chromosome is similar to the other chromosomes and contains many genes. The X and Y chromosomes form a very heterogeneous pair.

Reproduction
When cells divide, their full genome is copied and each daughter cell inherits one copy. This process, called mitosis, is the simplest form of reproduction and is the basis for asexual reproduction. Asexual reproduction can also occur in multicellular organisms, producing offspring that inherit their genome from a single parent. Offspring that are genetically identical to their parents are called clones.

Eukaryotic organisms often use sexual reproduction to generate offspring that contain a mixture of genetic material inherited from two different parents. The process of sexual reproduction alternates between forms that contain single copies of the genome (haploid) and [] double copies (diploid). Haploid cells fuse and combine genetic material to create a diploid cell with paired chromosomes. Diploid organisms form haploids by dividing, without replicating their DNA, to create daughter cells that randomly inherit one of each pair of chromosomes. Most animals and many plants are diploid for most of their lifespan, with the haploid form reduced to single cell gametes such as sperm or eggs. Although they do not use the haploid/diploid method of sexual reproduction, bacteria have many methods of acquiring new genetic information. Some bacteria can undergo conjugation, transferring a small circular piece of DNA to another bacterium.[] Bacteria can also take up raw DNA fragments found in the environment and integrate them into their genomes, a phenomenon known as transformation.[] These processes result in horizontal gene transfer, transmitting fragments of genetic information between organisms that would be otherwise unrelated.

Walther Flemming's 1882 diagram of eukaryotic cell division. Chromosomes are copied, condensed, and organized. Then, as the cell divides, chromosome copies separate into the daughter cells.

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Recombination and Genetic Linkage

The diploid nature of chromosomes allows for genes on different chromosomes to assort independently or be separated from their homologous pair during sexual reproduction wherein haploid gametes are formed. In this way new combinations of genes can occur in the offspring of a mating pair. Genes on the same chromosome would theoretically never recombine. However, they do via the cellular process of chromosomal crossover. During crossover, chromosomes exchange stretches of DNA, effectively shuffling the gene alleles between the chromosomes.[] This process of chromosomal crossover generally occurs during meiosis, a series of cell divisions that creates haploid cells. The probability of chromosomal crossover occurring between two given points on the chromosome is related to the distance between the points. For an arbitrarily long distance, the probability of crossover is high enough that the inheritance of the genes is effectively uncorrelated[citation needed]. For genes that are closer together, however, Thomas Hunt Morgan's 1916 illustration of a the lower probability of crossover means that the genes demonstrate double crossover between chromosomes genetic linkagealleles for the two genes tend to be inherited together. The amounts of linkage between a series of genes can be combined to form a linear linkage map that roughly describes the arrangement of the genes along the chromosome.[]

Gene expression
Genetic Code
Genes generally express their functional effect through the production of proteins, which are complex molecules responsible for most functions in the cell. Proteins are made up of one or more polypeptide chains, each of which is composed of a sequence of amino acids, and the DNA sequence of a gene (through an RNA intermediate) is used to produce a specific amino acid sequence. This process begins with the production of an RNA molecule with a sequence matching the gene's DNA sequence, a process called transcription.

The genetic code: DNA, through a messenger RNA intermediate, codes for protein with a triplet code.

This messenger RNA molecule is then used to produce a corresponding amino acid sequence through a process called translation. Each group of three nucleotides in the sequence, called a codon, corresponds either to one of the twenty possible amino acids in a protein or an instruction to end the amino acid sequence; this correspondence is called the genetic code.[24] The flow of information is unidirectional: information is transferred from nucleotide sequences into the amino acid sequence of proteins, but it never transfers from protein back into the sequence of DNAa phenomenon Francis Crick called the central dogma of molecular biology.[]

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The specific sequence of amino acids results in a unique three-dimensional structure for that protein, and the three-dimensional structures of proteins are related to their functions.[25][26] Some are simple structural molecules, like the fibers formed by the protein collagen. Proteins can bind to other proteins and simple molecules, sometimes acting as enzymes by facilitating chemical reactions within the bound molecules (without changing the structure of the protein itself). Protein structure is dynamic; the protein hemoglobin bends into slightly different forms as it facilitates the capture, transport, and release of oxygen molecules within mammalian blood. A single nucleotide difference within DNA can cause a change in the amino acid sequence of a protein. Because protein structures are the result of their amino acid sequences, some changes can dramatically change the properties of a protein by destabilizing the structure or changing the surface of the protein in a way that changes its interaction with other proteins and molecules. For example, A single amino acid change causes sickle-cell anemia is a human genetic disease that results from a single base hemoglobin to form fibers. difference within the coding region for the -globin section of hemoglobin, causing a single amino acid change that changes hemoglobin's physical properties.[27] Sickle-cell versions of hemoglobin stick to themselves, stacking to form fibers that distort the shape of red blood cells carrying the protein. These sickle-shaped cells no longer flow smoothly through blood vessels, having a tendency to clog or degrade, causing the medical problems associated with this disease. Some genes are transcribed into RNA but are not translated into protein productssuch RNA molecules are called non-coding RNA. In some cases, these products fold into structures which are involved in critical cell functions (e.g. ribosomal RNA and transfer RNA). RNA can also have regulatory effect through hybridization interactions with other RNA molecules (e.g. microRNA)[citation needed].

Nature versus Nurture

Although genes contain all the information an organism uses to function, the environment plays an important role in determining the ultimate phenotypes an organism displaysa phenomenon often referred to as "nature vs. nurture". The phenotype of an organism depends on the interaction of genes and the environment. An interesting example is the coat coloration of the siamese cat. In this case the body temperature of the cat plays the role of the environment. The cat's genes code for dark-hair, thus the hair producing cells in the cat make cellular proteins resulting in dark hair. But these dark-hair-producing proteins are sensitive to temperature (have a temperature sensitive mutation) and denature in higher temperature environments failing to function in producing dark-hair pigment in areas where the cat has a higher body temperature. In a low temperature environment, however, the protein's structure is stable and produces dark-hair pigment normally. The protein remains functional in areas of skin that are colder (e.g., legs, ears, tail, and face)- so the cat has dark-hair at its extremities.[28]

Siamese cats have a temperature-sensitive mutation in pigment production.

Environment plays a major role in effects of the human genetic disease phenylketonuria.[29] The mutation that causes phenylketonuria disrupts the ability of the body to break down the amino acid phenylalanine, causing a toxic build-up of an intermediate molecule that, in turn, causes severe symptoms of progressive mental retardation and seizures. However, if someone with the phenylketonuria mutation follows a strict diet that avoids this amino acid,

Genetics they remain normal and healthy. A popular method in determining how genes and environment (i.e. 'nature and nurture') contribute to a phenotype is by studying identical and fraternal twins or siblings of multiple births.[30] Because identical siblings come from the same zygote they are genetically the same. Fraternal siblings are as genetically different from one another as normal siblings. By analyzing statistics on how often a twin of a set has a certain disorder compared other sets of twins, scientists can determine whether that disorder is caused by genetic or environmental factors (i.e. whether it has 'nature' or 'nurture' causes). One famous example is the multiple birth study of the Genain quadruplets, whom were identical quadruplets all diagnosed with schizophrenia.[]

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Gene regulation
The genome of a given organism contains thousands of genes, but not all these genes need to be active at any given moment. A gene is expressed when it is being transcribed into mRNA and there exist many cellular methods of controlling the expression of genes such that proteins are produced only when needed by the cell. Transcription factors are regulatory proteins that bind to DNA, either promoting or inhibiting the transcription of a gene.[31] Within the genome of Escherichia coli bacteria, for example, there exists a series of genes necessary for the synthesis of the amino acid tryptophan. However, when tryptophan is already available to the cell, these genes for tryptophan synthesis are no longer needed. The presence of tryptophan directly affects the activity of the genestryptophan molecules bind to the tryptophan repressor (a transcription factor), changing the repressor's structure such that the repressor binds to the genes. The tryptophan repressor blocks the transcription and expression of the genes, thereby creating negative feedback regulation of the tryptophan synthesis process.[32] Differences in gene expression are especially clear within multicellular organisms, where cells all contain the same genome but have very different structures and behaviors due to the expression of different sets of genes. All the cells in a multicellular organism derive from a single cell, differentiating into variant cell types in response to external and intercellular signals and gradually establishing different patterns of gene expression to create different behaviors. As no single gene is responsible for the development of structures within multicellular organisms, these patterns arise from the complex interactions between many cells.

Within eukaryotes there exist structural features of chromatin that influence the transcription of genes, often in the form of modifications to DNA and chromatin that are stably inherited by daughter cells.[33] These features are called "epigenetic" because they exist "on top" of the DNA sequence and retain inheritance from one cell generation to the next. Because of epigenetic features, different cell types grown within the same medium can retain very different properties. Although epigenetic features are generally dynamic over the course of development, some, like the phenomenon of paramutation, have multigenerational inheritance and exist as rare exceptions to the general rule of DNA as the basis for inheritance.[34]

Transcription factors bind to DNA, influencing the transcription of associated genes.

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Genetic change
Mutations
During the process of DNA replication, errors occasionally occur in the polymerization of the second strand. These errors, called mutations, can have an impact on the phenotype of an organism, especially if they occur within the protein coding sequence of a gene. Error rates are usually very low1 error in every 10100million basesdue to the "proofreading" ability of DNA polymerases.[][] Processes that increase the rate of changes in DNA are called mutagenic: mutagenic chemicals promote errors in DNA replication, often by interfering with the structure of base-pairing, while UV radiation induces mutations by causing damage to the DNA structure.[] Chemical damage to DNA occurs naturally as well, and cells use DNA repair mechanisms to repair mismatches and breaks in DNAnevertheless, the repair sometimes fails to return the DNA to its original sequence. In organisms that use chromosomal crossover to exchange DNA and recombine Gene duplication allows genes, errors in alignment during meiosis can also cause mutations.[] Errors in diversification by providing redundancy: one gene can mutate crossover are especially likely when similar sequences cause partner and lose its original function without chromosomes to adopt a mistaken alignment; this makes some regions in harming the organism. genomes more prone to mutating in this way. These errors create large structural changes in DNA sequenceduplications, inversions, or deletions of entire regionsor the accidental exchanging of whole parts between different chromosomes (called translocation).

Natural selection and evolution

Mutations alter an organisms genotype and occasionally this causes different phenotypes to appear. Most mutations have little effect on an organism's phenotype, health, or reproductive fitness. Mutations that do have an effect are usually deleterious, but occasionally some can be beneficial. Studies in the fly Drosophila melanogaster suggest that if a mutation changes a protein produced by a gene, about 70 percent of these mutations will be harmful with the remainder being either neutral or weakly beneficial.[35] Population genetics studies the distribution of genetic differences within populations and how these distributions change over time.[] Changes in the frequency of an allele in a population are mainly influenced by natural selection, where a given allele provides a selective or reproductive advantage to the organism,[] as well as other factors such as mutation, genetic drift, genetic draft,[36] artificial selection, and migration.[] Over many generations, the genomes of organisms can change significantly, resulting in the phenomenon of evolution. Selection for beneficial mutations can cause a species to evolve into forms better An evolutionary tree of eukaryotic organisms, able to survive in their environment, a process called adaptation.[37] constructed by comparison of several orthologous New species are formed through the process of speciation, often caused gene sequences by geographical separations that prevent populations from exchanging genes with each other.[] The application of genetic principles to the study of population biology and evolution is referred to as the modern synthesis.

Genetics By comparing the homology between different species' genomes, it is possible to calculate the evolutionary distance between them and when they may have diverged (called a molecular clock). Genetic comparisons are generally considered a more accurate method of characterizing the relatedness between species than the comparison of phenotypic characteristics. The evolutionary distances between species can be used to form evolutionary trees; these trees represent the common descent and divergence of species over time, although they do not show the transfer of genetic material between unrelated species (known as horizontal gene transfer and most common in bacteria).[38]

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Research and technology

Model organisms
Although geneticists originally studied inheritance in a wide range of organisms, researchers began to specialize in studying the genetics of a particular subset of organisms. The fact that significant research already existed for a given organism would encourage new researchers to choose it for further study, and so eventually a few model organisms became the basis for most genetics research.[39] Common research topics in model organism genetics include the study of gene regulation and the involvement of genes in development and cancer. Organisms were chosen, in part, for convenienceshort generation The common fruit fly (Drosophila melanogaster) times and easy genetic manipulation made some organisms popular is a popular model organism in genetics research. genetics research tools. Widely used model organisms include the gut bacterium Escherichia coli, the plant Arabidopsis thaliana, baker's yeast (Saccharomyces cerevisiae), the nematode Caenorhabditis elegans, the common fruit fly (Drosophila melanogaster), and the common house mouse (Mus musculus).

Medicine
Medical genetics seeks to understand how genetic variation relates to human health and disease.[40] When searching for an unknown gene that may be involved in a disease, researchers commonly use genetic linkage and genetic pedigree charts to find the location on the genome associated with the disease. At the population level, researchers take advantage of Mendelian randomization to look for locations in the genome that are associated with diseases, a method especially useful for multigenic traits not clearly defined by a single gene.[41] Once a candidate gene is found, further research is often done on the corresponding gene (called an orthologous gene) in model organisms. In addition to studying genetic diseases, the increased availability of genotyping methods has led to the field of pharmacogeneticsstudying how genotype can affect drug responses.[42] Individuals differ in their inherited tendency to develop cancer,[43] and cancer is a genetic disease.[44] The process of cancer development in the body is a combination of events. Mutations occasionally occur within cells in the body as they divide. Although these mutations will not be inherited by any offspring, they can affect the behavior of cells, sometimes causing them to grow and divide more frequently. There are biological mechanisms that attempt to stop this process; signals are given to inappropriately dividing cells that should trigger cell death, but sometimes additional mutations occur that cause cells to ignore these messages. An internal process of natural selection occurs within the body and eventually mutations accumulate within cells to promote their own growth, creating a cancerous tumor that grows and invades various tissues of the body. Normally, a cell divides only in response to signals: "growth factors", it stops growing when making contact with surrounding cells (contact inhibition), and in response to growth inhibitory signals, it divides a limited number of times and dies (apoptosis), it stays inside the epithelium and is not able to migrate to invade other organs. To become

Genetics a cancer cell, a cell has to accumulate mutations in a number of genes (3-7) that allow it to bypass all these regulations: it no longer needs growth factors to divide, it continues growing when making contact to neighbor cells, and ignores inhibitory signals, it will keep growing indefinitely and is immortal, it will escape from the epithelium and ultimately may be able to escape from the primary tumor, cross the endothelium of a blood vessel, be transported by the bloodstream and will colonize a new organ, forming deadly metastasis. Although there are some genetic predispositions in a small fraction of cancers, the major fraction is due to a set of new genetic mutations that originally appear and accumulate in one or a small number of cells that will divide to form the tumor and are not transmitted to the progeny (somatic mutations). The most frequent mutations are a loss of function of p53 protein, a tumor suppressor, or in the p53 pathway, and gain of function mutations in the ras proteins, or in other oncogenes.

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Research methods
DNA can be manipulated in the laboratory. Restriction enzymes are commonly used enzymes that cut DNA at specific sequences, producing predictable fragments of DNA.[45] DNA fragments can be visualized through use of gel electrophoresis, which separates fragments according to their length. The use of ligation enzymes allows DNA fragments to be connected, and by ligating fragments of DNA together from different sources, researchers can create recombinant DNA. Often associated with genetically modified organisms, recombinant DNA is commonly used in the context of plasmidsshort circular DNA fragments with a few genes on them. By inserting plasmids into bacteria and growing those bacteria on plates of agar (to isolate clones of bacteria cells), researchers can clonally amplify the inserted fragment of DNA (a process known as molecular cloning). (Cloning can also refer to creating clonal organisms, by various means.) DNA can also be amplified using a procedure called the polymerase chain reaction (PCR).[46] By using specific short sequences of DNA, PCR can isolate and exponentially amplify a targeted region of DNA. Because it can amplify from extremely small amounts of DNA, PCR is also often used to detect the presence of specific DNA sequences.

DNA sequencing and genomics

One of the most fundamental technologies developed to study genetics, DNA sequencing allows researchers to determine the sequence of nucleotides in DNA fragments. Developed in 1977 by Frederick Sanger and coworkers, chain-termination sequencing is now routinely used to sequence DNA fragments.[47] With this technology researchers have been able to study the molecular sequences associated with many human diseases.

Colonies of E. coli on a plate of agar, an example of cellular cloning and often used in molecular cloning.

As sequencing has become less expensive, researchers have sequenced the genomes of many organisms, using computational tools to stitch together the sequences of many different fragments (a process called genome assembly).[48] These technologies were used to sequence the human genome, leading to the completion of the Human Genome Project in 2003.[] New high-throughput sequencing technologies are dramatically lowering the cost of DNA sequencing, with many researchers hoping to bring the cost of resequencing a human genome down to a thousand dollars.[49] The large amount of sequence data available has created the field of genomics, research that uses computational tools to search for and analyze patterns in the full genomes of organisms. Genomics can also be considered a subfield of bioinformatics, which uses computational approaches to analyze large sets of biological data.

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References
[4] Hartl D, Jones E (2005) [5] Lamarck, J-B (2008). In Encyclopdia Britannica. Retrieved from Encyclopdia Britannica Online (http:/ / www. search. eb. com/ eb/ article-273180) on 16 March 2008. [7] Peter J. Bowler, The Mendelian Revolution: The Emergency of Hereditarian Concepts in Modern Science and Society (Baltimore: Johns Hopkins University Press, 1989): chapters 2 & 3. [8] genetics, n., Oxford English Dictionary, 3rd ed. [9] Note that the letter was to an Adam Sedgwick, a zoologist and "Reader in Animal Morphology" at Trinity College, Cambridge [10] genetic, adj., Oxford English Dictionary, 3rd ed.

Initially titled the "International Conference on Hybridisation and Plant Breeding", the title was changed as a result of Bateson's speech. See:
[14] Reprint: [15] Cell and Molecular Biology", Pragya Khanna. I. K. International Pvt Ltd, 2008. p. 221. ISBN 81-89866-59-1, ISBN 978-81-89866-59-4 [23] Alberts et al. (2002), II.4. DNA and chromosomes: Chromosomal DNA and Its Packaging in the Chromatin Fiber (http:/ / www. ncbi. nlm. nih. gov/ books/ bv. fcgi?rid=mboc4. section. 608) [25] Alberts et al. (2002), I.3. Proteins: The Shape and Structure of Proteins (http:/ / www. ncbi. nlm. nih. gov/ books/ bv. fcgi?rid=mboc4. section. 388) [26] Alberts et al. (2002), I.3. Proteins: Protein Function (http:/ / www. ncbi. nlm. nih. gov/ books/ bv. fcgi?rid=mboc4. section. 452) [30] e.g. [32] Alberts et al. (2002), II.3. Control of Gene Expression The Tryptophan Repressor Is a Simple Switch That Turns Genes On and Off in Bacteria (http:/ / www. ncbi. nlm. nih. gov/ books/ bv. fcgi?rid=mboc4. section. 1269#1270) [37] Related earlier ideas were acknowledged in [44] Chapter 18: Cancer Genetics (http:/ / www. ncbi. nlm. nih. gov/ books/ bv. fcgi?rid=hmg. chapter. 2342) [45] Lodish et al. (2000), Chapter 7: 7.1. DNA Cloning with Plasmid Vectors (http:/ / www. ncbi. nlm. nih. gov/ books/ bv. fcgi?rid=mcb. section. 1582) [46] Lodish et al. (2000), Chapter 7: 7.7. Polymerase Chain Reaction: An Alternative to Cloning (http:/ / www. ncbi. nlm. nih. gov/ books/ bv. fcgi?highlight=PCR& rid=mcb. section. 1718) [48] Brown (2002), Section 2, Chapter 6: 6.2. Assembly of a Contiguous DNA Sequence (http:/ / www. ncbi. nlm. nih. gov/ books/ bv. fcgi?rid=genomes. section. 6481)

Further reading
Alberts B, Johnson A, Lewis J, Raff M, Roberts K, and Walter P (2002). Molecular Biology of the Cell (4th ed.). New York: Garland Science. ISBN0-8153-3218-1. Griffiths, William M.; Miller, Jeffrey H.; Suzuki, David T.; Lewontin, Richard C.; Gelbart, eds. (2000). An Introduction to Genetic Analysis (7th ed.). New York: W. H. Freeman. ISBN0-7167-3520-2. Hartl D, Jones E (2005). Genetics: Analysis of Genes and Genomes (6th ed.). Jones & Bartlett. ISBN0-7637-1511-5. Lodish H, Berk A, Zipursky LS, Matsudaira P, Baltimore D, and Darnell J (2000). Molecular Cell Biology (4th ed.). New York: Scientific American Books. ISBN0-7167-3136-3.

External links

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Library resources
About Genetics

Online books (http://tools.wmflabs.org/ftl/cgi-bin/ftl?st=&su=Genetics&library=OLBP) Resources in your library (http://tools.wmflabs.org/ftl/cgi-bin/ftl?st=&su=Genetics) Resources in other libraries (http://tools.wmflabs.org/ftl/cgi-bin/ftl?st=&su=Genetics&library=0CHOOSE0)

Genetics (http://www.bbc.co.uk/programmes/p00547md) on In Our Time at the BBC. ( listen now (http:// www.bbc.co.uk/iplayer/console/p00547md/In_Our_Time_Genetics)) Genetics (http://www.dmoz.org/Science/Biology/Genetics/) at the Open Directory Project

Gregor Mendel

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Gregor Mendel
Gregor Mendel

Born

Johann Mendel July 20, 1822 Heinzendorf bei Odrau, Austrian Empire (now Hynice, Czech Republic) January 6, 1884 (aged61) Brno (Brnn), Austria-Hungary (now Czech Republic) Empire of Austria-Hungary Genetics St Thomas's Abbey

Died

Nationality Fields Institutions

Alma mater University of Olomouc University of Vienna Knownfor Creating the science of genetics

Gregor Johann Mendel (July 20, 1822[1] January 6, 1884) was a German-speaking Silesian[2][3] scientist and Augustinian friar who gained posthumous fame as the founder of the new science of genetics. Mendel demonstrated that the inheritance of certain traits in pea plants follows particular patterns, now referred to as the laws of Mendelian inheritance. The profound significance of Mendel's work was not recognized until the turn of the 20th century, when the independent rediscovery of these laws initiated the modern science of genetics.[]

Biography
Gregor Mendel was born into an ethnic German family in Heinzendorf bei Odrau, Austrian Silesia, Austrian Empire (now Hynice, Czech Republic). He was the son of Anton and Rosine (Schwirtlich) Mendel, and had one older sister, Veronika, and one younger, Theresia. He was christened Johann Mendel and given the name Gregor when he joined the Augustinian monks.[4] They lived and worked on a farm which had been owned by the Mendel family for at least 130 years.[5] During his childhood, Mendel worked as a gardener, studied beekeeping, and as a young man attended gymnasium in Opava. He took four months off during his Gymnasium studies due to illness. From 1840 to 1843, he studied practical and theoretical philosophy as well as physics at the University of Olomouc Faculty of Philosophy, taking another year off because of illness. He also struggled financially to pay for his studies and Theresia gave him her dowry. Later he helped support her three sons, two of whom became doctors. He became a monk because it enabled him to obtain an education without having to pay for it himself.[6]

Gregor Mendel When Mendel entered the Faculty of Philosophy, the Department of Natural History and Agriculture was headed by Johann Karl Nestler, who conducted extensive research of hereditary traits of plants and animals, especially sheep. Upon recommendation of his physics teacher Friedrich Franz,[] Mendel entered the Augustinian St Thomas's Abbey and began his training as a priest. Born Johann Mendel, he took the name Gregor upon entering religious life. Mendel worked as a substitute high school teacher. In 1850 he failed the oral part, the last of three parts, of his exams to become a certified high school teacher. In 1851 he was sent to the University of Vienna to study under the sponsorship of Abbot C. F. Napp so that he could get more formal education.[7] At Vienna, his professor of physics was Christian Doppler.[] Mendel returned to his abbey in 1853 as a teacher, principally of physics. In 1856 he took the exam to become a certified teacher and again failed the oral part.[7]In 1867 he replaced Napp as abbot of the monastery.[] Mendel began his studies on heredity at St. Thomas's Abbey with mice, but his bishop did not like one of his monks studying animal sex, so Mendel switched to plants.[8] Mendel also bred bees in a bee house that was built for him, using bee hives that he designed.[9] He also studied astronomy and meteorology,[] founding the 'Austrian Meteorological Society' in 1865.[] The majority of his published works were related to meteorology.[]

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Experiments on plant hybridization

Gregor Mendel, who is known as the "father of modern genetics", was inspired by both his professors at the University of Olomouc (i.e. Friedrich Franz & Johann Karl Nestler) and his colleagues at the monastery (e.g., Franz Diebl) to study variation in plants, and he conducted his study in the monastery's 2 hectares (4.9acres) experimental garden,[10] which was originally planted by Napp in 1830.[] Unlike Nestler, who studied hereditary traits in sheep, Mendel focused on plants. After initial experiments with pea plants, Mendel settled on studying seven traits that seemed to inherit independently of other traits: seed shape, flower color, seed coat tint, pod shape, unripe pod color, flower location, and plant height. He first focused on seed shape, which was either angular or round.[11] Between 1856 and 1863 Mendel cultivated and tested some 29,000 pea plants (i.e., Pisum sativum). This study showed that one in four pea plants had purebred recessive alleles, two out of four were hybrid and one out of four were purebred dominant. His experiments led him to make two generalizations, the Law of Segregation and the Law of Independent Assortment, which later came to be known as Mendel's Laws of Inheritance. Mendel presented his paper, Versuche ber Pflanzenhybriden (Experiments on Plant Hybridization), at two meetings of the Natural History Society of Brno in Moravia on February 8 and March 8, 1865.[12] It was received favorably and generated reports in several local newspapers.[13] When Mendel's paper was published in 1866 in Verhandlungen des naturforschenden Vereins Brnn,[14] it was seen as essentially about hybridization rather than inheritance and had little impact and was cited about three times over the next thirty-five years. Notably, Charles Darwin was unaware of Mendel's paper, according to Jacob Bronowski's The Ascent of Man. His paper was criticized at the time, but is now considered a seminal work.

Life after the pea experiments

After completing his work with peas, Mendel turned to experimenting with honeybees to extend his work to animals. He produced a hybrid strain (so vicious they were destroyed) but failed to generate a clear picture of their heredity because of the difficulties in controlling mating behaviours of queen bees.Wikipedia:Disputed statement He also described novel plant species, and these are denoted with the botanical author abbreviation "Mendel". After he was elevated as abbot in 1868, his scientific work largely ended, as Mendel became consumed with his increased administrative responsibilities, especially a dispute with the civil government over their attempt to impose special taxes on religious institutions.[15] Mendel died on January 6, 1884, at the age of 61, in Brno, Moravia, Austria-Hungary (now Czech Republic), from chronic nephritis. Czech composer Leo Janek played the organ at his funeral. After his death, the succeeding abbot burned all papers in Mendel's collection, to mark an end to the disputes over taxation.[]

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Rediscovery of Mendel's work

Mendel's work was rejected at first, and was not widely accepted until after he died. During his own lifetime, most biologists held the idea that all characteristics were passed to the next generation through blending inheritance, in which the traits from each parent are averaged together. Instances of this phenomenon are now explained by the action of multiple genes with quantitative effects. Charles Darwin tried unsuccessfully to explain inheritance through a theory of pangenesis. It was not until the early 20th century that the importance of Mendel's ideas was realized. By 1900, research aimed at finding a successful theory of discontinuous inheritance rather than blending inheritance led to independent duplication of his work by Hugo de Vries and Carl Correns, and the rediscovery of Mendel's writings and laws. Both acknowledged Mendel's priority, and it is thought probable that de Dominant and recessive phenotypes. (1) Parental generation. (2) F1 generation. (3) F2 generation. Vries did not understand the results he had found until after reading [] Mendel. Though Erich von Tschermak was originally also credited with rediscovery, this is no longer accepted because he did not understand Mendel's laws.[16] Though de Vries later lost interest in Mendelism, other biologists started to establish genetics as a science.[] All three of these researchers, each from a different country, published their work rediscovering Mendel's work within a two-month span in the Spring of 1900.[17] Mendel's results were quickly replicated, and genetic linkage quickly worked out. Biologists flocked to the theory; even though it was not yet applicable to many phenomena, it sought to give a genotypic understanding of heredity which they felt was lacking in previous studies of heredity which focused on phenotypic approaches. Most prominent of these latter approaches was the biometric school of Karl Pearson and W.F.R. Weldon, which was based heavily on statistical studies of phenotype variation. The strongest opposition to this school came from William Bateson, who perhaps did the most in the early days of publicising the benefits of Mendel's theory (the word "genetics", and much of the discipline's other terminology, originated with Bateson). This debate between the biometricians and the Mendelians was extremely vigorous in the first two decades of the twentieth century, with the biometricians claiming statistical and mathematical rigor, whereas the Mendelians claimed a better understanding of biology. In the end, the two approaches were combined, especially by work conducted by R. A. Fisher as early as 1918. The combination, in the 1930s and 1940s, of Mendelian genetics with Darwin's theory of natural selection resulted in the modern synthesis of evolutionary biology.

Controversy
Mendel's experimental results have later been the object of considerable dispute.[] Fisher analyzed the results of the F2 (second filial) ratio and found them to be implausibly close to the exact ratio of 3 to 1.[18] Reproduction of his experiments has demonstrated the validity of his hypothesisbut, the results have continued to be a mystery for many, though it is often cited as an example of confirmation bias. This might arise if he detected an approximate 3 to 1 ratio early in his experiments with a small sample size, and continued collecting more data until the results conformed more nearly to an exact ratio. It is sometimes suggested that he may have censored his results, and that his seven traits each occur on a separate chromosome pair, an extremely unlikely occurrence if they were chosen at random. In fact, the genes Mendel studied occurred in only four linkage groups, and only one gene pair (out of 21 possible) is close enough to show deviation from independent assortment; this is not a pair that Mendel studied.

Gregor Mendel Some recent researchers have suggested that Fisher's criticisms of Mendel's work may have been exaggerated.[19][20]

107

References
[1] July 20 is his birthday; often mentioned is July 22, the date of his baptism. Biography of Mendel at the Mendel Museum (http:/ / www. mendel-museum. com/ eng/ 1online/ room1. htm) [4] Henig 2000, p.24. [5] Gregor Mendel, Alain F. Corcos, Floyd V. Monaghan, Maria C. Weber "Gregor Mendel's Experiments on Plant Hybrids: A Guided Study", Rutgers University Press, 1993. [6] Henig 2000, pp.19-21. [7] Henig 2000, pp.47-62. [8] Henig 2000, pp.15-17. [11] Henig 2000, pp.78-80. [12] Henig 2000, pp.134-138. [14] Mendel, J.G. (1866). Versuche ber Pflanzenhybriden Verhandlungen des naturforschenden Vereines in Brnn, Bd. IV fr das Jahr, 1865 Abhandlungen:347. For the English translation, see: [17] Henig 2000, pp.1-9. [18] Fisher, R. A. (1936). Has Mendel's work been rediscovered? Annals of Science 1:115137.

Bibliography
Smith, Jos A.; Cheryl Bardoe; Smith, Joseph A. (2006). Gregor Mendel: the friar who grew peas. Abrams Books for Young Readers. ISBN0-8109-5475-3. William Bateson Mendel, Gregor; Bateson, William (2009). Mendel's Principles of Heredity: A Defence, with a Translation of Mendel's Original Papers on Hybridisation (Cambridge Library Collection Life Sciences). Cambridge, UK: Cambridge University Press. ISBN1-108-00613-2. On-line Facsimile Edition: Electronic Scholarly Publishing, Prepared by Robert Robbins (http://www.esp.org/books/bateson/mendel/facsimile/ title3.html) Henig, Robin Marantz (2000). The Monk in the Garden: The Lost and Found Genius of Gregor Mendel, the Father of Genetics. Boston: Houghton Mifflin. ISBN978-0395-97765-1. Robert Lock, Recent Progress in the Study of Variation, Heredity and Evolution, London, 1906 Orel, Vtzslav (1996). Gregor Mendel: the first geneticist. Oxford [Oxfordshire]: Oxford University Press. ISBN0-19-854774-9. Reginald Punnett, Mendelism, Cambridge, 1905 Curt Stern and Sherwood ER (1966) The Origin of Genetics. Tudge, Colin (2000). In Mendel's footnotes: an introduction to the science and technologies of genes and genetics from the nineteenth century to the twenty-second. London: Vintage. ISBN0-09-928875-3. Waerden, B. L. V. D. (1968). "Mendel's Experiments". Centaurus 12 (4): 275288. doi: 10.1111/j.1600-0498.1968.tb00098.x (http://dx.doi.org/10.1111/j.1600-0498.1968.tb00098.x). PMID 4880928 (http://www.ncbi.nlm.nih.gov/pubmed/4880928). By Maria Malate about the Father of Genetics" Made by Gregor Mendel refutes allegations about "data smoothing" James Walsh, Catholic Churchmen in Science, Philadelphia: Dolphin Press, 1906 Ronald A. Fisher, "Has Mendel's Work Been Rediscovered?" Annals of Science, Volume 1, (1936): 115137. Discusses the possibility of fraud in his research.

Gregor Mendel

108

Further reading
Punnett, Reginald Crundall (1922). "Mendelism" (http://www.archive.org/stream/mendelism00punn#page/ n7/mode/2up). London: Macmillan. (1st Pub. 1905) Taylor, Monica (July/September 1922). "Abbot Mendel" (http://www.archive.org/stream/ dublinreview171londuoft#page/n7/mode/2up). Dublin Review. London: Burns, Oates and Washbourne. Windle, Bertram C. A. (1915). "Mendel and His Theory of Heredity" (http://www.archive.org/stream/ centuryofscienti00windrich#page/106/mode/2up). A Century of Scientific Thought and Other Essays. Burns & Oates.

External links
Works by Gregor Mendel (http://www.gutenberg.org/author/Gregor_Mendel) at Project Gutenberg Gregor Mendel (http://librivox.org/newcatalog/search.php?title=&author=Gregor+Mendel) public domain audiobooks from LibriVox 1913 Catholic Encyclopedia entry, "Mendel, Mendelism" (http://www.newadvent.org/cathen/10180b.htm) Augustinian Abbey of St. Thomas at Brno (http://www.opatbrno.cz/) Biography, bibliography and access to digital sources (http://vlp.mpiwg-berlin.mpg.de/people/ data?id=per116) in the Virtual Laboratory of the Max Planck Institute for the History of Science Biography of Gregor Mendel (http://mendel.imp.ac.at/mendeljsp/biography/biography.jsp) GCSE student (http://www.bbc.co.uk/schools/gcsebitesize/science/add_aqa/celldivision/inheritance3. shtml) Gregor Mendel (18221884) (http://www.accessexcellence.org/RC/AB/BC/Gregor_Mendel.html) Gregor Mendel Primary Sources (http://www.thomasmore.edu/library/mendel_collection.cfm?group =The Mendel Collection) Johann Gregor Mendel: Why his discoveries were ignored for 35 (72) years (http://www.weloennig.de/mendel. htm) (German) Masaryk University to rebuild Mendels greenhouse | Brno Now (http://brnonow.com/2009/02/ university-to-rebuild-mendels-greenhouse/) Mendel Museum of Genetics (http://www.mendelmuseum.muni.cz/en/) Mendel's Paper in English (http://www.mendelweb.org/Mendel.html) Online Mendelian Inheritance in Man (http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=OMIM) A photographic tour of St. Thomas' Abbey, Brno, Czech Republic (http://biology.clc.uc.edu/Fankhauser/ Travel/Berlin/for_web/Mendel_in_Brno.html) Works by or about Gregor Mendel (http://worldcat.org/identities/lccn-n50-36968) in libraries (WorldCat catalog)

Article Sources and Contributors

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Charles Darwin Source: http://en.wikipedia.org/w/index.php?oldid=568270158 Contributors: -x-Ultimate-Drummer-x-, 031586, 1 pwn nubs, 100% Christian, 11K, 168..., 1exec1, 200.191.188.xxx, 30daysinAK, 334a, 49erowens, 789suj98e7fh8sfdf98udfu8dfu9sdufdsf8377dh8nsdsncskj, 84user, @pple, A bit iffy, A vivid dreamer, A1 Aardvark, ABCD, ABIW, AGXD095, Aa77zz, Aaarrrggh, Aaron Bowen, Abdullais4u, Abeg92, Abhishikt, Academic Challenger, Acdun, Acetic Acid, Actam, Adam sk, Adamsan, Adashiel, Adraeus, Adrian.benko, AdultSwim, Afasmit, AgRince, Ahgrl123, Ahkitj, Ahoerstemeier, Aidan Croft, Akiioni, Aksi great, Al E., AlanHarmony, Alasdair, Alcoved id, AlexTG, Alexlange, Alexstep uk, Alfio, Ali K, Alienus, Alkaline Trio Rocks, All Hallow's Wraith, AllyUnion, Alphachimp, Alsandro, Altaar, Altofromgod, Amandajm, Amanduhh592, Amatulic, Amberrock, Amiodarone, Amitch, Anarchangel, Ancheta Wis, Andie142105, Andonic, Andre Engels, Andres, Andrevan, Andrew E. 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File:Charles Darwin seated crop.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Charles_Darwin_seated_crop.jpg License: Public Domain Contributors: Charles_Darwin_seated.jpg: Henry Maull (18291914) and John Fox (18321907) (Maull & Fox) UNIQ-ref-1-65981048430424dd-QINU derivative work: Beao File:Charles Darwin Signature.svg Source: http://en.wikipedia.org/w/index.php?title=File:Charles_Darwin_Signature.svg License: Public Domain Contributors: Connormah, Charles Darwin File:Charles Darwin 1816.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Charles_Darwin_1816.jpg License: Public Domain Contributors: ALE!, Anne97432, ArsniureDeGallium, Boo-Boo Baroo, Craigboy, Dave souza, Ecummenic, Finavon, Hystrix, LongLiveRock, Matt314, Mogelzahn, Richard001, Rotational, Wst, Zaphod, 5 anonymous edits File:Voyage of the Beagle-en.svg Source: http://en.wikipedia.org/w/index.php?title=File:Voyage_of_the_Beagle-en.svg License: Creative Commons Attribution-Share Alike Contributors: Smhur File:HMS Beagle by Conrad Martens.jpg Source: http://en.wikipedia.org/w/index.php?title=File:HMS_Beagle_by_Conrad_Martens.jpg License: Public Domain Contributors: Conrad Martens (1801 - 21 August 1878) File:Charles Darwin by G. Richmond.png Source: http://en.wikipedia.org/w/index.php?title=File:Charles_Darwin_by_G._Richmond.png License: Public Domain Contributors: Jdcollins13, Judithcomm, Shakko, 2 anonymous edits File:Darwin tree.png Source: http://en.wikipedia.org/w/index.php?title=File:Darwin_tree.png License: Public Domain Contributors: Cayambe, JMCC1, Jappalang, Jrockley, TimVickers, Trockennasenaffe, Werieth, 2 anonymous edits File:Emma Darwin.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Emma_Darwin.jpg License: Public Domain Contributors: Original uploader was Dave souza at en.wikipedia File:Charles and William Darwin.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Charles_and_William_Darwin.jpg License: Public Domain Contributors: Arria Belli, Closeapple, D-Kuru, Deadstar, Ecummenic, Fastfission, G.dallorto, Infrogmation, Onderwijsgek, Romary, Shakko, Wst, 4 anonymous edits File:Darwins Thinking Path.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Darwins_Thinking_Path.JPG License: Public Domain Contributors: Original uploader was Tedgrant at en.wikipedia File:Charles Darwin by Maull and Polyblank, 1855-crop.png Source: http://en.wikipedia.org/w/index.php?title=File:Charles_Darwin_by_Maull_and_Polyblank,_1855-crop.png License: Public Domain Contributors: Maull and Polyblank File:Charles Darwin by Julia Margaret Cameron 2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Charles_Darwin_by_Julia_Margaret_Cameron_2.jpg License: Public Domain Contributors: User:Davepape File:Editorial cartoon depicting Charles Darwin as an ape (1871).jpg Source: http://en.wikipedia.org/w/index.php?title=File:Editorial_cartoon_depicting_Charles_Darwin_as_an_ape_(1871).jpg License: Public Domain Contributors: Unknown, The Hornet is no longer in publication and it is very likely for a 20-year-old artist in 1871 to have died before 1939 File:1878 Darwin photo by Leonard from Woodall 1884 - cropped grayed partially cleaned.jpg Source: http://en.wikipedia.org/w/index.php?title=File:1878_Darwin_photo_by_Leonard_from_Woodall_1884_-_cropped_grayed_partially_cleaned.jpg License: Public Domain Contributors: Craigboy File:Man is But a Worm.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Man_is_But_a_Worm.jpg License: Public Domain Contributors: Punch magazine artist. File:Man dressed as Charles Darwin during Lyme Regis Fossil Festival 11.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Man_dressed_as_Charles_Darwin_during_Lyme_Regis_Fossil_Festival_11.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Mrjohncummings File:Annie Darwin.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Annie_Darwin.jpg License: Public Domain Contributors: Unknown photographer, uploaded by en:User:Duncharris File:VanityFair-Darwin2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:VanityFair-Darwin2.jpg License: Public Domain Contributors: (Source for author: Mark Bryant, "Going Ape Over Darwin", History Today, 1 April 2008) File:Charles Robert Darwin by John Collier.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Charles_Robert_Darwin_by_John_Collier.jpg License: Public Domain Contributors: User:Dcoetzee File:Wikisource-logo.svg Source: http://en.wikipedia.org/w/index.php?title=File:Wikisource-logo.svg License: logo Contributors: Guillom, INeverCry, Jarekt, MichaelMaggs, NielsF, Rei-artur, Rocket000 file:Fringilla coelebs chaffinch male edit2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Fringilla_coelebs_chaffinch_male_edit2.jpg License: Creative Commons Attribution-Sharealike 2.5 Contributors: MichaelMaggs Edited (sharpened, down-sampled, contrast corrected, noise removed) by: Arad File:Red Pencil Icon.png Source: http://en.wikipedia.org/w/index.php?title=File:Red_Pencil_Icon.png License: Creative Commons Zero Contributors: User:Peter coxhead File:DrepadinidaeSchnabel.jpg Source: http://en.wikipedia.org/w/index.php?title=File:DrepadinidaeSchnabel.jpg License: Public Domain Contributors: Artist/Knstler: John Gerrard Keulemans (1842-1912) File:Coccothraustes coccothraustes 1 (Marek Szczepanek).jpg Source: http://en.wikipedia.org/w/index.php?title=File:Coccothraustes_coccothraustes_1_(Marek_Szczepanek).jpg License: GNU Free Documentation License Contributors: Marek Szczepanek File:Cassin's Finch (male).jpg Source: http://en.wikipedia.org/w/index.php?title=File:Cassin's_Finch_(male).jpg License: Copyrighted free use Contributors: http://www.naturespicsonline.com/ File:Carpodacus roseus.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Carpodacus_roseus.jpg License: Creative Commons Attribution-Sharealike 3.0,2.5,2.0,1.0 Contributors: M.Nishimura File:PINTASSILGO ( Carduelis magellanica ).jpg Source: http://en.wikipedia.org/w/index.php?title=File:PINTASSILGO_(_Carduelis_magellanica_).jpg License: Creative Commons Attribution-Sharealike 2.0 Contributors: Dario Sanches File:Iiwi.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Iiwi.jpg License: Public Domain Contributors: Dysmorodrepanis, Maksim, 1 anonymous edits File:Euphonia violacea-2.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Euphonia_violacea-2.jpg License: Creative Commons Attribution-Sharealike 2.0 Contributors: Dario Sanches from So Paulo, Brasil file:Darwin's finches by Gould.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Darwin's_finches_by_Gould.jpg License: Public Domain Contributors: John Gould (14.Sep.1804 3.Feb.1881) File:Galapagos Islands topographic map-en.svg Source: http://en.wikipedia.org/w/index.php?title=File:Galapagos_Islands_topographic_map-en.svg License: Creative Commons Attribution-Sharealike 3.0,2.5,2.0,1.0 Contributors: Galapagos_Islands_topographic_map-de.svg: Eric Gaba (Sting - fr:Sting), translated by NordNordWest derivative work: MatthewStevens (talk) File:Flag of Ecuador.svg Source: http://en.wikipedia.org/w/index.php?title=File:Flag_of_Ecuador.svg License: Public Domain Contributors: President of the Republic of Ecuador, Zscout370 File:Orthographic projection centred over the Galapagos.png Source: http://en.wikipedia.org/w/index.php?title=File:Orthographic_projection_centred_over_the_Galapagos.png License: GNU Free Documentation License Contributors: FSII, Geo Swan, KJG2007, Telim tor, 3 anonymous edits File:School of Hammerhead sharks.jpg Source: http://en.wikipedia.org/w/index.php?title=File:School_of_Hammerhead_sharks.jpg License: Creative Commons Attribution-Sharealike 2.5 Contributors: Clark Anderson/Aquaimages () File:Sphyrna lewini galapagos.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Sphyrna_lewini_galapagos.jpg License: Creative Commons Attribution 2.0 Contributors: daren_ck File:Grapsus grapsus 2011.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Grapsus_grapsus_2011.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:P.Kurmelis File:Galapagos-satellite-esislandnames.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Galapagos-satellite-esislandnames.jpg License: Public Domain Contributors: labeled by Storpilot File:Galapagos SPOT 1178.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Galapagos_SPOT_1178.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: Cnes - 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File:Galapagos2007--39--08-22-07.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Galapagos2007--39--08-22-07.JPG License: Creative Commons Attribution-ShareAlike 3.0 Unported Contributors: Iris Diensthuber File:Tour of the Galapagos.OGG Source: http://en.wikipedia.org/w/index.php?title=File:Tour_of_the_Galapagos.OGG License: Public Domain Contributors: Kevin Ward File:Journey to Galapagos.OGG Source: http://en.wikipedia.org/w/index.php?title=File:Journey_to_Galapagos.OGG License: Public Domain Contributors: Kevin Ward File:North Seymour Island in the Galapagos about to land on shore photo by Alvaro Sevilla Design.JPG Source: http://en.wikipedia.org/w/index.php?title=File:North_Seymour_Island_in_the_Galapagos_about_to_land_on_shore_photo_by_Alvaro_Sevilla_Design.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Alvarosevilladesign File:Alvaro Sevilla Design Isla Santa Cruz Galapagos foto tomada desde el avin.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Alvaro_Sevilla_Design_Isla_Santa_Cruz_Galapagos_foto_tomada_desde_el_avin.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Alvarosevilladesign File:Chlorophyll concentration off the Galapagos archipelago during El Nio and La Nia.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Chlorophyll_concentration_off_the_Galapagos_archipelago_during_El_Nio_and_La_Nia.jpg License: Public Domain Contributors: GSFC Ocean Color team and GeoEye. File:Sea surface temperature & chlorophyll concentrations off the Galapagos archipelago.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Sea_surface_temperature_&_chlorophyll_concentrations_off_the_Galapagos_archipelago.jpg License: Public Domain Contributors: Robert Simmon and Jesse Allen File:The Galpagos tortoise or Galpagos giant tortoise (Chelonoidis nigra) - Santa Cruz Island.jpeg Source: http://en.wikipedia.org/w/index.php?title=File:The_Galpagos_tortoise_or_Galpagos_giant_tortoise_(Chelonoidis_nigra)_-_Santa_Cruz_Island.jpeg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Alvarosevilladesign File:Gallapagos Islands 1684.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Gallapagos_Islands_1684.jpg License: Public Domain Contributors: James Burney File:The marine iguana (Amblyrhynchus cristatus) Galpagos Islands Santa Cruz.JPG Source: http://en.wikipedia.org/w/index.php?title=File:The_marine_iguana_(Amblyrhynchus_cristatus)_Galpagos_Islands_Santa_Cruz.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Alvarosevilladesign File:Flag of the Galpagos Islands.svg Source: http://en.wikipedia.org/w/index.php?title=File:Flag_of_the_Galpagos_Islands.svg License: Public Domain Contributors: Original uploader was Zscout370 at en.wikipedia File:Water taxi in Puerto Ayora on the Island of Santa Cruz in the Galapagos photo by Alvaro Sevilla Design.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Water_taxi_in_Puerto_Ayora_on_the_Island_of_Santa_Cruz_in_the_Galapagos_photo_by_Alvaro_Sevilla_Design.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Alvarosevilladesign File:Iguanamarina.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Iguanamarina.jpg License: GNU Free Documentation License Contributors: Anrie, Eugene van der Pijll, John feather, Juiced lemon, Kilom691, Martini, Oersted, Rmih, SCEhardt, 3 anonymous edits File:Land Iguana.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Land_Iguana.jpg License: GNU Free Documentation License Contributors: Badseed, Dr Brains, Eugene van der Pijll, Haplochromis, Juiced lemon, Kersti Nebelsiek, Maksim, Martini, Romanm, Rmih, 6 anonymous edits File:SulaNebouxi.jpg Source: http://en.wikipedia.org/w/index.php?title=File:SulaNebouxi.jpg License: Creative Commons Attribution-ShareAlike 3.0 Unported Contributors: Marc Figueras File:Tortuga Bay Galapagos Island of Santa Cruz.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Tortuga_Bay_Galapagos_Island_of_Santa_Cruz.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Alvarosevilladesign File:Great Frigatebird bird 2011.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Great_Frigatebird_bird_2011.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:P.Kurmelis File:Gigantic Turtle on the Island of Santa Cruz in the Galapagos.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Gigantic_Turtle_on_the_Island_of_Santa_Cruz_in_the_Galapagos.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Alvarosevilladesign File:Two ducks in the galapagos islands - santa cruz, ecuador.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Two_ducks_in_the_galapagos_islands_-_santa_cruz,_ecuador.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Alvarosevilladesign File:Tortuga Bay - Island of Santa Cruz, Galapagos.JPG Source: http://en.wikipedia.org/w/index.php?title=File:Tortuga_Bay_-_Island_of_Santa_Cruz,_Galapagos.JPG License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Alvarosevilladesign File:Grapsus grapsus Galapagos Islands.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Grapsus_grapsus_Galapagos_Islands.jpg License: Public Domain Contributors: Lieutenant Elizabeth Crapo, NOAA Corps File:Spheniscus mendiculus juvenile.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Spheniscus_mendiculus_juvenile.jpg License: Creative Commons Attribution-Sharealike 2.5 Contributors: Original uploader was Aquaimages at en.wikipedia File:PD-icon.svg Source: http://en.wikipedia.org/w/index.php?title=File:PD-icon.svg License: Public Domain Contributors: Alex.muller, Anomie, Anonymous Dissident, CBM, MBisanz, PBS, Quadell, Rocket000, Strangerer, Timotheus Canens, 1 anonymous edits File:PSM V57 D097 Hms beagle in the straits of magellan.png Source: http://en.wikipedia.org/w/index.php?title=File:PSM_V57_D097_Hms_beagle_in_the_straits_of_magellan.png License: Public Domain Contributors: Ineuw, 1 anonymous edits File:Naval Ensign of the United Kingdom.svg Source: http://en.wikipedia.org/w/index.php?title=File:Naval_Ensign_of_the_United_Kingdom.svg License: Public Domain Contributors: AnonMoos, Avicennasis, Bender235, Cycn, Dancingwombatsrule, Ec.Domnowall, Fry1989, Homo lupus, Pumbaa80, Stunteltje, Xiengyod, Yaddah, 3 anonymous edits File:HMS Beagle 1832 longitudinal section larger.jpg Source: http://en.wikipedia.org/w/index.php?title=File:HMS_Beagle_1832_longitudinal_section_larger.jpg License: Public Domain Contributors: HMS_Beagle_1832_longitudinal_section.jpg: Original drawing by F. G. King, digital photo by User:Alexei Kouprianov derivative work: Themadchopper (talk) File:TheBeagleLaidAshore.png Source: http://en.wikipedia.org/w/index.php?title=File:TheBeagleLaidAshore.png License: Public Domain Contributors: Conrad Martens (1834), engraved (1838) by Thomas Landseer and published in the year by H. Colburn in The Narrative of the Surveying Voyages of HMS Adventure and Beagle File:HMSBeagle.jpg Source: http://en.wikipedia.org/w/index.php?title=File:HMSBeagle.jpg License: Public Domain Contributors: Dave souza, Interpretix, Kresspahl, Makthorpe, Poppy File:General Chart of Australia (Discoveries in Australia).jpg Source: http://en.wikipedia.org/w/index.php?title=File:General_Chart_of_Australia_(Discoveries_in_Australia).jpg License: Public Domain Contributors: John Arrowsmith File:The construction of the HMS Beagle Full Size Replica in the Nao Victoria Museum - Punta Arenas - Chile.jpg Source: http://en.wikipedia.org/w/index.php?title=File:The_construction_of_the_HMS_Beagle_Full_Size_Replica_in_the_Nao_Victoria_Museum_-_Punta_Arenas_-_Chile.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Symcovington File:Construccin de la rplica del HMS Beagle 143 - State of the art 2013 03 20 portside view from the Nao Victoria.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Construccin_de_la_rplica_del_HMS_Beagle_143_-_State_of_the_art_2013_03_20_portside_view_from_the_Nao_Victoria.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Symcovington File:Construccin de la rplica del HMS Beagle 144 - State of the art 2013 03 21 starboard view.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Construccin_de_la_rplica_del_HMS_Beagle_144_-_State_of_the_art_2013_03_21_starboard_view.jpg License: Creative Commons Attribution-Sharealike 3.0 Contributors: User:Symcovington File:Biological_classification_L_Pengo_vflip.svg Source: http://en.wikipedia.org/w/index.php?title=File:Biological_classification_L_Pengo_vflip.svg License: unknown Contributors: Adrignola, ArnoLagrange, Nisetpdajsankha, Pavel55, Pengo, 1 anonymous edits File:Undiscovered species chart.png Source: http://en.wikipedia.org/w/index.php?title=File:Undiscovered_species_chart.png License: Public Domain Contributors: KVDP File:John Ray from NPG.jpg Source: http://en.wikipedia.org/w/index.php?title=File:John_Ray_from_NPG.jpg License: Public Domain Contributors: User:Dcoetzee File:Carolus Linnaeus (cleaned up version).jpg Source: http://en.wikipedia.org/w/index.php?title=File:Carolus_Linnaeus_(cleaned_up_version).jpg License: Public Domain Contributors: Original painting by Alexander Roslin. 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File:Office-book.svg Source: http://en.wikipedia.org/w/index.php?title=File:Office-book.svg License: Public Domain Contributors: Chris Down/Tango project File:Lichte en zwarte versie berkenspanner.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Lichte_en_zwarte_versie_berkenspanner.jpg License: GNU Free Documentation License Contributors: Original uploader was Martinowksy at nl.wikipedia (Original text : Maarten Sanne) Image:Darwin's finches.jpeg Source: http://en.wikipedia.org/w/index.php?title=File:Darwin's_finches.jpeg License: Public Domain Contributors: John Gould (14.Sep.1804 - 3.Feb.1881) Image:Life cycle of a sexually reproducing organism.svg Source: http://en.wikipedia.org/w/index.php?title=File:Life_cycle_of_a_sexually_reproducing_organism.svg License: Public Domain Contributors: Wykis/Wykis on Wikipedia Image:Antibiotic resistance.svg Source: http://en.wikipedia.org/w/index.php?title=File:Antibiotic_resistance.svg License: Public Domain Contributors: Wykis Image:Pavo cristatus albino001xx.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Pavo_cristatus_albino001xx.jpg License: GNU Free Documentation License Contributors: Pavo cristatus; 2004 by M. 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