Abstract
Any biological species of biparental organisms necessarily includes, and is fundamentally dependent on, sign processes between individuals. In this case, the natural category of the species is based on family resemblances (in the Wittgensteinian sense), which is why a species is not a natural kind. We describe the mechanism that generates the family resemblance. An individual recognition window and biparental reproduction almost suffice as conditions to produce species naturally. This is due to assortativity of mating which is not based on certain individual traits, but on the difference between individuals. The biosemiotic model described here explains what holds a species together. It also implies that boundaries of a species are fundamentally fuzzy, and that character displacement occurs in case of sympatry. Speciation is a special case of discretisation that is an inevitable result of any communication system in work. The biosemiotic mechanism provides the conditions and communicative restrictions for the origin and persistence of diversity in the realm of living (communicative and semiotic) systems.



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That is, no single diagnostic trait can characterise all individuals of the category. Also, common origin is not obligatory for the category; thus, polyphyly is possible.
The concept of family resemblance itself was known long before Wittgenstein; for instance, it was defined precisely by Dugald Stewart already in 1818 (see a review in Mizak 2005).
Krzeszowski (1990: 215–216) writes: „natural categories, characterized by lack of clear-cut category boundaries (fuzziness, ability to stretch), […] permeate natural languages. […] Classical phonology was incapable of handling the well-known phenomenon of phonemic overlap, which involves situations in which many non-prototypical variants of some phonemes belong to a given category because of family resemblance […].“
At the same time, we still retain the assumption that every organism belongs to some genus, familia, and other upper taxa — because (differently from the species) we do not assume that these taxa have been produced and delimited by the organisms themselves.
This is equivalent to the absence of natural selection, the latter being defined as the differential reproduction of genotypes (see also Kull 2014).
Therefore this mechanism is not the one of stabilising selection. In case of stabilising selection, the lower fitness of extreme specimen is due to their particular genotypes, not due to their difference from the average.
In many cases, however, we can observe a “double assortativity” (or multi-stage recognition), which means that the recognition of a mate has (at least) two stages — the first, which restricts the range of potential mates (a wider “window”), and the second, where the more detailed comparison (fitting) of the (usually sexual) characteristics of the mates takes place (a narrower “window”); this is obviously a factor that allows the coexistence of close species.
See also the critique of the isolationist concept of species in Paterson (1993).
Thus it is not the same as the Wallace effect, either. See Pfennig and Pfennig (2010); they mention that E. Mayr did not make a clear distinction between character displacement and character divergence.
W. Petersen would belong to Romanes’ tradition, while K. Jordan, who argued against W. Petersen, to the tradition of Wagner.
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Acknowledgments
I thank Hugh Paterson and the late John Maynard Smith for inspiring discussions, and Ene-Reet Soovik and Timo Maran for technical help. The work is related to IUT2-44.
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Kull, K. The Biosemiotic Concept of the Species. Biosemiotics 9, 61–71 (2016). https://doi.org/10.1007/s12304-016-9259-2
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DOI: https://doi.org/10.1007/s12304-016-9259-2