Was Alfred Russel Wallace a Group Selectionist?

By Dr George Beccaloni, September 2022

The quick answer is "no", but ironically, Charles Darwin was - see https://gardner.wp.st-andrews.ac.uk/files/2015/05/Gardner_2013b.pdf There is, however, a widespread misconception that Wallace was the group selectionist - so why is this the case? The fallacy originated from a misinterpretation (by non-biologists) of what Wallace meant by the term "variety". Historian Peter Bowler (Bowler, 1976), for example, has argued that that the "varieties" Wallace spoke about in his 1858 essay on natural selection, were subspecies rather than variant individuals (aberrations or sports) within a population. Bowler's assertion was, however, comprehensively rebuked by Malcolm Kottler (Kottler, 1985. See pdf HERE). Wallace's paper would simply not make much sense if you replaced "subspecies" for "variety"! In any case, reading entries made in 1855 in his unpublished 'Species Notebook' in the Linnean Society of London leaves one in no doubt that by "variety" he meant one or more individuals in a population which differ in some way from the ancestral state.

As long ago as 1896 Poulton considered this issue and concluded that:

"Although Wallace used the term "variety" as contrasted with "species," the whole context proves that he, equally with Darwin, recognised the importance of individual variations and of variations in single characters. This becomes clear when we remember his argument about the neck of the giraffe, the changes of colour and hairiness, the shorter legs of the antelope, and the less powerful wings of the passenger pigeon. Wallace has kindly written to me (May 12th, 1896) stating the case as I have given it, and he further explains—

'I used the term 'varieties' because 'varieties' were alone recognised at that time, individual variability being ignored or thought of no importance. My 'varieties' therefore included 'individual variations.'" [Read the full letter HERE]

It is important to note that the term 'variety' as used by zoologists (especially ones with a taxonomic bent, like Wallace) has always been ambiguous - even into the 20th century - and the meaning of the term depended on the context it was used. This issue is recognised by the International Commission on Zoological Nomenclature and discussed in the Code i.e.

45.6.3. [A taxonomic name]... is deemed to be infrasubspecific if it was first published after 1960 and the author expressly used one of the terms "variety" or "form" (including use of the terms "var.", "forma", "v." and "f.");

45.6.4. it is subspecific if first published before 1961 and its author expressly used one of the terms "variety" or "form" (including use of the terms "var.", "forma", "v." and "f."), unless its author also expressly gave it infrasubspecific rank, or the content of the work unambiguously reveals that the name was proposed for an infrasubspecific entity, in which case it is infrasubspecific [see also Art. 45.6.1]; except that a name that is infrasubspecific under Article 45.6.4 is nevertheless deemed to be subspecific from its original publication if, before 1985, it was either adopted as the valid name of a species or subspecies or was treated as a senior homonym.

Examples. Spencer (1896) described and named Sminthopsis murina var. constricta, a small carnivorous marsupial, from a specimen which he considered morphologically intermediate between two congeneric species, Sminthopsis murina and S. crassicaudata; his work does not unambiguously reveal that the name was proposed for an infrasubspecific entity, and accordingly constricta has subspecific rank from its original publication.

In the Heteroptera, Westhoff (1884) explicitly gave the name Pyrrhocoris apterus var. pennata to a macropterous form as such, and Wagner (1947) explicitly gave the name Stenodema trispinosum f. pallescens to freshly emerged adults as such; the names pennata and pallescens are therefore of infrasubspecific rank, and since neither was adopted for a species or subspecies before 1985 they are both unavailable.

Polinski (1929) described a terrestrial gastropod Fruticicola unidentata subtecta as a "variété (natio) n.", explicitly stating that it was only "une forme" which did not merit subspecific rank. However, Klemm (1954) adopted Trichia (Petasinaunidentata subtecta (Polinski) as the valid name of a subspecies, and the subspecific name subtecta is therefore deemed to be available from Polinski, 1929.

Richard Dawkins (2002), sumed it up nicely: "Historians of science have raised the suggestion that Wallace's version of natural selection was not quite so Darwinian as Darwin himself believed. Wallace persistently used the word 'variety' as the level of entity at which natural selection acts. You heard an example in the long passage I have just read out. And some have suggested that Wallace, unlike Darwin who clearly saw selection as choosing among individuals, was proposing what modern theorists rightly denigrate as 'group selection'. This would be true if, by 'varieties', Wallace meant geographically separated groups or races of individuals. At first I wondered about this myself. But I believe a careful reading of Wallace's paper rules it out. I think that by 'variety' Wallace meant what we would nowadays call 'genetic type', even what a modern writer might mean by a gene. I think that, to Wallace in this paper, variety meant not local race of eagles, for example, but 'that set of individual eagles whose talons were hereditarily sharper than usual.'"


Bowler, P. J. 1976. Alfred Russel Wallace's concepts of variation. Journal of the History of Medicine and Allied Sciences, 31(1): 17-29.

Dawkins, R. 2002. The reading of the Darwin-Wallace papers commemorated - in the Royal Academy of Arts. The Linnean, 18(4): 17-24. [Click HERE for a pdf]

Kottler, M. J. 1985. Charles Darwin and Alfred Russel Wallace: Two decades of debate over natural selection. In: Kohn, D. (Ed.), The Darwinian Heritage. Princeton NJ: Princeton University Press: 367-432.

Scratchpads developed and conceived by (alphabetical): Ed Baker, Katherine Bouton Alice Heaton Dimitris Koureas, Laurence Livermore, Dave Roberts, Simon Rycroft, Ben Scott, Vince Smith