Early evolutionary thinking Wallace began his career as a travelling naturalist already believing in the
transmutation of species. The concept had been advocated by
Jean-Baptiste Lamarck,
Geoffroy Saint-Hilaire,
Erasmus Darwin, and
Robert Grant, among others. It was widely discussed, but not generally accepted by leading naturalists, and was considered to have
radical, even revolutionary connotations. Prominent anatomists and geologists such as
Georges Cuvier,
Richard Owen,
Adam Sedgwick, and Lyell attacked transmutation vigorously. It has been suggested that Wallace accepted the idea of the transmutation of species in part because he was always inclined to favour radical ideas in politics, religion and science, and because he was unusually open to marginal, even fringe, ideas in science. Wallace was profoundly influenced by
Robert Chambers's
Vestiges of the Natural History of Creation, a controversial work of popular science published anonymously in 1844. It advocated an evolutionary origin for the
Solar System, the Earth, and living things. Wallace wrote to Henry Bates in 1845 describing it as "an ingenious hypothesis strongly supported by some striking facts and analogies, but which remains to be proven by ... more research". In 1847, he wrote to Bates that he would "like to take some one family [of beetles] to study thoroughly, ... with a view to the theory of the origin of species." Wallace planned fieldwork to test the evolutionary hypothesis that closely related species should inhabit neighbouring territories. During his work in the
Amazon basin, he came to realise that geographical barriers—such as the Amazon and its major tributaries—often separated the ranges of closely allied species. He included these observations in his 1853 paper "On the Monkeys of the Amazon". Near the end of the paper he asked the question, "Are very closely allied species ever separated by a wide interval of country?".
Sarawak Law Paper For sometime Wallace had been compiling notes for an argument against the concept of new species being introduced through
progressive creationism found in
Charles Lyell's
Principles of Geology. This was Lyell's rebuttal to transmutational ideas. When Wallace read an 1854 paper written by
Edward Forbes that proposed an even more quasi-mystical form of progressive creationism he decided to write a response. In February 1855, while working in
Sarawak on the island of
Borneo, Wallace wrote "On the Law which has Regulated the Introduction of New Species". The paper was published in the
Annals and Magazine of Natural History in September 1855. In this paper, he discussed observations of the geographic and geologic distribution of both living and fossil species, a field that became biogeography. His observation (his notebook shows it was inspired by
Pictet's Palaeontology) that "Every species has come into existence coincident both in space and time with a closely allied species" has come to be known as the "Sarawak Law", answering his own question in his paper on the monkeys of the Amazon basin. Although it does not mention possible mechanisms for evolution or even explicitly discuss transmutation, it did say that each new species was a small variation on an already existing species and that this process resulted in a branching tree of relationships between species that should be the basis of taxonomic classification. This paper foreshadowed the momentous paper he would write three years later. The paper challenged Lyell's belief that species were immutable. Although Darwin had written to him in 1842 expressing support for transmutation, Lyell had continued to be strongly opposed to the idea. Around the start of 1856, he told Darwin about Wallace's paper, as did
Edward Blyth who thought it "Good! Upon the whole! ... Wallace has, I think put the matter well; and according to his theory the various domestic races of animals have been fairly developed into
species." Despite this hint, Darwin mistook Wallace's conclusion for the progressive creationism of the time, writing that it was "nothing very new ... Uses my simile of tree [but] it seems all creation with him." Lyell was more impressed, and opened a notebook on species in which he grappled with the consequences, particularly for human ancestry. Darwin had already shown his theory to their mutual friend
Joseph Hooker and now, for the first time spelt out the full details of natural selection to Lyell. Although Lyell could not agree, he urged Darwin to publish to establish priority. Darwin demurred at first, but began writing up a
species sketch of his continuing work in May 1856.
Natural selection and Darwin By February 1858, Wallace had been convinced by his biogeographical research in the Malay Archipelago that evolution was real. He later wrote in his autobiography that the problem was of how species change from one well-marked form to another. He stated that it was while he was in bed with a fever that he thought about Malthus's idea of positive checks on human population, and had the idea of natural selection. His autobiography says that he was on the island of
Ternate at the time; but the evidence of his journal suggests that he was in fact on the island of
Gilolo. From 1858 to 1861, he rented a house on Ternate from the Dutchman
Maarten Dirk van Renesse van Duivenbode, which he used as a base for expeditions to other islands such as Gilolo. Wallace describes how he discovered natural selection as follows: was issued by the
Linnean Society on the 50th anniversary of the reading of Darwin and Wallace's papers on
natural selection. Wallace received the only gold example.|alt=photograph of the Darwin-Wallace medal Wallace had once briefly met Darwin, and was one of the correspondents whose observations Darwin used to support his own theories. Although Wallace's first letter to Darwin has been lost, Wallace carefully kept the letters he received. In the first letter, dated 1 May 1857, Darwin commented that Wallace's letter of 10 October which he had recently received, as well as Wallace's paper "On the Law which has regulated the Introduction of New Species" of 1855, showed that they thought alike, with similar conclusions, and said that he was preparing his own work for publication in about two years time. The second letter, dated 22 December 1857, said how glad he was that Wallace was theorising about distribution, adding that "without speculation there is no good and original observation" but commented that "I believe I go much further than you". Wallace believed this and sent Darwin his February 1858 essay, "
On the Tendency of Varieties to Depart Indefinitely From the Original Type", asking Darwin to review it and pass it to
Charles Lyell if he thought it worthwhile. Although Wallace had sent several articles for journal publication during his travels through the Malay archipelago, the Ternate essay was in a private letter. Darwin received the essay on 18 June 1858. Although the essay did not use Darwin's term "natural selection", it did outline the mechanics of an evolutionary divergence of species from similar ones due to environmental pressures. In this sense, it was very similar to the theory that Darwin had worked on for 20 years, but had yet to publish. Darwin sent the manuscript to Charles Lyell with a letter saying "he could not have made a better short abstract! Even his terms now stand as heads of my chapters ... he does not say he wishes me to publish, but I shall, of course, at once write and offer to send to any journal." Distraught about the illness of his baby son, Darwin put the problem to Charles Lyell and
Joseph Hooker, who decided to publish the essay in a joint presentation together with unpublished writings which highlighted Darwin's priority. Wallace's essay was presented to the
Linnean Society of London on 1 July 1858, along with excerpts from an essay which Darwin had disclosed privately to Hooker in 1847 and a letter Darwin had written to
Asa Gray in 1857. Communication with Wallace in the far-off Malay Archipelago involved months of delay, so he was not part of this rapid publication. Wallace accepted the arrangement after the fact, happy that he had been included at all, and never expressed bitterness in public or in private. Darwin's social and scientific status was far greater than Wallace's, and it was unlikely that, without Darwin, Wallace's views on evolution would have been taken seriously. Lyell and Hooker's arrangement relegated Wallace to the position of co-discoverer, and he was not the social equal of Darwin or the other prominent British natural scientists. All the same, the joint reading of their papers on natural selection associated Wallace with the more famous Darwin. This, combined with Darwin's (as well as Hooker's and Lyell's) advocacy on his behalf, would give Wallace greater access to the highest levels of the scientific community. The reaction to the reading was muted, with the president of the Linnean Society remarking in May 1859 that the year had not been marked by any striking discoveries; but, with Darwin's publication of
On the Origin of Species later in 1859, its significance became apparent. When Wallace returned to the UK, he met Darwin. Although some of Wallace's opinions in the ensuing years would test Darwin's patience, they remained on friendly terms for the rest of Darwin's life. Over the years, a few people have questioned this version of events. In the early 1980s, two books, one by
Arnold Brackman and another by
John Langdon Brooks, suggested not only that there had been a conspiracy to rob Wallace of his proper credit, but that Darwin had actually stolen a key idea from Wallace to finish his own theory. These claims have been examined and found unconvincing by a number of scholars. Shipping schedules show that, contrary to these accusations, Wallace's letter could not have been delivered earlier than the date shown in Darwin's letter to Lyell.
Defence of Darwin and his ideas After Wallace returned to England in 1862, he became one of the staunchest defenders of Darwin's
On the Origin of Species. In an incident in 1863 that particularly pleased Darwin, Wallace published the short paper "Remarks on the Rev. S. Haughton's Paper on the Bee's Cell, And on the Origin of Species". This rebutted a paper by a professor of geology at the University of Dublin that had sharply criticised Darwin's comments in the
Origin on how hexagonal honey bee cells could have evolved through natural selection. An even longer defence was an 1867 article in the
Quarterly Journal of Science, "Creation by Law". It reviewed
George Campbell, the 8th Duke of Argyll's book,
The Reign of Law, which aimed to refute natural selection. After an 1870 meeting of the
British Science Association, Wallace wrote to Darwin complaining that there were "no opponents left who know anything of natural history, so that there are none of the good discussions we used to have". Explorers in Madagascar had discovered the orchid
Angraecum sesquipedale, with a sixteen-inch nectary. Darwin predicted the existence of a moth with a proboscis long enough to pollinate it. In a footnote to "Creation by Law", Wallace wrote "That such a moth exists in Madagascar may be safely predicted; and naturalists who visit that island should search for it with as much confidence as astronomers
searched for the planet Neptune,—and they will be equally successful!"
Differences between Darwin and Wallace Historians of science have noted that, while Darwin considered the ideas in Wallace's paper to be essentially the same as his own, there were differences. Darwin emphasised competition between individuals of the same species to survive and reproduce, whereas Wallace emphasised environmental pressures on varieties and species forcing them to become adapted to their local conditions, leading populations in different locations to diverge. The historian of science
Peter J. Bowler has suggested that in the paper he mailed to Darwin, Wallace might have been discussing
group selection. Against this, Malcolm Kottler showed that Wallace was indeed discussing individual variation and selection. Others have noted that Wallace appeared to have envisioned natural selection as a kind of feedback mechanism that kept species and varieties adapted to their environment (now called 'stabilizing", as opposed to 'directional' selection). Bateson revisited the topic in his 1979 book
Mind and Nature: A Necessary Unity, and other scholars have continued to explore the connection between natural selection and
systems theory.
Warning colouration and sexual selection : a wasp (top) mimicked by a beetle in Wallace's 1889 book
Darwinism|alt=see caption
Warning colouration was one of Wallace's contributions to the evolutionary biology of
animal colouration. In 1867, Darwin wrote to Wallace about a problem in explaining how some caterpillars could have evolved conspicuous colour schemes. Darwin had come to believe that many conspicuous animal colour schemes were due to sexual selection, but he saw that this could not apply to caterpillars. Wallace responded that he and Bates had observed that many of the most spectacular butterflies had a peculiar odour and taste, and that he had been told by
John Jenner Weir that birds would not eat a certain kind of common white moth because they found it unpalatable. Since the moth was as conspicuous at dusk as a coloured caterpillar in daylight, it seemed likely that the conspicuous colours served as a warning to predators and thus could have evolved through natural selection. Darwin was impressed by the idea. At a later meeting of the Entomological Society, and in a letter published in
Field magazine, Wallace asked for any evidence anyone might have on the topic. In 1869, Weir published data from experiments and observations involving brightly coloured caterpillars that supported Wallace's idea. Wallace attributed less importance than Darwin to sexual selection. One area where he debated Darwin involved explaining sexual dimorphism in the colouration and nesting behaviour of many bird species where males are more brightly coloured than females. Darwin attributed this to sexual selection by females between competing males. In the 1860s Wallace wrote a series of papers that emphasized instead selective pressure from predation for drab colouration in females that spend a lot of time sitting on nests. In his 1878 book
Tropical Nature and Other Essays, he wrote extensively about the colouration of animals and plants, and proposed alternative explanations for a number of cases Darwin had attributed to sexual selection. He revisited the topic at length in his 1889 book
Darwinism. In 1890, he wrote a critical review in
Nature of his friend
Edward Bagnall Poulton's
The Colours of Animals which supported Darwin on sexual selection, attacking especially Poulton's claims on the "aesthetic preferences of the insect world".
Wallace effect In 1889, Wallace wrote the book
Darwinism, which explained and defended natural selection. In it, he proposed the hypothesis that natural selection could drive the reproductive isolation of two varieties by encouraging the development of barriers against hybridisation. Thus it might contribute to the development of new species. He suggested the following scenario: When two populations of a species had diverged beyond a certain point, each adapted to particular conditions, hybrid offspring would be less adapted than either parent form and so natural selection would tend to eliminate the hybrids. Furthermore, under such conditions, natural selection would favour the development of barriers to hybridisation, as individuals that avoided hybrid matings would tend to have more fit offspring, and thus contribute to the reproductive isolation of the two incipient species. This idea came to be known as the
Wallace effect, later called reinforcement. He was motivated to spell the idea out in detail in 1889 in response to a paper by
George Romanes that suggested that natural selection alone was insufficient to explain the separation of incipient species. Wallace had suggested to Darwin that natural selection could play a role in preventing hybridisation in private correspondence as early as 1868, but had not worked it out in detail. It continues to be a topic of research in evolutionary biology today, with both computer simulation and empirical results supporting its validity.
Evolution of humans to humans in Wallace's 1889 book
Darwinism shows a chimpanzee.|alt=illustration of a chimpanzee from one of Wallace's books In January of 1864 Wallace presented a paper, "On the Varieties of Man in the Malay Archipelago", to the
Ethnological Society of London. This paper summarized his ethnological observations during his travels in the East Indies. Wallace stated that the people in the archipelago could be divided into two groups. The first were members of what he called the
Malay race and the second were members of the
Papuan race. He described the physical, linguistic, and cultural differences between them, and he traced an east west dividing line that separated the 2 groups. He compared this with the famous dividing line he had discovered between Asian and Australian affiliated faunas, suggesting that the distribution of human races could be explained by the geographical history of a region the same way as the biogeographical distribution of species. A few weeks later he presented a follow up paper, "The Origin of Human Races and the Antiquity of Man Deduced from the Theory of 'Natural Selection to the
Anthropological Society of London , applying the theory of natural selection to humankind. Darwin had not yet publicly addressed the subject, although
Thomas Huxley had in ''
Evidence as to Man's Place in Nature''. Wallace explained the apparent stability of the human stock by pointing to the vast gap in cranial capacities between humans and the
great apes. Unlike some other Darwinists, including Darwin himself, he did not "regard modern primitives as almost filling the gap between man and ape". He saw the evolution of humans in two stages: achieving a bipedal posture that freed the hands to carry out the dictates of the brain, and the "recognition of the human brain as a totally new factor in the history of life". He argued that the natives die out due to an unequal struggle. Charles Lyell wrote to Wallace praising the paper and saying that it had made a major contribution to resolving the dispute between monogenists and polygenists.
Non material causes and teleology in evolution Prior to 1864, Wallace believed that humans were a result of only natural selection, however, by 1869 Wallace had changed his view since he felt that natural selection was an insufficient mechanism for the development of multiple features in man (e.g high capacity for rationality) and, according to historians of science, postulated that "higher intelligences guiding man's development were required."Shortly afterwards, Wallace became a
spiritualist. At about the same time, he began to maintain that natural selection could not account for mathematical, artistic, or musical genius, metaphysical musings, or wit and humour. He stated that something in "the unseen universe of Spirit" had interceded at least three times in history: the creation of life from inorganic matter; the introduction of consciousness in the higher animals; and the generation of the higher mental faculties in humankind. He believed that the
raison d'être of the universe was the development of the human spirit. While some historians have concluded that Wallace's belief that natural selection was insufficient to explain the development of consciousness and the higher functions of the human mind was directly caused by his adoption of spiritualism, other scholars have disagreed, and some maintain that Wallace never believed natural selection applied to those areas. Reaction to Wallace's ideas on this topic among leading naturalists at the time varied. Lyell endorsed Wallace's views on human evolution rather than Darwin's. Wallace's belief that human consciousness could not be entirely a product of purely material causes was shared by a number of prominent intellectuals in the late 19th and early 20th centuries. All the same, many, including Huxley, Hooker, and Darwin himself, were critical of Wallace's views. As the historian of science and sceptic
Michael Shermer has stated, Wallace's views in this area were at odds with two major tenets of the emerging Darwinian philosophy. These were that evolution was not
teleological (purpose-driven), and that it was not
anthropocentric (human-centred). Much later in his life Wallace returned to these themes, that evolution suggested that the universe might have a purpose, and that certain aspects of living organisms might not be explainable in terms of purely materialistic processes. He set out his ideas in a 1909 magazine article entitled
The World of Life. In 1910, Wallace published his final views on evolution, natural selection, and design in
The World of Life : A Manifestation of Creative Power, Directive Mind and Ultimate Purpose where he discussed among other things, the distribution of species, the evolution of life, the geological record, the emergence of research into living cells and microbiology, elements and water in relation to life, pain among living creatures. Wallace anticipated ideas about design in nature and directed evolution that would arise from religious traditions throughout the 20th century.
Wallace in the history of evolutionary theory In many accounts of the development of evolutionary theory, Wallace is mentioned only in passing as simply being the stimulus to the publication of Darwin's own theory. In reality, Wallace developed his own distinct evolutionary views which diverged from Darwin's, and was considered by many (especially Darwin) to be a leading thinker on evolution in his day, whose ideas could not be ignored. One historian of science has pointed out that, through both private correspondence and published works, Darwin and Wallace exchanged knowledge and stimulated each other's ideas and theories over an extended period. Wallace is the most-cited naturalist in Darwin's
Descent of Man, occasionally in strong disagreement. Darwin and Wallace agreed on the importance of natural selection, and some of the factors responsible for it: competition between species and geographical isolation. But Wallace believed that evolution had a purpose ("teleology") in maintaining species' fitness to their environment, whereas Darwin hesitated to attribute any purpose to a random natural process. Scientific discoveries since the 19th century support Darwin's viewpoint, by identifying additional mechanisms and triggers such as mutations triggered by environmental radiation or mutagenic chemicals. Wallace remained an ardent defender of natural selection for the rest of his life. By the 1880s, evolution was widely accepted in scientific circles, but natural selection less so. Wallace's 1889
Darwinism was a response to the scientific critics of natural selection. Of all Wallace's books, it is the most cited by scholarly publications. == Other scientific contributions ==