Abstract
This work takes a stand on whether Wallace should be regarded as co-author of the theory of natural selection alongside Darwin as he is usually considered on behalf of his alleged essential contribution to the conception of the theory. It does so from a perspective unexplored thus far: we will argue for Darwin’s priority based on a rational reconstruction of the theory of natural selection (following a hierarchical perspective for scientific theories, we present its conceptual framework together with its fundamental and special laws) as it appears in the writings of both authors. We show that the theory does not appear in exactly the same manner in the writings of each of its alleged co-discoverers: tough we find the same fundamental elements in both works, even in Darwin’s early texts, we discern a more complex unifying and ramified structure than the one we find in Wallace’s Ternate manuscript. Even when we think the badge of the “Darwin–Wallace” theory is well deserved, the unifying force of Darwin’s version has proved to be one of the keys for the ulterior success of the so-called Darwinian revolution in Biology.
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Notes
Remarkably, Wallace did not use the expression “natural selection”. See note 14 on his opinion regarding the use of this nomenclature.
In Wallace’s own words (Wallace 1908, pp. 5–7):
Since the death of Darwin in 1882, I have found myself in the somewhat unusual position of receiving credit and praise from popular writers under a complete misapprehension of what my share in Darwin’s work really amounted to. It has been stated (not unfrequently) in the daily and weekly press, that Darwin and myself discovered “natural selection” simultaneously, while a more daring few have declared that I was the first to discover it, and that I gave way to Darwin! [...] He [Darwin] would have been at once recognised, and should be ever recognised, as the sole and undisputed discoverer and patient investigator of the great law of “Natural Selection” in all its far-reaching consequences.
Despite subsequent debates [see the well documented work of Malcolm Kottler on this (Kottler 1985)], both Darwin and Wallace stressed the similarities when discussing originality. In 1858, and after receiving Wallace’s Ternate paper, Darwin wrote two famous letters to his close friend Charles Lyell, expressing desperate worries regarding priority. In the first one (dated June \(18{\mathrm{th}}\)) he wrote: “I never saw a more striking coincidence [between his ideas and Wallace’s]. If Wallace had my M.S. sketch written out in 1842 he could not have made a better short abstract! Even his terms now stand as Heads of my Chapters.” (Burkhardt and Smith 1992, p. 107) In the second letter (dated June \(25{\mathrm{th}}\)) he wrote that “there is nothing in Wallace’s sketch which is not written out much fuller in my sketch copied in 1844” (Burkhardt and Smith 1992, p. 117). Several years later, on May \(14{\mathrm{th}}\), 1864, Hooker called Darwin’s attention to a recent writing of Wallace’s (Wallace 1864) in which the latter did not claim any authorship whatsoever in the theory –he uses there expressions such as “Mr. Darwin’s celebrated theory of natural selection”, and “the theory promulgated by Mr. Darwin”. This perplexed Hooker who said “I am struck [...] with his negation of all credit or share in the Natural Selection theory” (Hooker 1864). Darwin’s reaction was to write to Wallace and admonish him thus: “You ought not [...] speak of the theory as mine; it is just as much yours as mine.” (Burkhardt 2001, p. 216)
Endler is not alone in this. Many other authors claim that fitness can be determined independently of survival (Ginnobili 2010, 2016; Gould 1976; Kutschera 2003, p. 354; Lerner 1959; Naylor and Handford 1985; Wassermann 1978). On the other hand, Lennox and Wilson think that, for Darwin, the struggle for existence is necessary for natural selection to occur (Lennox and Wilson 1994).
The same idea can be found in other authors. For example, Castrodeza (1988, pp. 182–183), Bateman (1948), Mayr (1972), Ghiselin (1969, p. 242), Gruber (1974, pp. 87–88), Thornhill and Alcock (1983), Spencer and Masters (1992), Dobzhansky et al. (1977, p. 118), Ayala (1995, pp. 80–82) and especially Ghiselin (1997), p. 226.
Darwin himself told Lyell (in his letter of June \(18{\mathrm{th}}\), 1858) that “all the labour” in his Book “consists in the application of the theory” (Burkhardt and Smith 1992, p. 107). As a matter of fact, one of us performed just this task by going through every kind of practical application of the theory that we can find in OS, identifying both what they all share and their differences (Ginnobili 2010, 2012, 2016).
To conceiving sexual selection as a form of natural selection may be considered controversial. If natural selection is confined to what relates to survival, it is clear that sexual selection then constitutes a different construct. In fact, it could be argued that sometimes Darwin himself seemed to regard them as different for that very reason. See, for instance, this statement from the 1842 text:
Besides selection by death, in bisexual animals \(\langle \hbox {illegible}\rangle \) the selection in time of fullest vigour, namely struggle of males; even in animals which pair there seems a surplus \(\langle ? \rangle \) and a battle, possibly as in man more males produced than females, struggle of war or charms. Hence that male which at that time is in fullest vigour, or best armed with arms or ornaments of its species, will gain in hundreds of generations some small advantage and transmit such characters to its offspring. (Darwin 1909a, p. 10)
In OS this opposition is still more explicit:
[Sexual Selection] depends, not on a struggle for existence, but on a struggle between the males for possession of the females; the result is not death to the unsuccessful competitor, but few or no offspring. Sexual selection is, therefore, less rigorous than natural selection. (Darwin 1859, p. 88, our emphasis)
In these terms, sexual selection will have applications that exclude those of natural selection. On the other hand, and this is crucial, putting these discussions aside, it is enough for our purposes that the use of the same concepts be recognized in all the proposals we review here. This clue will enable us to see unity in the diversity of theories, as Endler and Tuomi did. Our treatment will be justified, we hope, later on in this section.
We insist: the connection between the trait that is adequate for the environment and success in differential reproduction may be due to different reasons: either an improvement in survival, or an improvement in mating, or an improvement in the ability to attract pollinators, etc., all of which may, in turn, be due to different causes.
This terminological choice may be strange for the reader. It is a predicament hard to overcome. We either choose terms are not typically used to talk about natural selection, which makes the text harder to read; or, rather, we opt for terms that are more familiar and run the risk of being misinterpreted or of upsetting those who use these terms in a different way. This is a consequence of the fact that in this discussion the number of concepts involved outruns the number of terms to express them.
Note that the theory of artificial selection can also be thought of as different from that of natural selection restricted to survival. This is hinted at in several passages from Darwin himself in later editions of OS (Darwin 1872, pp. 39, 80–81, 137–138), and from many commentators that follow him on this when they present these theories as being in competition (Beer 2001, p. 166; Karlin and Carmelli 1975; Minvielle 1981; Nicholas and Robertson 1980; Secord 1981) and deserving separate treatment. Even though the relationship between artificial selection and natural selection is undoubtedly debatable, here we assume the position that artificial selection can be judged as a case of natural selection (Álvarez 2010; Brandon 1978, 1990, p. 13, Gayon 1998, pp. 54–59; Campbell 1984, pp. 149–152; Richards 2014; Wilner 2006; Ginnobili 2011; among others). For our purposes, it is sufficient to know that that relationship has been discussed in Darwin’s version of the theory, and that it is absent in Wallace’s version because he explicitly denied that possibility (at least in the treated works).
We thank an anonymous reviewer for pointing this out to us.
Gray is one the few people we are aware of that had come to know Darwin’s transmutationist ideas before 1858 directly from Darwin’s own mouth or pen. Others were, chronologically, Joseph Hooker; Leonard Jenyns; Charles Lyell; William Fox; George Thwaites; Samuel Woodward; and James Dana (see Costa 2013, p. 229; Kottler 1985; Porter 1993). On the other hand, Darwin personally explained in detail the theory of natural selection to Lyell, Hooker and Thomas Huxley.
In a letter to Lyell, Darwin himself wrote that Wallace and him “differ only that I [Darwin] was led to my views from what artificial selection has done for domestic animals” (Burkhardt and Smith 1992, p. 117).
Let us remember that for this reason Wallace urges Darwin to substitute the expression “natural selection” for “survival of the fittest”. Darwin ended up following his colleague’s advice. In the fifth edition of OS he wrote:
I have called this principle, by which each slight variation, if useful, is preserved, by the term Natural Selection, in order to mark its relation to man’s power of selection. But the expression often used by Mr. Herbert Spencer of the Survival of the Fittest is more accurate, and is sometimes equally convenient. (Darwin 1869, p. 72)
Wallace’s proposal was delivered (once again) by correspondence, in a letter dated July \(2{\mathrm{nd}}\) 1866. As Darwin admits, Herbert Spencer (1820–1903) coined the expression, more specifically, in his The Principles of Biology of 1864 (Spencer 1864). Darwin adopted the Spencerian language in this fifth edition of OS and in the sixth one. However, he never dropped “natural selection” altogether, but preserved it, using both expressions interchangeably up to the end (Darwin 1872, pp. 80, 102, 134–135, 180, 471) (Adopting this terminology is inadequate for multiple reasons. To mention an oft-quoted one, it was precisely this formulation that drove many authors to (wrongly) declare the theory to be unfalsifiable on account of circularity.)
Cf. (Kutschera 2003), where we can find this and other interesting differences between Darwin’s and Wallace’s theoretical proposals. On account of our approach, we only pick up two: the difference in how they treat natural selection and Wallace’s omission of sexual selection, about which we will speak later in this section.
Note what Darwin claims at the beginning of his longer work The Variation of Animals and Plants under Domestication:
No doubt man selects varying individuals, sows their seeds, and again selects their varying offspring. [...] Man, therefore, may be said to have been trying an experiment on a gigantic scale; and it is an experiment which nature during the long lapse of time has incessantly tried. Hence, it follows that the principles of domestication are important for us. (Darwin 1868, vol. 1, p. 3; cf. Wilner 2006)
Just as breeders intervene positively selecting those traits which will be useful for their ends and they are interested in preserving, thereby creating in relatively few generations products that satisfy their practical or aesthetic agenda, in an analogous manner nature “chooses”, unintentionally but not randomly, “creating” and preserving new varieties for different reasons (Darwin 1859, Chapters 1–2). Again, Darwin attributes a greater creative power to the selective force of nature than he does to artificial selection.
So extraordinary that it can hardly be attributed to chance. Perhaps a historiographical analysis of their interaction before 1858 could shed light to an explanation (see Kottler 1985). Notwithstanding this, we should remember that these coincidences surprised Darwin himself (cf. note 3).
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Acknowledgements
We want to thank Roberto Biaggi, Damien Rochford and two anonymous referees for their helpful comments on previous versions of this paper. This research was supported by the research Projects PICT-2014-1741 (ANPCyT, Argentina), Research Program “Philosophy and History of Science” PUNQ, 1719/11 (UNQ, Argentina), and 32/15 255 (UNTREF, Argentina).
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Equal time was dedicated to the production of this article. Order of authorship was decided by chance.
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Ginnobili, S., Blanco, D. Wallace’s and Darwin’s natural selection theories. Synthese 196, 991–1017 (2019). https://doi.org/10.1007/s11229-017-1491-z
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DOI: https://doi.org/10.1007/s11229-017-1491-z