Sunday 10 August 2014

Natural selection before Darwin and Wallace

[22.08.2016: update on Prichard (1826)]
[13.08.2016: update on Diderot (1749)]
[19.06.2016: update on Fennel (1836)]
[18.06.2016: update on Lyell (1832)]
[30.05.2016: update on Maupertuis (1751)]
[12.03.2016: update on Corbaux (1829)]
[12.06.2015: update on Lamarck (1809), Erasmus Darwin (1818), A.P. de Candolle (1820), Naudin (1852)]
[14.06.2015: update on Blyth (1835)]
[27.02.2015: update on Loudon (1822)]
[13.11.2014: update: Conway Zirkle (1941. Natural selection before the "Origin of Species." Proceedings of the American Philosophical Society 84.1: pp. 71-123) reviews the history of the idea starting in antiquity with Empedocles. He also quotes translations showing, for example, that al-Jâhiz described the struggle for survival (though not the survival of the fittest it seems).]
[29.10.2014: update on Malthus (1798)]
[08.10.2014: update on Townsend (1786), Adams (1814), Wells (1813/18) and Matthew (1831)]

By other names, selection has been a household concept of breeders and the survival of the fittest has been a widespread idea. It was part of Natural Theology, but it could not transform a species in that context. On the contrary, many early formulations of natural selection saw it as conserving rather than transforming principle. In retrospective, natural selection was paradoxically taken as an argument for the fixity of species. For natural selection to be, what it is today, all the other parts of the Darwinian theory (e.g., heritable variation) also need to be in place. Therefore, it is wrong to speak of the 'discovery' of natural selection as if nobody had ever thought of it before. It may have been the last piece of the huge puzzle for Charles Darwin, but the piece has been known long before as an integral part to various puzzles showing different pictures.

Denis Diderot, in 1749, anonymously published the Lettre sur les aveugles. The pages 77-112 therein contain a passage of which I give the English translation by Margaret Jourdain (1916, pp. 111-112). The scene is a talk between a blind philosopher Saounderson and a clergyman, Mr. Holmes, summoned to the philosopher's deathbed: 
"Think, if you choose, that the design ['l'ordre' in the original French] which strikes you so powerfully has always subsisted, but allow me my own contrary opinion, and allow me to believe that if we went back to the origin ['naissance' in the original] of things and scenes and perceived matter in motion and the evolution from chaos ['le chaos se débrouiller' in original], we should meet with a number of shapeless creatures, instead of a few creatures highly organised. I make no criticism on the present state of things, but I can ask you some questions as to the past. For instance, I may ask you and Leibniz and Clarke and Newton, who told you that in the first instance of the formation some were not headless and others footless? I might affirm that such an one had no stomach, another no intestines, that some which seemed to deserve a long duration from their possession of a stomach, palate, and teeth came to an end owing to some defect in the heart or lungs; that monsters mutually destroyed one another; that all the defective combinations of matter disappeared, and that those only survived whose mechanism was not defective in any important particular and who were able to support and perpetuate themselves.1
      Suppose the first man had his larynx closed, or had lacked suitable food, or had been defective in the organs of generation, or had failed to find a mate, or had propagated in another species, what then, Mr Holmes, would have been the fate of the human race? It would have been still merged in the general depuration of the universe, and that proud being who calls himself man, dissolved and dispersed among the molecules of matter, would have remained perhaps for ever hidden among the number of mere possibilities. If shapeless creatures had never existed, you would not fail to assert that none will ever appear, and that I am throwing myself headlong into chimerical fancies, but the order is not even now so perfect as to exclude the occasional appearance of monstrosities."

1 This is the thesis of Lucretius, and the theory of the survival of the fittest.—(A)
Pierre Louis Moreau de Maupertuis (1751. Essay de Cosmologie) is, so far, the earliest source I know, in which the principle of natural selection or survival of the fittest was clearly stated. As typical for this time, however, natural selection was thought to keep the species fixed to their place in nature (we'd say niche). Natural selection was a mechanism of god's providence and thought to be mutually exclusive with species transmutation. This is why the sub-heading of this essay reads: "Which examines the evidence for God's existence from the Wonders of Nature."  

“Mais ne pourroit-on pas dire, que dans la combinaison fortuite des productions de la Nature, comme il n'y avoit que celles où se trouvoient certains rapports de convenance, qui pussent subsister, il n'est pas merveilleux que cette convenance se trouve dans toutes les eípèces qui actuellement existent? Le hazard, diroit-on, avoit produit une multitude innombrable d'Individus; un petit nombre se trouvoit construit de manière que les parties de l Animal pou voient íàtisfaire à ses besoins; dans un autre infiniment plus grand, il n'y avoit ni convenance, ni ordre: tous ces derniers ont péri: des Animaux sans bouche ne pouvoient pas vivre, d'autres qui manquoient d'organes pour la génération ne pouvoient pas se perpétuer: les seuls qui sòient restés sont ceux où se trouvoient l'ordre & la convenance & ces espèces, que nous voyons aujourd'hui, ne sont que la plus petite partie de ce qu'un destin aveugle avoit produit.” (Maupertius 1751. Essay de Cosmologie, p. 24-26)
This passage is often, falsely, attributed to Maupertuis' earlier publication Venus Physique. Its reproduction is then, usually, deviant (e.g., starting with "Ne pourrait-on" instead of "Mais ne pourroit-on") and cited without giving page numbers. Anyway, the above passage roughly translates:

“Could one not say that, in the fortuitous combinations of the productions of nature, none but those that found themselves in certain relations of appropriateness could subsist, is it not wonderful that this appropriateness is present in all the species that are currently in existence? Chance, one would say, produced an innumerable multitude of individuals; a small number found themselves constructed in such a manner that the parts of the animal were able to satisfy its needs; in another infinitely greater number, there was neither appropriateness nor order: all of these latter have perished. Animals lacking a mouth could not live; others lacking reproductive organs could not perpetuate themselves: the species we see today are but the smallest part of what blind destiny has produced.”
Some translate "convenace" with "fitness" instead of "appropriateness," in order to make the semblance clearer.

Joseph Tonwsend 1786.
republished in 1817, "A Dissertation on the Poor Laws, by a well-wisher to mankind." (London: Ridgeways) seems to have inspired Thomas Mathus's (1798) Essay on the Principle of Population (see here for a separate blog post on this issue).

In it is a passage containing a short statement of natural selection (survival of the fittest). It is also highly reminiscent of later population ecological reasoning. But it is given in a context of discussing the poor laws and not organic evolution. That is, it is not proposed as a mechanism for the transformation of species. It could not possibly do so on its own without heritable variation and the other parts of the Darwinian explanatory system in place.

The passage begins with an example of an island that is regularly visited by English sailors, because of a population of goats used for food.
"When the Spaniards found that the English privateers resorted to this island for provisions, they resolved on the total extirpation of these goats, and for this purpose they put on shore a greyhound dog and a bitch. [reference omitted] These in their turn increased and multiplied, in proportion to the quantity of food they met with; but in consequence, as the Spaniards had foreseen, the breed of goats diminished. Had they been totally destroyed, the dogs likewise must have perished. But as many of the goats retired to the craggy rocks, where the dogs could never follow them, descending only for short intervals to feed with fear and circumspection in the vallies, few of these, besides the careless and the rash, became a prey, and none but the most watchful, strong, and active of the dogs could get a sufficiency of food. Thus a new kind of balance was established. The weakest of both species were among the first to pay the debt of nature; the most active and vigorous preserved their lives. It is the quantity of food of food which regulates the number of the human species." (Townsend 1817, Sect. VIII, p. 44f)
He then gets back to his topic of poor laws, commons and his economic/political agenda.

James Hutton 1794. "Investigation of the Principles of Knowledge and of the Progress of Reason, from Sense to Science and Philosophy." (Vol. 2, section 13, chap. 3, p. 501") reproduced in the Supplementary information to Paul N. Pearson's "In Retrospect." (2003. Nature 425, p. 665):
"Now, this [adaptation through seminal variation and selection] will be evident, when we consider, that if an organised body is not in the situation and circumstances best adapted to its sustenance and propagation, then, in conceiving an indefinite variety among the individuals of that species, we must be assured, that, on the one hand, those which depart most from the best adapted constitution, will be the most liable to perish, while, on the other hand, those organised bodies, which most approach to the best constitution for the present circumstances, will be best adapted to continue, in preserving themselves and multiplying the individuals of their race." 
Hutton continues with the human breeding and selection of different dog races as an example and generalizes the principle to selection under natural conditions, though he does not apply the term selection to the natural process. He also did not believe that this natural process could lead to the transmutation of species, it seems, but only to the adaptation of races within species. That is, the idea of natural selection is as old as that of species transmutation, but they used to be regarded as mutually exclusive. It is therefore a false abridgement to reduce the evolutionary paradigm shift to the 'discovery of natural selection' or the 'discovery of the mechanism of evolution.' The mechanism was long since known, but it was fitted into theory so as to prevent the transmutation of species. The paradigm shift was to combine two old ideas that previously seemed to be mutually exclusive. 
     As Pearson (2003) explains, Hutton's Investigation is an intimidating philosophical magnum opus in three volumes running up to more than 2000 pages that were already deemed obscure by his contemporaries. It is, therefore, no wonder that the chapter on natural selection in the second volume has been ignored for so long. It's an example of the theory of evolution through natural selection being proposed in an odd context.

Thomas R. Malthus 1798. An Essay on the Principle of Population ) was a major source of inspiration for Matthew, Wallace and Darwin for their insight into the importance of natural selection in evolution. Malthus (1798, p. 11) starts from two simple postulates:
"I think I may fairly make two postulata. First, That food is necessary to the existence of man. Secondly, That the passion between the sexes is necessary and will remain nearly in its present state."
From these postulates he deduces a law of nature (Malthus 1798, p. 13f):
"Assuming then my postulata as granted, I say, that the power of population is indefinitely greater than the power in the earth to produce subsistence for man.
Population, when unchecked, increases in a geometrical ratio. Subsistence increases only in an arithmetical ratio. A slight acquaintance with numbers will shew the immensity of the first power in comparison of the second.
By that law of our nature which makes food necessary to the life of man, the effects of these two unequal powers must be kept equal."
And he illustrates the effect of that law of nature (Malthus 1798, p. 15):
The race of plants and the race of animals shrink under this great restrictive law. And the race of man cannot, by any efforts of reason, escape from it. Among plants and animals its effects are waste of seed, sickness, and premature death. Among mankind, misery and vice.
Malthus's law of nature can justly be called by its current name, which is natural selection. The only thing that is still missing, here, is the idea of heritable variation acting in concert with this natural selection. That a lot of variation is heritable, however, was a triviality for plant and animal breeders of all stripes. It is therefore quite understandable that the following predecessors all independently hit on the same idea of combining these two explanatory parts. Though this did not necessarily mean the transformation of species.

Jean-Baptiste Lamarck 1809. Lamarck (Philosophie Zoologique) had use-inheritance in place of natural selection and does therefore not belong to the anticipators of natural selection in the strict sense. He nevertheless suggested an identity of the causal relations underlying culture (domestication) and nature that would now be read as an identification of artificial with natural selection (see here). Charles Lyell (Principles of Geology 1830-33), by the way, translated much of Lamarck's ideas and was likely an important secondary source for English readers.

William Charles Wells 1813/1818. ("Two Essays: one upon single vision with two eyes; the other on dew...". Archibald Constable and Co., Edinburgh). Pages 423-437 contain An Account of a Female of the White Race of Mankind, part of whose skin resembles that of a negro; with some observations on the causes of the differences in colour and form between the white and negro races of men This has been read before the Royal Society of London in 1813, but not been published in its transactions. This is interesting because the next anticipator (Adams 1814) may have misquoted Wells:
"Again, those who attend to the improvement of domestic animals, when they find individuals possessing, in a greater degree than common, the qualities they desire, couple a male and female of these together, then take the best of their offspring as a new stock, and in this way proceed, till they approach as near the point in view, as the nature of things will permit. But, what is here done by art, seems to be done, with equal efficacy, though more slowly, by nature, in the formation of varieties of mankind fitted for the country which they inhabit." (p. 435)    
"Similar facts occur in respect to other species of animals." (p. 437) 
The context is a collection of pieces—not just two as the title of the book suggests. Starting out rather medically, about a woman with a mosaic skin pigmentation, it descends into speculations about human races and their adaptation to their environment - much of it racist and no longer politically correct I guess. Again, this is an example of odd context for a theory of evolution through natural selection. The whole collection in the book can also be considered odd, as it contains mixture of essays medical, physical, autobiographical etc.

Joseph Adams 1814. (A Treatise on Hereditary Disease." London: J. Callow). Mind that the context is a treatise on hereditary disease meant for the general public. The public seems to have been frightened by the misconception that hereditary meant doom to the children of those who had a hereditary disease in the family. Adams anticipated many concepts of medical genetics, which would now be called recessiveness, penetrance etc. (see Weiss 2008). 

Adams (1814, p. 32f):
"In a state of nature the race of all gregarious animals is probably progressively improving, as far as is consistent with their capacity for improvement. The strongest male becomes the virgregis, and consequently, the father of most of the offspring. In a ruder state of human society, or rather in its earliest formation, something of the same kind may prevail; but in a more advanced stage, sufficient provision is made by the preferences which health and intellect will for the most part produce in either sex.
Another provision arises out of climate; which we have seen is, in some cases, the only means of exciting a diseased susceptibility into action. Those constitutions, which are peculiarly susceptible of such diseases as are excited by climate, fall an early sacrifice; hence, the propagation from sources gradually lessens, and the disease would cease altogether, were it not that parents, free from such susceptibility, occasionally produce an offspring in whom the susceptibility originates.  [...]
By these means a race is gradually reared with constitutions best calculated for the climate: a law which, I suspect, has been too much overlooked, in our inquiries after the causes of the more marked varieties in the human species."
He continues with examples of artificial selection by breeders, especially the ill consequences of inbreeding, ruminates on eugenic laws against inbreeding, and so on. See for yourself or read the article by Weiss (2008), who also discussed other anticipations including Matthew's.

Erasmus Darwin 1818. The grandfather's Zoonomia contains an anticipation of natural or rather sexual selection (see also here):
"The birds, which do not carry food to their young, and do not
therefore marry, are armed with spurs for the purpose of fighting
for the exclusive possession of the females, as cocks and quails.
It is certain that these weapons are not provided for their defence
against other adversaries, because the females of these species are
without this armour. The final cause of this contest amongst the
males seems to be, that the strongest and most active animal should
propagate the species, which should thence become improved."

Augustin Pyramus de Candolle 1820. Géographie Botanique contained the following passage:
"All the plants of a country, those of a given location, are in a state of war each. All are equipped with means of reproduction and nutrition more or less effective. The first that establish themselves by chance in a given location, tend, by the mere fact that they occupy the ground, to exclude other species: the biggest stifle the smaller; the more perennial replace those with a shorter duration; the most fertile gradually seize the space that could otherwise be filled by slower multiplying ones." (my translation)
For the French original passage and further details see also here. Again, Lyell's Principles of Geology seem to have provided the first English translation.

John Claudius Loudon, 1822 ("An Encyclopaedia of Gardening." London: Longman, Hurst, Rees, Brown and Green), is particularly relevant to the claim, sometimes aired, that Darwin and Wallace plagiarised Matthew (see below) because Matthew, in turn, lifted his insights from Loudon (see here). The standards of citation were just different, I guess, and authors felt no need to cite the sources from which they have taken the pieces of a puzzle if their solution of the puzzle was new. The following quote is from page 230 (bold emphasis mine):
"465. From the various sources of vegetable reproduction, but particularly from the fertility and dispersion of the seed, the earth would soon be over-run with plants of the most prolific species, and converted again into a desert, if it were not that nature has set bounds to their propagation by subjecting them to the control of man, and to the depredations of the great mass of animals; as well as in confining the germination of their seeds to certain and peculiar habitations arising from soil, climate, altitude, and other circumstances. In order to form an idea of the manner in which these act upon vegetation; imagine that every year an enormous quantity of seeds, produced by the existing vegetables, are spread over the surface of the globe, by the winds and other causes already mentioned, all of these seeds which fall in places suitable for their vegetation, and are not destroyed by animals, germinate and produce plants; then among these plants, the strongest, and largest, and those to which the soil is best suited, developed themselves in number and magnitude so as to choke the others. Such is the general progress of nature, and among plants, as among animals, the strong flourish at the expense of the weak. These causes have operated for such a length of time, that the greater number of species are now fixed and considered as belonging to certain soils, situations, and climates, beyond which they seldom propagate themselves otherwise than by the hands of man."  

James Cowles Prichard 1826. Researches into the physical history of mankind. Second edition, Vol. 2. John and Arthur Arch, Cornhill. Spoiler: racism. Page 573:
It seems that people descended from the European nations bear with difficulty an abode in Africa between the tropics. The insalubrity of the intertropical African climate to the constitutions of Europeans is extreme, and has hitherto been sufficient, notwithstanding innumerable attempts at colonization, to prevent the growth of any white population on the African coasts. By Phoenician and Arabian traders, the eastern coast, or Zanguebar, had been visited almost before the era of history; many colonies were doubtless settled there in early times, and very many, as we know, in the middle ages; many after the Portugueze discovery, both on the old coast of Zanguebar and on the shores of Guinea, and of Congo. Yet no white population exists in tropical Africa. At Mozambique indeed the mortality is so great, that a banishment to that colony is held at Goa equivalent to an execution, and only malefactors are sent thither, where they die in a few years. The mortality in the late English expedition to the river Zaire was surprising and mysterious. These are apparently the reasons why white colonies have never thrived in Africa, where we should otherwise have now a white population as great as that descended from the English or the Spaniards in America.
     With respect to the West Indies, it is well known that the incapability of Europeans to support labour in that tropical country was the first occasion of introducing into the islands an African population. On the other hand, Negroes seem under a physical disability to establish themselves in Europe and other northern countries, otherwise we should find them here in numbers. They are here, more than any other class of people, subject to phthisical and scrofulous complaints, and are seldom, under the most favourable circumstances, healthy. The diseases to which both kinds of people are subject in the climate appropriated to the other, is the impediment which has prevented large colonies of whites from forming themselves and multiplying in tropical Africa, and of Negroes in the North. We are told indeed by Herodotus, that there was once founded at Colchis a colony of African blacks, but they have long since dwindled away and disappeared.
     It appears to result from the foregoing facts, to which a great many others might probably be added, that in mankind, as in some other races, particular varieties are adapted by constitution and physical peculiarities to particular local situations. These remarks, if they are well founded, serve to illustrate the doctrine of variation, or deviation, in the races of animals in general, and they seem to lead us to the conclusion, that this is not merely an accidental phaenomenon, but a part of the provision of nature for furnishing to each region an appropriate stock of inhabitants, or for modifying the structure and constitution of species, in such a way as to produce races fitted for each mode and condition of existence. A great part of this plan of local adaptation appears to have been accomplished by the original modification of a genus into a variety of species. It has been further continued, and the same end promoted, by the ramification of a species into several varieties.

And from page 581:
A question now presents itself; how these varieties are developed and preserved in connexion with particular climates and differences of local situation?
      One cause which tends to maintain this relation is obvious. Individuals and families, and even whole colonies, perish and disappear in climates for which they are, by peculiarity of constitution, not adapted. Of this fact proofs have been already mentioned.
      Besides, it appears probable that those local circumstances, which are most congenial to particular races, do in fact promote the appearance of those varieties which are best suited to them, or tend to give rise to their production in the breed.

Francis Corbaux 1829. ("On the laws of mortality, and the intensity of human life." The Philosophical Magazine 5: 198-205.) Corbaux formulated an idea of natural selection in an odd combination with senescence. In one passage he even wrote about the "natural selection" due to individual competition between centenaries. This does not only show that he had not fully understood that natural selection can only have an effect when working on individuals that can still reproduce and inherit their anomalies, it also proves that the next anticipator, Patrick Matthew, did not coin the term "natural selection." This is important with respect to some people who claim that Darwin plagiarized Matthew based mainly on such logomachy. Here's the passage:
"At a certain age, which may vary from the eighty-third to the ninetieth year, according to the description of a whole population or any select portion of it, an anomaly is exhibited in the shape of apparent increase, as to the intensity of life, during a few years. Not that individual lives have actually improved; but considered in the aggregate, such as were originally constituted for outliving their contemporaries, and who continued to exist under the most favourable circumstances, ultimately stand prominent, competing amongst themselves for protracted longevity, to the exclusion of all the rest. Indeed, this natural selection of particular lives, out of a very considerable mass, repeatedly occurs among centenaries, at later periods, and according to their respective degrees of constitutional vigour; so that very little difference may appear in the probabilities of living one more year, between two individuals of whom the ages differed even to the extent of twenty years." (Corbaux 1829, p. 201)
Patrick Matthew 1831. ("On Naval Timber and Arboriculture." Edinburgh: Adam Black, Edinburgh). On my closer reading of Matthew (1831), I noticed that various scattered passages in the main text have been informed by his idea of natural selection with heritable variation. Some of these occur in Part IV called Notice of Authors who treat of Arboriculture, in which Matthew reviewed works of other scholars on arboriculture. That is, Matthew (1831) would head a chapter reviewing a certain scholar by, for example, The Forester's Guide, by Mr. Monteath. The running head throughout that chapter would alternate between Notice of Authors and Monteath's Forster's Guide on every other page. From my perusal, I got away with the impression that selection was a household term in the trade of tree breeding, as much as it used to be among breeders of any other kind of organism. As is well known, Darwin consulted many breeders during his years of incubating The Origin. It is not hard to imagine that, from this context, both scholars took the little step from human to natural selection independently of each other. 
      Appart from these scattered rubies in the rubbish of the main text, the Appendix is remarkable. It consists of two parts. The first is a collection of Endnotes A to F. These endnotes were too long for inclusion as footnotes at the bottom of the pages in the main text. Therefore, the respective footnotes in the main text refer to these endnotes instead. For example, a footnote at page 3 goes "See App. B." Page 3, however, belongs to the introduction of the book, which extols the relevance of navigation for trade, culture and Britain's dominion. Said Note B begins (p. 364f):
"There is a universal law in nature, tending to render every reproductive being the best suited to its condition that its kind, or that organized matter, is susceptible of, which appears intended to model the physical and mental or instinctive powers, to their highest perfection, and to continue them so. This law sustains the lion in his strength, the hare in her swiftness, and the fox in his wiles. As nature in all her modifications of life, has a power to increase far beyond what is needed to supply the place of what falls by Time's decay, those individuals who possess not the requisite strength, swiftness, hardihood, or cunning, fall prematurely without reproducing--either a prey to their natural devourers, or  sinking uner disease, generally induced by want of nourishment, their place being occupied by the more perfect of their own kind, who pressing on the means of subsistenece." 
The rest of Note B drifts off into a rant against feudal laws such as the law of entail of aristocracy as being against this natural law of selection. The persistent reader will get a big surprise after the end of the last endnote (Note F). After a horizontal line, Matthew (1831, pp. 381-388) simply continues with an appendix to the Appendix, which is nowhere referred to in the main text. This, however, is the place where he coherently explains his idea of natural selection of heritable variants and even the origin of new species:
"The progeny of the same parents, under great difference of circumstances, might, in several generations, even become distinct species, incapable of co-reproduction." (Matthew 1831, p. 384)
For many more blog posts concerned with Matthew click the corresponding tab at the top of this blog see here

I have myself overlooked the appendix to the Appendix at first, and the rubies in the rubbish of the main text are also easy to overlook. It seems fair to say that these are obscure places in an obscurely written book, even if the author was not unknown, but was widely read and cited for his practical advice on planting, pruning etc.

Charles Lyell 1832. Although Lyell's Principles of Geology are usually taken to be on geology only, volume 2 is a book on biology rather than geology. For example, chapter 2 summarizes and dismissed Lamarck's theory of species transmutation. However, Lyell also mentioned the "struggle for existence" verbatim at the pages 55-56
"If we consider the vegetable kingdom generally, it must be recollected, that even of the seeds which are well ripened, the greater part are either eaten by insects, birds, and other animals, or decay for want of room and opportunity to germinate. Unhealthy plants are the first which are cut off by causes prejudicial to the species, being usually stifled by more vigorous individuals of their own kind. If, therefore, the relative fecundity or hardiness of hybrids be in the least degree inferior, they cannot maintain their footing for many generations, even if they were ever produced beyond one generation in a wild state. In the universal struggle for existence, the right of the strongest eventually prevails; and the strength and durability of a race depends mainly on its prolificness, in which hybrids are acknowledged to be deficient." 
Edward Blyth 1835. An Attempt to classify the "Varieties" of Animals, with Observations on the marked Seasonal and other Changes which naturally take place in various British Species, and which do not constitute Varieties. (The Magazine of Natural History 8: 40-46).
"It is a general law of nature for all creatures to propagate the like of themselves: and this extends even to the most trivial minutiae, to the slightest individual peculiarities; and thus, among ourselves, we see a family likeness transmitted from generation to generation. When two animals are matched together, each remarkable for a certain given peculiarity, no matter how trivial, there is also a decided tendency in nature for that peculiarity to increase; and if the produce of these animals be set apart, and only those in which the same peculiarity is most apparent, be selected to breed from, the next generation will possess it in a still more remarkable degree; and so on, till at length the variety I designate a breed, is formed, which may be very unlike the original type.
      The examples of this class of varieties must be too obvious to need specification: many of the varieties of cattle, and, in all probability, the greater number of those of domestic pigeons, have been generally brought about in this manner. It is worthy of remark, however, that the original and typical form of an animal is in great measure kept up by the same identical means by which a true breed is produced. The original form of a species is unquestionably better adapted to its natural habits than any modification of that form; and, as the sexual passions excite to rivalry and conflict, and the stronger must always prevail over the weaker, the latter, in a state of nature, is allowed but few opportunities of continuing its race. In a large herd of cattle, the strongest bull drives from him all the younger and weaker individuals of his own sex, and remains sole master of the herd; so that all the young which are produced must have had their origin from one which possessed the maximum of power and physical strength; and which, consequently, in the struggle for existence, was the best able to maintain his ground, and defend himself from every enemy. In like manner, among animals which procure their food by means of their agility, strength, or delicacy of sense, the one best organised must always obtain the greatest quantity; and must, therefore, become physically the strongest, and be thus enabled, by routing its opponents, to transmit its superior qualities to a greater number of offspring. The same law, therefore, which was intended by Providence to keep up the typical qualities of a species, can be easily converted by man into a means of raising different varieties; but it is also clear that, if man did not keep up these breeds by regulating the sexual intercourse, they would all naturally soon revert to the original type. Farther, it is only on this principle that we can satisfactorily account for the degenerating effects said to be produced by the much-censured practice of "breeding in and in." There would almost seem, in some species, to be a tendency, in every separate family, to some particular kind of deviation; which is only counteracted by the various crossings which, in a state of nature, must take place, and by the above-mentioned law, which causes each race to be chiefly propagated by the most typical and perfect individuals." Blyth (1835, 45f)

"Still, however, it may not be impertinent to remark here, that, as in the brute creation, by a wise provision, the typical characters of a species are, in a state of nature, preserved by those individuals chiefly propagating, whose organisation is the most perfect, and which, consequently, by their superior energy and physical powers, are enabled to vanquish and drive away the weak and sickly, so in the human race degeneration is, in great measure, prevented by the innate and natural preference which is always given to the most comely; and this is the principal and main reason why the varieties which are produced in savage tribes, must generally either become extinct in the first generation, or, if propagated, would most likely be left to themselve, and so become the origin of a new race; and in this we see an adequate cause for the obscurity in which the origin of different races is involved." Blyth (1835, 48f)  

Fennel 1836. James H. Fennel replied to Blyth's above quoted article in the Short Communications section of the same journal. While he did not repeat the principle of natural selection, he contradicted Blyth on the fixity of species (Magazine of Natural History 9: 647-648). Remembering that Darwin owned these volumes and scrutinized them carefully, it is clear that he put the pieces of a huge puzzle into place rather than creating the pieces of the puzzle from scratch. 
"Now, it appears to me that he does not make a sufficient allowance for the modifications of an animal, gradually incurred by a difference of circumstance. [...] I believe that any habit or structure is liable to, or capable of, modification according to locality, with its peculiar climate, soil, produce, &c. [...] Though a species may have continued to propagate, without any intermixture with other species, which would produce different appearances, if not habits, yet I do not think that the order of nature has offered any opposition to the structure and habits of any animal becoming, from the cause I have mentioned, gradually altered, in a greater or less degree, from those of its primitive parents. My views may seem to involve the Lamarckian theory; but they do not. On some more convenient occasion, I hope to speak at large on the changes which I believe, after much observation and reflection, to have taken place in the works of nature, not only in the habit, but in the structure."

Charles Naudin 1852. Published a highly important article in the Revue Horticole, that is unjustly forgotten. He anticipated much more than merely the principle of natural selection, for example, the tree-of-life metaphor (see here and here for further details). 

"We do not think that Nature has made its species in a different fashion from that in which we proceed ourselves in order to make our varieties; or better, we carried it's [Nature's] process into our practice." (p. 104)
"Such is, in our ideas, the course followed by nature; like us, it wanted to form races appropriate for their needs; and with a relatively small number of primordial kinds, she gave birth in succession and at various times, to all plant and animal species that inhabit the globe." (p. 104)
"Nature has operated on an immense scale and with immense resources; we, on the contrary, we do so with extremely limited means; but between its processes and ours, between his results and those we get, the difference is in any amount; between its species and those we create, there are only the more and less."

Herbert Spencer 1852. ("A theory of population, deduced from the general law of animal fertility." Westminster Review 57: 468-501):
"And here it must be remarked, that the effect of pressure of population, in increasing the ability to maintain life, and decreasing the ability to multiply, is not a uniform effect, but an average one. In this case, as in many others, Nature secures each step in advance by a succession of trials, which are perpetually repeated, and cannot fail to be repeated, until success is achieved. All mankind in turn subject themselves more or less to the discipline described; they either may or may not advance under it; but, in the nature of things, only those who do advance under it eventually survive. For, necessarily, families and races [page break: 499/500] whom this increasing difficulty of getting a living which excess of fertility entails, does not stimulate to improvements in production—that is, to greater mental activity—are on the high road to extinction; and must ultimately be supplanted by those whom the pressure does so stimulate. This truth we have recently seen exemplified in Ireland. And here, indeed, without further illustration, it will be seen that premature death, under all its forms, and from all its causes, cannot fail to work in the same direction. For as those prematurely carried off must, in the average of cases, be those in whom the power of self-preservation is the least, it unavoidably follows, that those left behind to continue the race are those in whom the power of self-preservation is the greatest—are the select of their generation. So that, whether the dangers to existence be of the kind produced by excess of fertility, or of any other kind, it is clear, that by the ceaseless exercise of the faculties needed to contend with them, and by the death of all men who fail to contend with them successfully, there is ensured a constant progress towards a higher degree of skill, intelligence, and self-regulation—a better co-ordination of actions—a more complete life." [pp. 499/500.]
The Westminster Review was established in 1823 as the official organ of the Philosophical Radicals, a group of English political radicals with philosophical inspirations. In this respect, it can be regarded as an odd place of publication quite unlike the journals and outlets of scientific societies. This group of philosophers believed in a loosely defined evolutionism or 'Law of Progress,' but was divided over the merit of Malthus's principle. Some rejected it as a justification for workhouse morals. Herbert Spencer published his article in order to show that the adaptedness or fitness of organisms for their environment does not result from some benevolent government of nature, but from the painful Malthusian principle combined with some self-correcting mechanism, as it were, natural selection. I would not regard this context as very odd because it seems to have been the larger (political/ideological) context for evolutionary theory ever since.

There might even be yet unknown anticipations in other languages.