document_title: History of Australia (1788–1850)

question_text: when did the first english settlers come to australia

text: History of Australia (1788–1850)

text: Colonisation[edit]

text: It is commonly reported that the  colonisation of Australia was driven by the need to address overcrowding in the British prison system, and the fact of the British losing the Thirteen Colonies of America in the American Revolution; however, it was simply not economically viable to transport convicts halfway around the world for this reason alone.[1] Many convicts were either skilled tradesmen or farmers who had been convicted for trivial crimes and were sentenced to seven years transportation, the time required to set up the infrastructure for the new colony. Convicts were often given pardons prior to or on completion of their sentences and were allocated parcels of land to farm.

text: Sir Joseph Banks, the eminent scientist who had accompanied Lieutenant James Cook on his 1770 voyage, recommended Botany Bay as a suitable site.[2] Banks accepted an offer of assistance which by the American Loyalist James Matra in July 1783. Matra had visited Botany Bay with Banks in 1770 as a junior officer on the Endeavour commanded by James Cook. Under Banks's guidance, he rapidly produced "A Proposal for Establishing a Settlement in New South Wales" (24 August 1783), with a fully developed set of reasons for a colony composed of American Loyalists, Chinese and South Sea Islanders (but not convicts).[3]

text: Following an interview with Secretary of State Lord Sydney in March 1784, Matra amended his proposal to include convicts as settlers.[4] Matra’s plan can be seen to have “provided the original blueprint for settlement in New South Wales”.[5] A cabinet memorandum December 1784 shows the Government had Matra’s plan in mind when considering the creation of a settlement in New South Wales.[5][6] The London Chronicle of 12 October 1786 said: “Mr. Matra, an Officer of the Treasury, who, sailing with Capt. Cook, had an opportunity of visiting Botany Bay, is the Gentleman who suggested the plan to Government of transporting convicts to that island”. The Government also incorporated into the colonisation plan the project for settling Norfolk Island, with its attractions of timber and flax, proposed by Banks’s Royal Society colleagues, Sir John Call and Sir George Young.[7]

text: On 13 May 1787, the First Fleet of 11 ships and about 1,530 people (736 convicts, 17 convicts' children, 211 marines, 27 marines' wives, 14 marines' children and about 300 officers and others) under the command of Captain Arthur Phillip set sail for Botany Bay.[8][9][10] A few days after arrival at Botany Bay the fleet moved to the more suitable Port Jackson where a settlement was established at Sydney Cove on 26 January 1788.[11] This date later became Australia's national day, Australia Day. The colony was formally proclaimed by Governor Phillip on 7 February 1788 at Sydney. Sydney Cove offered a fresh water supply and a safe harbour, which Philip famously described as:[12]

text:
“

being with out exception the finest Harbour in the World [...] Here a Thousand Sail of the Line may ride in the most perfect Security.

”

text: Phillip named the settlement after the Home Secretary, Thomas Townshend, 1st Baron Sydney (Viscount Sydney from 1789). The only people at the flag raising ceremony and the formal taking of possession of the land in the name of King George III were Phillip and a few dozen marines and officers from the Supply, the rest of the ship's company and the convicts witnessing it from on board ship. The remaining ships of the Fleet were unable to leave Botany Bay until later on 26 January because of a tremendous gale.[13] The new colony was formally proclaimed as the Colony of New South Wales on 7 February.[14]

text: On 24 January 1788 a French expedition of two ships led by Admiral Jean-François de La Pérouse had arrived off Botany Bay, on the latest leg of a three-year voyage that had taken them from Brest, around Cape Horn, up the coast from Chile to California, north-west to Kamchatka, south-east to Easter Island, north-west to Macao, and on to the Philippines, the Friendly Isles, Hawaii and Norfolk Island.[15] Though amicably received, the French expedition was a troublesome matter for the British, as it showed the interest of France in the new land.

text: Nevertheless, on 2 February Lieutenant King, at Phillip's request, paid a courtesy call on the French and offered them any assistance they may need.[13] The French made the same offer to the British, as they were much better provisioned than the British and had enough supplies to last three years.[13] Neither of these offers was accepted. On 10 March[13] the French expedition, having taken on water and wood, left Botany Bay, never to be seen again. Phillip and La Pérouse never met. La Pérouse is remembered in a Sydney suburb of that name. Various other French geographical names along the Australian coast also date from this expedition.

text: Governor Phillip was vested with complete authority over the inhabitants of the colony. Phillip's personal intent was to establish harmonious relations with local Aboriginal people and try to reform as well as discipline the convicts of the colony. Phillip and several of his officers – most notably Watkin Tench – left behind journals and accounts of which tell of immense hardships during the first years of settlement. Often Phillip's officers despaired for the future of New South Wales. Early efforts at agriculture were fraught and supplies from overseas were few and far between. Between 1788 and 1792 about 3546 male and 766 female convicts were landed at Sydney – many "professional criminals" with few of the skills required for the establishment of a colony. Many new arrivals were also sick or unfit for work and the conditions of healthy convicts only deteriorated with hard labour and poor sustenance in the settlement. The food situation reached crisis point in 1790 and the Second Fleet which finally arrived in June 1790 had lost a quarter of its "passengers" through sickness, while the condition of the convicts of the Third Fleet appalled Phillip. From 1791 however, the more regular arrival of ships and the beginnings of trade lessened the feeling of isolation and improved supplies.[16]

text: In 1792, two French ships, La Recherche and L'Espérance anchored in a harbour near Tasmania's southernmost point they called Recherche Bay. This was at a time when Britain and France were trying to be the first to discover and colonise Australia. The expedition carried scientists and cartographers, gardeners, artists and hydrographers who, variously, planted, identified, mapped, marked, recorded and documented the environment and the people of the new lands that they encountered at the behest of the fledgling Société D'Histoire Naturelle.

text: White settlement began with a consignment of English convicts, guarded by a detachment of the Royal Marines, a number of whom subsequently stayed in the colony as settlers. Their view of the colony and their place in it was eloquently stated by Captain David Collins: "From the disposition to crimes and the incorrigible character of the major part of the colonists, an odium was, from the first, illiberally thrown upon the settlement; and the word "Botany Bay" became a term of reproach that was indiscriminately cast upon every one who resided in New South Wales. But let the reproach light upon those who have used it as such... if the honour of having deserved well of one's country be attainable by sacrificing good name, domestic comforts, and dearest connections in her service, the officers of this settlement have justly merited that distinction".[17]

text: Convicts and free settlers[edit]

text: When the Bellona transport came to anchor in Sydney Cove on 16 January 1793, she brought with her the first immigrant free settlers. They were: Thomas Rose, a farmer from Dorset, his wife and four children; he was allowed a grant of 120 acres; Frederic Meredith, who had formerly been at Sydney with HMS Sirius; Thomas Webb (who had also been formerly at Sydney with the Sirius), his wife, and his nephew, Joseph Webb; Edward Powell, who had formerly been at Sydney with the Juliana transport, and who married a free woman after his arrival. Thomas Webb and Edward Powell each received a grant of 80 acres; and Joseph Webb and Frederic Meredith received 60 acres each.

text: The conditions they had come out under were that they should be provided with a free passage, be furnished with agricultural tools and implements by the Government, have two years' provisions, and have grants of land free of expense. They were likewise to have the labour of a certain number of convicts, who were also to be provided with two years' rations and one year's clothing from the public stores. The land assigned to them was some miles to the westward of Sydney, at a place named by the settlers, "Liberty Plains". It is now the area covered mainly by the suburbs of Strathfield and Homebush.

text: One in three convicts transported after 1798 was Irish, about a fifth of whom were transported in connection with the political and agrarian disturbances common in Ireland at the time. While the settlers were reasonably well-equipped, little consideration had been given to the skills required to make the colony self-supporting – few of the first wave convicts had farming or trade experience (nor the soldiers), and the lack of understanding of Australia's seasonal patterns saw initial attempts at farming fail, leaving only what animals and birds the soldiers were able to shoot. The colony nearly starved, and Phillip was forced to send a ship to Batavia (Jakarta) for supplies. Some relief arrived with the Second Fleet in 1790, but life was extremely hard for the first few years of the colony.

text: Convicts were usually sentenced to seven or fourteen years' penal servitude, or "for the term of their natural lives". Often these sentences had been commuted from the death sentence, which was technically the punishment for a wide variety of crimes. Upon arrival in a penal colony, convicts would be assigned to various kinds of work. Those with trades were given tasks to fit their skills (stonemasons, for example, were in very high demand) while the unskilled were assigned to work gangs to build roads and do other such tasks. Female convicts were usually assigned as domestic servants to the free settlers, many being forced into prostitution.[18]

text: Where possible, convicts were assigned to free settlers who would be responsible for feeding and disciplining them; in return for this, the settlers were granted land. This system reduced the workload on the central administration. Those convicts who weren't assigned to settlers were housed at barracks such as the Hyde Park Barracks or the Parramatta female factory.

text: Convict discipline was harsh; convicts who would not work or who disobeyed orders were punished by flogging, being put in stricter confinement (e.g. leg-irons), or being transported to a stricter penal colony. The penal colonies at Port Arthur and Moreton Bay, for instance, were stricter than the one at Sydney, and the one at Norfolk Island was strictest of all. Convicts were assigned to work gangs to build roads, buildings, and the like. Female convicts, who made up 20% of the convict population, were usually assigned as domestic help to soldiers. Those convicts who behaved were eventually issued with ticket of leave, which allowed them a certain degree of freedom. Those who saw out their full sentences or were granted a pardon usually remained in Australia as free settlers, and were able to take on convict servants themselves.

text: In 1789 former convict James Ruse produced the first successful wheat harvest in NSW. He repeated this success in 1790 and, because of the pressing need for food production in the colony, was rewarded by Governor Phillip with the first land grant made in New South Wales. Ruse's 30 acre grant at Rose Hill, near Parramatta, was aptly named 'Experiment Farm'.[19] This was the colony's first successful farming enterprise, and Ruse was soon joined by others. The colony began to grow enough food to support itself, and the standard of living for the residents gradually improved.

text: In 1804 the Castle Hill convict rebellion was led by around 200 escaped, mostly Irish convicts, although it was broken up quickly by the New South Wales Corps. On 26 January 1808, there was a military rebellion against Governor Bligh led by John Macarthur. Following this, Governor Lachlan Macquarie was given a mandate to restore government and discipline in the colony. When he arrived in 1810, he forcibly deported the NSW Corps and brought the 73rd regiment to replace them.

text:
13 May 1787 – The 11 ships of the First Fleet leave Portsmouth under the command of Captain Arthur Phillip. Different accounts give varying numbers of passengers but the fleet consisted of at least 1,350 persons of whom 780 were convicts and 570 were free men, women and children and the number included four companies of marines. About 20% of the convicts were women and the oldest convict was 82. About 50% of the convicts had been tried in Middlesex and most of the rest were tried in the county assizes of Devon, Kent and Sussex
18 January 1788 – The First Fleet arrived in Botany Bay but the landing party was not impressed with the site, and moved the fleet to Port Jackson, landing in Sydney Cove on 26 January 1788 (now celebrated as Australia Day)
 April 1789  – A suspicious outbreak of smallpox decimates local tribes. Research by Michael J Bennett[20] published in 2009, corroborated independently by Christopher Warren[21] in 2013, demonstrates this outbreak was most likely deliberate.
1790 – the Second Fleet of convicts arrives in Sydney Cove.
1791 – Third Fleet of convicts arrives
1793 – January: the first free settlers arrive in NSW.
1793 – March–April: visit of the expedition led by Alessandro Malaspina.
1825 - May: Founding of Brisbane
14 June 1825 – the colony of Van Diemen's Land is established in its own right; its name is officially changed to Tasmania on 1 January 1856. The first settlement was made at Risdon, Tasmania on 11 September 1803 when Lieut John Bowen landed with about 50 settlers, crew, soldiers and convicts. The site proved unsuitable and was abandoned in August 1804. Lieut-Col David Collins finally established a successful settlement at Hobart in February 1804 with a party of about 260 people, including 178 convicts. (Collins had previously attempted a settlement in Victoria.) Convict ships were sent from England directly to the colony from 1812 to 1853 and over the 50 years from 1803–1853 around 67,000 convicts were transported to Tasmania. About 14,492 were Irish but many of them had been sentenced in English and Scottish courts. Some were also tried locally in other Australian colonies. The Indefatigable brought the first convicts direct from England on 19 October 1812 and by 1820 there were about 2,500 convicts in the colony. By the end of 1833 the number had increased to 14,900 convicts of whom 1864 were females. About 1,448 held ticket of leave, 6,573 were assigned to settlers and 275 were recorded as "absconded or missing". In 1835 there were over 800 convicts working in chain-gangs at the penal station at Port Arthur which operated from 1830 to 1877. Convicts were transferred to Van Diemen's Land from Sydney and, in later years, from 1841 to 1847, from Melbourne. Between 1826 and 1840, there were at least 19 ship loads of convicts sent from Van Diemen's Land to Norfolk Island and at other times they were sent from Norfolk Island to Van Diemen's Land.
21 January 1827 – Western Australia was established when a small British settlement was established at King George's Sound (Albany) by Major Edmund Lockyer who was to provide a deterrent to the French presence in the area. On 18 June 1829 the new Swan River Colony was officially proclaimed with Captain James Stirling as the first Governor. Except for the settlement at King George's Sound, the colony was never really a part of NSW. King George's Sound was handed over in 1831. In 1849 the colony was proclaimed a British penal settlement and the first convicts arrived in 1850. Rottnest Island, off the coast of Perth, became the colony's convict settlement in 1838 and was used for local colonial offenders. Around 9,720 British convicts were sent directly to the colony in 43 ships between 1850 and 1868. The convicts were sought by local settlers because of the shortage of labour needed to develop the region. On 9 January 1868, Australia's last convict ship, the Hougoumont brought its final cargo of 269 convicts. Convicts sent to Western Australia were sentenced to terms of 6, 7, 10, 14 and 15 years and some reports suggest that their literacy rate was around 75% as opposed to 50% for those sent to NSW and Tasmania. About a third of the convicts left the Swan River Colony after serving their time.
1835 – the Proclamation of Governor Bourke, issued by the Colonial Office and sent to the Governor with Despatch 99 of 10 October 1835, implements the doctrine of terra nullius upon which British settlement was based. Reinforcing the British assertion that the land belonged to no one prior to the British Crown taking possession of it, it effectively quashes pre-existing treaties with Aboriginal peoples (e.g. that signed by John Batman). Its publication in the Colony means that from then on, all people found occupying land without the authority of the government would be considered illegal trespassers. Aboriginal people therefore could not sell or assign the land, nor could an individual person acquire it, other than through distribution by the Crown.[22]
28 December 1836 – the British province of South Australia was established. In 1842 it became a crown colony and on 22 July 1861 its area was extended westwards to its present boundary and more area was taken from New South Wales. South Australia was never a British convict colony and between 1836–1840 about 13,400 immigrants arrived in the area. 24,900 more arrived between 1841–1850. Some escaped convicts did settle in the area and no doubt a number of ex-convicts moved there from other colonies. On 4 January 1837 Governor Hindmarsh proclaimed that any offenders convicted in South Australia, and being under sentence of transportation, were to be transported to either New South Wales or Van Diemens Land, by the first opportunity.[23]
1841 – New Zealand is separated from New South Wales
1851 – Victoria is separated from New South Wales (formerly known as the Port Phillip District of NSW). Apart from castaways and runaway convicts in the 1790s, the first attempt at settlement was made on 13 October 1803 by Lieut. David Collins and his party of soldiers and convicts. Harsh conditions convinced him to abandon the settlement in January 1804. He moved on to Tasmania and it was not until the Henty brothers landed in Portland Bay on 19 November 1834 and John Batman settled on the site of Melbourne that the Port Phillip District was officially sanctioned on 10 April 1837. The first immigrant ships arrived at Port Phillip in 1839. Apart from those involved in early attempts at settlement in 1803 and 1826, the only convicts sent directly to Victoria from Britain were about 1,750 convicts known as the "Exiles" and they arrived between 1844–1849. They were sometimes called the "Pentonvillians" because most of them came from Pentonville Probationary Prison in England. Many ex-convicts and convicts on Tickets of Leave and Conditional Pardons also moved to Port Phillip from Van Diemen's Land.
10 December 1859 – Queensland is separated from New South Wales. In 1824 the explorer Lieut. John Oxley took a party of 30 convicts and established a penal colony at Redcliffe. Known as the Moreton Bay Settlement, this later moved to the site now called Brisbane. The name Brisbane Town was in use by 1825 and the main inhabitants in the area were the convicts of the Moreton Bay Penal Station until it was closed in 1839. Around 2,280 convicts were sent to the settlement between 1824–1839 and at the end of 1836 the convict population numbered 337. The first free settlers moved to the district in 1838 and others followed in 1840.
23 December 1862 – the area of Queensland is increased.
1863 – control of the Northern Territory is granted to the Province (later State) of South Australia. In 1825 the area occupied today by Northern Territory was incorporated into the colony of New South Wales. It was first settled by Europeans in 1824 at Fort Dundas, Port Essington. Its capital city, Darwin was established in 1869 and was originally known as Palmerston. On 1 January 1911, the Northern Territory as we know it today was separated from South Australia and became part of the Commonwealth of Australia.
1 January 1901 – the  Federation of Australian States to form the Commonwealth of Australia.
1911 – The Federal Capital Territory is established.

text: Exploration[edit]

text: In October 1795 George Bass and Matthew Flinders, accompanied by William Martin sailed the boat Tom Thumb out of Port Jackson to Botany Bay and explored the Georges River further upstream than had been done previously by the colonists. Their reports on their return led to the settlement of Banks' Town.[25] In March 1796 the same party embarked on a second voyage in a similar small boat, which they also called the Tom Thumb.[26] During this trip they travelled as far down the coast as Lake Illawarra, which they called Tom Thumb Lagoon. They discovered and explored Port Hacking. In 1798–99, Bass and Flinders set out in a sloop and circumnavigated Van Diemen's Land, thus proving it to be an island.[27]

text: Aboriginal guides and assistance in the European exploration of the colony were common and often vital to the success of missions. In 1801–02 Matthew Flinders in The Investigator lead the first circumnavigation of Australia. Aboard ship was the Aboriginal explorer Bungaree, of the Sydney district, who became the first person born on the Australian continent to circumnavigate the Australian continent.[27] Previously, the famous Bennelong and a companion had become the first people born in the area of New South Wales to sail for Europe, when, in 1792 they accompanied Governor Phillip to England and were presented to King George III.[27]

text: In 1813, Gregory Blaxland, William Lawson and William Wentworth succeeded in crossing the formidable barrier of forested gulleys and sheer cliffs presented by the Blue Mountains, west of Sydney, by following the ridges instead of looking for a route through the valleys. At Mount Blaxland they looked out over "enough grass to support the stock of the colony for thirty years", and expansion of the British settlement into the interior could begin.[28]

text: In 1824 the Governor Sir Thomas Brisbane, commissioned Hamilton Hume and former Royal Navy Captain William Hovell to lead an expedition to find new grazing land in the south of the colony, and also to find an answer to the mystery of where New South Wales's western rivers flowed. Over 16 weeks in 1824–25, Hume and Hovell journeyed to Port Phillip and back. They made many important discoveries including the Murray River (which they named the Hume), many of its tributaries, and good agricultural and grazing lands between Gunning, New South Wales and Corio Bay, Victoria.[29]

text: Charles Sturt led an expedition along the Macquarie River in 1828 and discovered the Darling River. A theory had developed that the inland rivers of New South Wales were draining into an inland sea. Leading a second expedition in 1829, Sturt followed the Murrumbidgee River into a 'broad and noble river', the Murray River, which he named after Sir George Murray, secretary of state for the colonies. His party then followed this river to its junction with the Darling River, facing two threatening encounters with local Aboriginal people along the way. Sturt continued down river on to Lake Alexandrina, where the Murray meets the sea in South Australia. Suffering greatly, the party had to then row back upstream hundreds of kilometres for the return journey.[30]

text: Surveyor General Sir Thomas Mitchell conducted a series of expeditions from the 1830s to 'fill in the gaps' left by these previous expeditions. He was meticulous in seeking to record the original Aboriginal place names around the colony, for which reason the majority of place names to this day retain their Aboriginal titles.[31]

text: The Polish scientist/explorer Count Paul Edmund Strzelecki conducted surveying work in the Australian Alps in 1839 and became the first European to ascend Australia's highest peak, which he named Mount Kosciuszko in honour of the Polish patriot Tadeusz Kosciuszko.[32]

text: Growth of free settlement[edit]

text: Traditional Aboriginal society had been governed by councils of elders and a corporate decision-making process, but the first European-style governments established after 1788 were autocratic and run by appointed governors – although English law was transplanted into the Australian colonies by virtue of the doctrine of reception, thus notions of the rights and processes established by the Magna Carta and the Bill of Rights 1689 were brought from Britain by the colonists. Agitation for representative government began soon after the settlement of the colonies.[33]

text: The Second Fleet in 1790 brought to Sydney two men who were to play important roles in the colony's future. One was D'Arcy Wentworth, whose son, William Charles, went on to be an explorer, to found Australia's first newspaper and to become a leader of the movement to abolish convict transportation and establish representative government. The other was John Macarthur, a Scottish army officer and founder of the Australian wool industry, which laid the foundations of Australia's future prosperity. Macarthur was a turbulent element: in 1808 he was one of the leaders of the Rum Rebellion against the governor, William Bligh.

text: From about 1815 the colony, under the governorship of Lachlan Macquarie, began to grow rapidly as free settlers arrived and new lands were opened up for farming. Despite the long and arduous sea voyage, settlers were attracted by the prospect of making a new life on virtually free Crown land. From the late 1820s settlement was only authorised in the limits of location, known as the Nineteen Counties.

text: Many settlers occupied land without authority and beyond these authorised settlement limits: they were known as squatters and became the basis of a powerful landowning class. As a result of opposition from the labouring and artisan classes, transportation of convicts to Sydney ended in 1840, although it continued in the smaller colonies of Van Diemen's Land (first settled in 1803, later renamed Tasmania) and Moreton Bay (founded 1824, and later renamed Queensland) for a few years more.

text: The Swan River Settlement (as Western Australia was originally known), centred on Perth, was founded in 1829. The colony suffered from a long-term shortage of labour, and by 1850 local capitalists had succeeded in persuading London to send convicts. (Transportation did not end until 1868.) New Zealand was part of New South Wales until 1840 when it became a separate colony.

text: The first governments established after 1788 were autocratic and each colony was governed by a British Governor, appointed by the British monarch. There was considerable unhappiness with the way some of the colonies were run. In most cases the administration of the early colonies was carried out by the British military. The New South Wales Corps, which was in charge of New South Wales, became known as the "Rum Corps", due to its stranglehold on the distribution of rum, which was used as a makeshift currency at the time. In New South Wales this led to the "Rum Rebellion". Although English law was transplanted into the Australian colonies by virtue of the doctrine of reception, thus notions of the rights and processes established by the Magna Carta and the Bill of Rights 1689 were brought from Britain by the colonists. Agitation for representative government began soon after the settlement of the colonies.[33]

text: The oldest legislative body in Australia, the New South Wales Legislative Council, was created in 1825 as an appointed body to advise the Governor of New South Wales. William Wentworth established the Australian Patriotic Association (Australia's first political party) in 1835 to demand democratic government for New South Wales. The reformist attorney general, John Plunkett, sought to apply Enlightenment principles to governance in the colony, pursuing the establishment of equality before the law, first by extending jury rights to emancipists, then by extending legal protections to convicts, assigned servants and Aborigines. Plunkett twice charged the colonist perpetrators of the Myall Creek massacre of Aborigines with murder, resulting in a conviction and his landmark Church Act of 1836 disestablished the Church of England and established legal equality between Anglicans, Catholics, Presbyterians and later Methodists.[34]

text: In 1840, the Adelaide City Council and the Sydney City Council were established. Men who possessed 1000 pounds worth of property were able to stand for election and wealthy landowners were permitted up to four votes each in elections. Australia's first parliamentary elections were conducted for the New South Wales Legislative Council in 1843, again with voting rights (for males only) tied to property ownership or financial capacity. Voter rights were extended further in New South Wales in 1850 and elections for legislative councils were held in the colonies of Victoria, South Australia and Tasmania.[35]

text: By the mid-19th century, there was a strong desire for representative and responsible government in the colonies of Australia, later fed by the democratic spirit of the goldfields and the ideas of the great reform movements sweeping Europe, the United States and the British Empire. The end of convict transportation accelerated reform in the 1840s and 1850s. The Australian Colonies Government Act [1850] was a landmark development which granted representative constitutions to New South Wales, Victoria, South Australia and Tasmania and the colonies enthusiastically set about writing constitutions which produced democratically progressive parliaments – though the constitutions generally maintained the role of the colonial upper houses as representative of social and economic "interests" and all established constitutional monarchies with the British monarch as the symbolic head of state.[36]

text: Timeline[edit]

text:
1788 – New South Wales, according to Arthur Phillip's amended Commission dated 25 April 1787, includes "all the islands adjacent in the Pacific Ocean" and running westward to the 135th meridian east. These islands included the current islands of New Zealand, which was administered as part of New South Wales.[a]
1824 — Brisbane is founded
1825 – New South Wales's western border is extended to 129° E. In the same year, Van Diemen's Land is proclaimed.
1829 – Swan River Colony is declared by Charles Fremantle for Britain.
1832 – Swan River Colony has its name changed to Western Australia.
1836 – South Australia is proclaimed with its western border at 132° E.
1840 – New Zealand is proclaimed.
1846 – The colony of North Australia was proclaimed by Letters Patent on 17 February. This was all of New South Wales north of 26° S. Although revoked in December 1846, the colony did formally exist.

text: Economy and trade[edit]

text:
The colonies relied heavily on imports from England for survival.
The official currency of the colonies was the British pound, but the unofficial currency and most readily accepted trade good was rum. During this period Australian businessmen began to prosper. For example, the partnership of Berry and Wollstonecraft made enormous profits by means of land grants, convict labour, and exporting native cedar back to England.

text: Religion, education, and culture[edit]

text: Since time immemorial in Australia, indigenous people had performed the rites and rituals of the animist religion of the Dreamtime. The permanent presence of Christianity in Australia however, came with the arrival of the First Fleet of British convict ships at Sydney in 1788. As a British colony, the predominant Christian denomination was the Church of England, but one-tenth of all the convicts who came to Australia on the First Fleet were Catholic, and at least half of them were born in Ireland.[37]

text: A small proportion of British marines were also Catholic. Some of the Irish convicts had been Transported to Australia for political crimes or social rebellion in Ireland, so the authorities were suspicious of the minority religion for the first three decades of settlement.[38] It was therefore the crew of the French explorer La Pérouse who conducted the first Catholic ceremony on Australian soil in 1788 – the burial of Father Louis Receveur, a Franciscan monk, who died while the ships were at anchor at Botany Bay, while on a mission to explore the Pacific.[39]

text: In early colonial times, Church of England clergy worked closely with the governors. Richard Johnson, Anglican chaplain to the First Fleet, was charged by the governor, Arthur Phillip, with improving "public morality" in the colony, but he was also heavily involved in health and education.[40] The Reverend Samuel Marsden (1765–1838) had magisterial duties, and so was equated with the authorities by the convicts. He became known as the 'floging parson' for the severity of his punishments[41]

text: Catholic convicts were compelled to attend Church of England services and their children and orphans were raised by the authorities as Protestant.[42] The first Catholic priest colonists arrived in Australia as convicts in 1800 – James Harold, James Dixon, and Peter O'Neill, who had been convicted for 'complicity' in the Irish 1798 Rebellion. Fr Dixon was conditionally emancipated and permitted to celebrate Mass. On 15 May 1803, in vestments made from curtains and with a chalice made of tin he conducted the first Catholic Mass in New South Wales.[42]

text: The Irish led Castle Hill Rebellion of 1804 alarmed the British authorities and Dixon's permission to celebrate Mass was revoked. Fr Jeremiah Flynn, an Irish Cistercian, was appointed as Prefect Apostolic of New Holland, and set out from Britain for the colony, uninvited. Watched by authorities, Flynn secretly performed priestly duties before being arrested and deported to London. Reaction to the affair in Britain led to two further priests being allowed to travel to the Colony in 1820 – John Joseph Therry and Philip Connolly.[38] The foundation stone for the first St Mary's Cathedral, Sydney was laid on 29 October 1821 by Governor Lachlan Macquarie.

text: The absence of a Catholic mission in Australia before 1818 reflected the legal disabilities of Catholics in Britain and the difficult position of Ireland within the British Empire. The government therefore endorsed the English Benedictines to lead the early Church in the Colony.[43] The Church of England lost its legal privileges in the Colony of New South Wales by the Church Act of 1836. Drafted by the reformist attorney-general John Plunkett, the Act established legal equality for Anglicans, Catholics and Presbyterians and was later extended to Methodists.[44] Catholic missionary William Ullathorne criticised the convict system, publishing a pamphlet, The Horrors of Transportation Briefly Unfolded to the People, in Britain in 1837.[45] Laywoman Caroline Chisolm did ecumenical work to alleviate the suffering of female migrants.

text: Initially, education was informal, primarily occurring in the home.[citation needed] At the instigation of the then British Prime Minister, the Duke of Wellington, and with the patronage of King William IV, Australia's oldest surviving independent school, The King's School, Parramatta, was founded in 1831 as part of an effort to establish grammar schools in the colony.[46] By 1833, there were around ten Catholic schools in the Australian colonies.[38] Today one in five Australian students attend Catholic schools.[47]

text: Sydney's first Catholic Bishop, John Bede Polding requested a community of nuns be sent to the colony and five Irish Sisters of Charity arrived in 1838 to set about pastoral care of convict women and work in schools and hospitals before going on to found their own schools and hospitals.[48] At Polding's request, the Christian Brothers arrived in Sydney in 1843 to assist in schools. Establishing themselves first at Sevenhill, in South Australia in 1848, the Jesuits were the first religious order of priests to enter and establish houses in South Australia, Victoria, Queensland and the Northern Territory – where they established schools and missions.

text: Some Australian folksongs date to this period.

text: Among the first true works of Australian literature produced over this period was the accounts of the settlement of Sydney by Watkin Tench, a captain of the marines on the First Fleet to arrive in 1788. In 1819, poet, explorer, journalist and politician William Wentworth published the first book written by an Australian: A Statistical, Historical, and Political Description of the Colony of New South Wales and Its Dependent Settlements in Van Diemen's Land, With a Particular Enumeration of the Advantages Which These Colonies Offer for Emigration and Their Superiority in Many Respects Over Those Possessed by the United States of America,[49] in which he advocated an elected assembly for New South Wales, trial by jury and settlement of Australia by free emigrants rather than convicts. In 1838 The Guardian: a tale by Anna Maria Bunn was published in Sydney. It was the first Australian novel printed and published in mainland Australia and the first Australian novel written by a woman. It is a Gothic romance.[50]

text: European traditions of Australian theatre also came with the First Fleet, with the first production being performed in 1789 by convicts : The Recruiting Officer by George Farquhar.[51] Two centuries later, the extraordinary circumstances of the foundations of Australian theatre were recounted in Our Country's Good by Timberlake Wertenbaker: the participants were prisoners watched by sadistic guards and the leading lady was under threat of the death penalty. The play is based on Thomas Keneally's novel The Playmaker.[51] The Theatre Royal, Hobart, opened in 1837 and it remains the oldest theatre in Australia.[52] The Melbourne Athenaeum is one of the oldest public institutions in Australia, founded in 1839 and it served as library, school of arts and dance hall (and later became Australia's first cinema, screening The Story of the Kelly Gang, the world's first feature film in 1906).[53] The Queen's Theatre, Adelaide opened with Shakespeare in 1841 and is today the oldest theatre on the mainland.[54]

text: Aboriginal resistance[edit]

text: Aboriginal reactions to the sudden arrival of British settlers were varied, but often hostile when the presence of the colonisers led to competition over resources, and to the occupation by the British of Aboriginal lands. European diseases decimated Aboriginal populations, and the occupation or destruction of lands and food resources led to starvation. By contrast with New Zealand, where the Treaty of Waitangi was seen to legitimise British settlement, no treaty was signed with Aborigines, who never authorised British colonisation.

text: According to the historian Geoffrey Blainey, in Australia during the colonial period:

text: "In a thousand isolated places there were occasional shootings and spearings. Even worse, smallpox, measles, influenza and other new diseases swept from one Aboriginal camp to another... The main conqueror of Aborigines was to be disease and its ally, demoralisation".[55]

text: Since the 1980s, the use of the word "invasion" to describe the British colonisation of Australia has been highly controversial. According to Australian Henry Reynolds however, government officials and ordinary settlers in the eighteenth and nineteenth centuries frequently used words such as "invasion" and "warfare" to describe their presence and relations with Indigenous Australians. In his book The Other Side of the Frontier,[56] Reynolds described in detail armed resistance by Aboriginal people to white encroachments by means of guerilla warfare, beginning in the eighteenth century and continuing into the early twentieth.

text: In the early years of colonisation, David Collins, the senior legal officer in the Sydney settlement, wrote of local Aborigines:

text: While they entertain the idea of our having dispossessed them of their residences, they must always consider us as enemies; and upon this principle they [have] made a point of attacking the white people whenever opportunity and safety concurred.[57]

text: In 1847, Western Australian barrister E.W. Landor stated: "We have seized upon the country, and shot down the inhabitants, until the survivors have found it expedient to submit to our rule. We have acted as Julius Caesar did when he took possession of Britain."[58] In most cases, Reynolds says, Aborigines initially resisted British presence. In a letter to the Launceston Advertiser in 1831, a settler wrote:

text: We are at war with them: they look upon us as enemies – as invaders – as oppressors and persecutors – they resist our invasion. They have never been subdued, therefore they are not rebellious subjects, but an injured nation, defending in their own way, their rightful possessions which have been torn from them by force.[59]

text: Reynolds quotes numerous writings by settlers who, in the first half of the nineteenth century, described themselves as living in fear and even in terror due to attacks by Aborigines determined to kill them or drive them off their lands. He argues that Aboriginal resistance was, in some cases at least, temporarily effective; the Aboriginal killings of men, sheep and cattle, and burning of white homes and crops, drove some settlers to ruin. Aboriginal resistance continued well beyond the middle of the nineteenth century, and in 1881 the editor of The Queenslander wrote:

text: During the last four or five years the human life and property destroyed by the aborigines in the North total up to a serious amount. [...] [S]ettlement on the land, and the development of the mineral and other resources on the country, have been in a great degree prohibited by the hostility of the blacks, which still continues with undiminished spirit.[60]

text: Reynolds argues that continuous Aboriginal resistance for well over a century belies the "myth" of peaceful settlement in Australia. Settlers in turn often reacted to Aboriginal resistance with great violence, resulting in numerous indiscriminate massacres by whites of Aboriginal men, women and children.[61] Among the most famous massacres of the early nineteenth century were the Pinjarra massacre and the Myall Creek massacre.

text: Famous Aborigines who resisted British colonisation in the eighteenth and early nineteenth centuries include Pemulwuy and Yagan. In Tasmania, the "Black War" was fought in the first half of the nineteenth century.

text: Representations in literature and film[edit]

text:
Marcus Clarke's 1874 novel, For the Term of his Natural Life, and the 1983 television adaptation of the novel.
Eleanor Dark's 1947 Timeless Land trilogy, which spans the colonisation from 1788–1811. The 1980s television drama, The Timeless Land, was based on this trilogy.
D. Manning Richards. Destiny in Sydney: An epic novel of convicts, Aborigines, and Chinese embroiled in the birth of Sydney, Australia. First book in Sydney series. Washington DC: Aries Books, 2012. ISBN 978-0-9845410-0-3


document_title: Natural selection

question_text: who developed the idea of evolution by natural selection

root

text: Natural selection

text: Historical development[edit]

text: Pre-Darwinian theories[edit]

text: Several philosophers of the classical era, including Empedocles[1] and his intellectual successor, the Roman poet Lucretius,[2] 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 persist. Empedocles' idea that organisms arose entirely by the incidental workings of causes such as heat and cold was criticised by Aristotle in Book II of Physics.[3] He posited natural teleology in its place, and believed that form was achieved for a purpose, citing the regularity of heredity in species as proof.[4][5] Nevertheless, he accepted in his biology that new types of animals, monstrosities (τερας), can occur in very rare instances (Generation of Animals, Book IV).[6] As quoted in Darwin's 1872 edition of The Origin of Species, Aristotle considered whether different forms (e.g., of teeth) might have appeared accidentally, but only the useful forms survived:

tag: p

text: 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 the 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.

text: But Aristotle rejected this possibility in the next paragraph, making clear that he is talking about the development of animals as embryos with the phrase "either invariably or normally come about", not the origin of species:

text: ... 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.

text: The struggle for existence was later described by the Islamic writer Al-Jahiz in the 9th century.[9][10][11]

text: The classical arguments were reintroduced in the 18th century by Pierre Louis Maupertuis[12] and others, including Darwin's grandfather, Erasmus Darwin.

text: 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 ideals (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.[13]

text: The early 19th-century zoologist 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.[14] This theory, Lamarckism, was an influence on the Soviet biologist Trofim Lysenko's antagonism to mainstream genetic theory as late as the mid 20th century.[15]

text: Between 1835 and 1837, the zoologist Edward Blyth worked on the area of variation, artificial selection, and how a similar process occurs in nature. Darwin acknowledged Blyth's ideas in the first chapter on variation of On the Origin of Species.[16]

text: Darwin's theory[edit]

text: 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".[17] 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 and that variation is heritable, there will be an inevitable selection of individuals with the most advantageous variations. If the variations are heritable, then differential reproductive success leads to a progressive evolution of particular populations of a species, and populations that evolve to be sufficiently different eventually become different species.[18][19]

text: Darwin's ideas were inspired by the observations that he had made on the second voyage of HMS Beagle (1831–1836), and by the work of a political economist, Thomas Robert Malthus, who, in An Essay on the Principle of Population (1798), 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".[20] When Darwin read Malthus in 1838 he was already primed by his work as a naturalist to appreciate the "struggle for existence" in nature. 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."[21] Darwin wrote:

text: 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.

text: Once he had his theory, Darwin was meticulous about gathering and refining evidence before making his idea public. He was in the process of writing his "big book" to present his research 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 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 of London announcing co-discovery of the principle in July 1858.[23] 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—like 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.[24]

text: 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 Natural Selection has been the main but not exclusive means of modification."[25] In a letter to Charles Lyell in September 1860, Darwin regretted the use of the term "Natural Selection", preferring the term "Natural Preservation".[26]

text: 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,[27] 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,[28] a response that has been characterised as the single most significant impediment to the idea's acceptance.[29] However, some thinkers enthusiastically embraced natural selection; after reading Darwin, Herbert Spencer introduced the phrase survival of the fittest, which became a popular summary of the theory.[30][31] 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."[32] 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.[33]

text: The modern synthesis[edit]

text: Natural selection relies crucially on the idea of heredity, but developed before the basic concepts of genetics. Although the Moravian monk Gregor Mendel, the father of modern genetics, was a contemporary of Darwin's, his work lay in obscurity, only being rediscovered in 1900.[34] With the early 20th century integration of evolution with Mendel's laws of inheritance, the so-called modern synthesis, scientists generally came to accept natural selection.[35][36] The synthesis grew from advances in different fields. Ronald Fisher developed the required mathematical language and wrote The Genetical Theory of Natural Selection (1930).[37] J. B. S. Haldane introduced the concept of the "cost" of natural selection.[38][39] Sewall Wright elucidated the nature of selection and adaptation.[40] In his book Genetics and the Origin of Species (1937), Theodosius Dobzhansky established the idea that mutation, once seen as a rival to selection, actually supplied the raw material for natural selection by creating genetic diversity.[41][42]

text: A second synthesis[edit]

text: Ernst Mayr recognised the key importance of reproductive isolation for speciation in his Systematics and the Origin of Species (1942).[44] W. D. Hamilton conceived of kin selection in 1964.[45][46] This synthesis cemented natural selection as the foundation of evolutionary theory, where it remains today. A second synthesis was brought about at the end of the 20th century by advances in molecular genetics, creating the field of evolutionary developmental biology ("evo-devo"), which seeks to explain the evolution of form in terms of the genetic regulatory programs which control the development of the embryo at molecular level. Natural selection is here understood to act on embryonic development to change the morphology of the adult body.[47][48][49][50]

text: Terminology[edit]

text: The term natural selection is most often defined to operate on heritable traits, because these directly participate in evolution. However, natural selection is "blind" in the sense that changes in phenotype can give a reproductive advantage regardless of whether or not the trait is heritable. Following Darwin's primary usage, the term is used to refer both to the evolutionary consequence of blind selection and to its mechanisms.[27][37][51][52] It is sometimes helpful to explicitly distinguish between selection's mechanisms and its effects; when this distinction is important, scientists define "(phenotypic) 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.[53][54][55] Traits that cause greater reproductive success of an organism are said to be selected for, while those that reduce success are selected against.[56]

text: Mechanism[edit]

text: Heritable variation, differential reproduction[edit]

text: Natural variation occurs among the individuals of any population of organisms. Some differences may improve an individual's chances of surviving and reproducing such that its lifetime reproductive rate is increased, which means that it leaves more offspring. If the traits that give these individuals a reproductive advantage are also heritable, that is, passed from parent to offspring, then there will be differential reproduction, that is, a slightly higher proportion of fast rabbits or efficient algae in the next generation. Even if the reproductive advantage is very slight, over many generations any advantageous heritable trait becomes dominant in the population. In this way the natural environment of an organism "selects for" traits that confer a reproductive advantage, causing evolutionary change, as Darwin described.[57] This gives the appearance of purpose, but in natural selection there is no intentional choice. Artificial selection is purposive where natural selection is not, though biologists often use teleological language to describe it.[58]

text: The peppered moth exists in both light and dark colours in Great Britain, but during the industrial revolution, many of the trees on which the moths rested became blackened by soot, giving the dark-coloured moths an advantage in hiding from predators. This gave dark-coloured moths a better chance of surviving to produce dark-coloured 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.[59]

text: Fitness[edit]

text: 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", but 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 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.[60]

text: Competition[edit]

text: In biology, competition is an interaction between organisms in which the fitness of one is lowered by the presence of another. This may be because both rely on a limited supply of a resource such as food, water, or territory.[61] Competition may be within or between species, and may be direct or indirect.[62] Species less suited to compete should in theory either adapt or die out, since competition plays a powerful role in natural selection, but according to the "room to roam" theory it may be less important than expansion among larger clades.[62][63]

text: Competition is modelled by r/K selection theory, which is based on Robert MacArthur and E. O. Wilson's work on island biogeography.[64] In this theory, selective pressures drive evolution in one of two stereotyped directions: r- or K-selection.[65] These terms, r and K, can be illustrated in a logistic model of population dynamics:[66]

text: where r is the growth rate of the population (N), and K is the carrying capacity of its local environmental setting. Typically, r-selected species exploit empty niches, and produce many offspring, each with a relatively low probability of surviving to adulthood. In contrast, K-selected species are strong competitors in crowded niches, and invest more heavily in much fewer offspring, each with a relatively high probability of surviving to adulthood.[66]

text: Types of selection[edit]

text: Natural selection can act on any heritable phenotypic trait,[67] 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.[68][69] 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.[57]

text: Selection can be classified in several different ways, such as by its effect on a trait, on genetic diversity, by the life cycle stage where it acts, by the unit of selection, or by the resource being competed for.

text: Selection has different effects on traits. Stabilizing selection acts to hold a trait at a stable optimum, and in the simplest case all deviations from this optimum are selectively disadvantageous. Directional selection favours extreme values of a trait. The uncommon disruptive selection also acts during transition periods when the current mode is sub-optimal, but alters the trait in more than one direction. In particular, if the trait is quantitative and univariate then both higher and lower trait levels are favoured. Disruptive selection can be a precursor to speciation.[57]

text: Alternatively, selection can be divided according to its effect on genetic diversity. Purifying or negative selection acts to remove genetic variation from the population (and is opposed by de novo mutation, which introduces new variation.[70][71] In contrast, balancing selection acts to maintain genetic variation in a population, even in the absence of de novo mutation, by negative frequency-dependent selection. One mechanism for this is heterozygote advantage, where individuals with two different alleles have a selective advantage over individuals with just one allele. The polymorphism at the human ABO blood group locus has been explained in this way.[72]

text: Another option is to classify selection by the life cycle stage at which it acts. Some biologists recognise just two types: viability (or survival) selection, which acts to increase an organism's probability of survival, and fecundity (or fertility or reproductive) selection, which acts to increase the rate of reproduction, given survival. Others split the life cycle into further components of selection. Thus viability and survival selection may be defined separately and respectively as acting to improve the probability of survival before and after reproductive age is reached, while fecundity selection may be split into additional sub-components including sexual selection, gametic selection, acting on gamete survival, and compatibility selection, acting on zygote formation.[73]

text: Selection can also be classified by the level or unit of selection. Individual selection acts on the individual, in the sense that adaptations are "for" the benefit of the individual, and result from selection among individuals. Gene selection acts directly at the level of the gene. In kin selection and intragenomic conflict, gene-level selection provides a more apt explanation of the underlying process. Group selection, if it occurs, acts on groups of organisms, on the assumption that groups replicate and mutate in an analogous way to genes and individuals. There is an ongoing debate over the degree to which group selection occurs in nature.[74]

text: Finally, selection can be classified according to the resource being competed for. Sexual selection results from competition for mates. Sexual selection typically proceeds via fecundity selection, sometimes at the expense of viability. Ecological selection is natural selection via any means other than sexual selection, such as kin selection, competition, and infanticide. Following Darwin, natural selection is sometimes defined as ecological selection, in which case sexual selection is considered a separate mechanism.[75]

text: Sexual selection[edit]

text: Sexual selection as first articulated by Darwin (using the example of the peacock's tail)[76] refers specifically to competition for mates,[78] which can be intrasexual, between individuals of the same sex, that is male–male competition, or intersexual, where one gender chooses mates, most often with males displaying and females choosing.[79] However, in some species, mate choice is primarily by males, as in some fishes of the family Syngnathidae.[80][81]

text: Phenotypic traits can be displayed in one sex and desired in the other sex, causing a positive feedback loop called a Fisherian runaway, for example, the extravagant plumage of some male birds such as the peacock.[77] An alternate theory proposed by the same Ronald Fisher in 1930 is the sexy son hypothesis, that mothers want promiscuous sons to give them large numbers of grandchildren and so choose promiscuous fathers for their children. 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.[79]

text: Natural selection in action[edit]

text: Natural selection is seen in action in the development of antibiotic resistance in microorganisms. Since the discovery of penicillin in 1928, antibiotics have been used to fight bacterial diseases. The widespread misuse of antibiotics has selected for 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.[82] 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.[83] This is an evolutionary arms race, in which bacteria develop strains less susceptible to antibiotics, while medical researchers attempt 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[84][85]

text: Evolution by means of natural selection[edit]

text: 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, genetic 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 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, many mutations in non-coding DNA have deleterious effects.[86][87] Although both mutation rates and average fitness effects of mutations are dependent on the organism, a majority of mutations in humans are slightly deleterious.[88]

text: Some mutations occur in "toolkit" or regulatory genes. Changes in these often 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 embryos. Some nonlethal regulatory mutations occur in HOX genes in humans, which can result in a cervical rib[89] or polydactyly, an increase in the number of fingers or toes.[90] When such mutations result in a higher fitness, natural selection favours these phenotypes and the novel trait spreads 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 becomes 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.[91]

text: Speciation[edit]

text: Speciation requires a degree of reproductive isolation—that is, a reduction in gene flow. However, it is intrinsic to the concept of a species that hybrids are selected against, opposing the evolution of reproductive isolation, a problem that was recognised by Darwin. The problem does not occur in allopatric speciation with geographically separated populations, which can diverge with different sets of mutations. E. B. Poulton realized in 1903 that reproductive isolation could evolve through divergence, if each lineage acquired a different, incompatible allele of the same gene. Selection against the heterozygote would then directly create reproductive isolation, leading to the Bateson–Dobzhansky–Muller model, further elaborated by H. Allen Orr[92] and Sergey Gavrilets.[93] With reinforcement, however, natural selection can favor an increase in pre-zygotic isolation, influencing the process of speciation directly.[94]

text: Genetic basis[edit]

text: Genotype and phenotype[edit]

text: 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. 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 differences in phenotype. An example is the ABO blood type antigens in humans, where three alleles govern the phenotype.[95]

text: 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.[96]

text: Directionality of selection[edit]

text: When some component of a trait is heritable, selection alters 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: directional, stabilizing, and purifying selection.[97] Directional selection occurs when an allele has a greater fitness than others, so that it increases in frequency, gaining an increasing share in the population. This process can continue until the allele is fixed and the entire population shares the fitter phenotype.[98] Far more common is stabilizing selection, which lowers the frequency of alleles that have a deleterious effect on the phenotype – that is, produce organisms of lower fitness. This process can continue until the allele is eliminated from the population. Purifying selection conserves functional genetic features, such as protein-coding genes or regulatory sequences, over time by selective pressure against deleterious variants.[99]

text: Some forms of balancing selection do not result in fixation, but maintain an allele at intermediate frequencies in a population. This can occur in diploid species (with pairs of chromosomes) when heterozygous individuals (with just one copy of the allele) have a higher fitness than homozygous individuals (with two copies). This is called heterozygote advantage or over-dominance, of which the best-known example is the resistance to malaria in humans heterozygous for sickle-cell anaemia. Maintenance of allelic variation can also occur through disruptive or diversifying selection, which favours genotypes that depart from the average in either direction (that is, the opposite of over-dominance), 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.[100][101]

text: Selection, genetic variation, and drift[edit]

text: A portion of all genetic variation is functionally neutral, producing no phenotypic effect or significant difference in fitness. Motoo Kimura's neutral theory of molecular evolution by genetic drift proposes that this variation accounts for a large fraction of observed genetic diversity.[102] Neutral events can radically reduce genetic variation through population bottlenecks.[103] which among other things can cause the founder effect in initially small new populations.[104] 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 is higher than at sites where variation does influence fitness.[97] However, after a period with no new mutations, the genetic variation at these sites is eliminated due to genetic drift. Natural selection reduces genetic variation by eliminating maladapted individuals, and consequently 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 unfavourable the mutation proves to be.[105]

text: Genetic linkage occurs when the loci of two alleles are in close proximity on a chromosome. During the formation of gametes, recombination reshuffles the alleles. The chance that such a reshuffle occurs between two alleles is inversely related to the distance between them. 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, closely linked alleles can also become more common by "genetic hitchhiking", whether they are neutral or even slightly deleterious. A strong selective sweep results in a region of the genome where the positively selected haplotype (the allele and its neighbours) are in essence the only ones that exist in the population. Selective sweeps can be detected by measuring linkage disequilibrium, or whether a given haplotype is overrepresented in the population. Since a selective sweep also results in selection of neighbouring alleles, the presence of a block of strong linkage disequilibrium might indicate a 'recent' selective sweep near the centre of the block.[106]

text: Background selection is the opposite of a selective sweep. If a specific site experiences strong and persistent purifying selection, linked variation tends 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.[107]

text: Impact[edit]

text: Darwin's ideas, along with those of Adam Smith and Karl Marx, had a profound influence on 19th century thought, including his radical claim 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.[108] This inspired some of Darwin's most ardent supporters—and provoked the strongest opposition. Natural selection had the power, according to Stephen Jay Gould, to "dethrone some of the deepest and most traditional comforts of Western thought", such as the belief that humans have a special place in the world.[109]

text: In the words of the philosopher Daniel Dennett, "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.[110] Thus, in the last decades, the concept of natural selection has spread from evolutionary biology to other disciplines, including evolutionary computation, quantum Darwinism, evolutionary economics, evolutionary epistemology, evolutionary psychology, and cosmological natural selection. This unlimited applicability has been called universal Darwinism.[111]

text: Origin of life[edit]

text: How life originated from inorganic matter remains an unresolved problem in biology. One prominent hypothesis is that life first appeared in the form of short self-replicating RNA polymers.[112] On this view, life may have come into existence when RNA chains first experienced the basic conditions, as conceived by Charles Darwin, for natural selection to operate. These conditions are: heritability, variation of type, and competition for limited resources. The fitness of an early RNA replicator would likely have been a function of adaptive capacities that were intrinsic (i.e., determined by the nucleotide sequence) and the availability of resources.[113][114] The three primary adaptive capacities could logically have been: (1) the capacity to replicate with moderate fidelity (giving rise to both heritability and variation of type), (2) the capacity to avoid decay, and (3) the capacity to acquire and process resources.[113][114] These capacities would have been determined initially by the folded configurations (including those configurations with ribozyme activity) of the RNA replicators that, in turn, would have been encoded in their individual nucleotide sequences.[115]

text: Cell and molecular biology[edit]

text: In 1881, the embryologist Wilhelm Roux published Der Kampf der Theile 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.[116] In recent years, a modern version of this theory has been proposed by Jean-Jacques Kupiec. According to this cellular Darwinism, random variation at the molecular level generates diversity in cell types whereas cell interactions impose a characteristic order on the developing embryo.[117]

text: Social and psychological theory[edit]

text: 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, and especially on his countrymen, 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."[118] Herbert Spencer and the eugenics advocate Francis Galton's interpretation of natural selection as necessarily progressive, leading to supposed advances in intelligence and civilisation, became a justification for colonialism, eugenics, and social Darwinism. For example, in 1940, Konrad Lorenz, in writings that he subsequently disowned, 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."[119] Others have developed ideas that human societies and culture evolve by mechanisms analogous to those that apply to evolution of species.[120]

text: More recently, work among anthropologists and psychologists has led to the development of sociobiology and later of evolutionary psychology, a field that attempts to explain features of human psychology in terms of adaptation to the ancestral environment. The most prominent example of evolutionary psychology, notably advanced in the early work of Noam Chomsky and later by Steven Pinker, is the hypothesis that the human brain has adapted to acquire the grammatical rules of natural language.[121] Other aspects of human behaviour and social structures, from specific cultural norms such as incest avoidance to broader patterns such as gender roles, have been hypothesised to have similar origins as adaptations to the early environment in which modern humans evolved. By analogy to the action of 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 in 1976[122] and subsequently expanded upon by philosophers such as Daniel Dennett as explanations for complex cultural activities, including human consciousness.[123]

text: Information and systems theory[edit]

text: In 1922, Alfred J. 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,[124][125] a concept later developed by Howard T. Odum as the maximum power principle in thermodynamics, whereby evolutionary systems with selective advantage maximise the rate of useful energy transformation.[126]

text: 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.[127] For example, a class of heuristic optimisation algorithms known as genetic algorithms, pioneered by John Henry Holland in the 1970s and expanded upon by David E. Goldberg,[128] identify optimal solutions by simulated reproduction and mutation of a population of solutions defined by an initial probability distribution.[129] Such algorithms are particularly useful when applied to problems whose energy landscape is very rough or has many local minima.[130]