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Interbreeding
Defining species by the criteria of not being able to breed is problematic:

  • The concept loses its meaning in phylogenetic prospective, since closely related species were able to breed between themselves in past, during the speciation event (an "event" that often lasted hundreds or thousands of generations)
  • The definition is problematic for simpler life forms, such as bacteria exchanging genetic material, or viruses where different species from the same family may differ only quantitatively (e.g., HIV vs. SIV).

(Excellent exposition of this view on the difficulty/impossibility of defining species is presented in this answer, as well as in the answer by @SPr below.)

Emergence
On the other hand, species are not reducible to merely accumulation of genetic variation. When looked from more general prospective (e.g., here), species are an example of an emergent phenomenon (the same inability to interbreed is a qalitative rather than quantitative difference.) We thus should be able to define a critical boundary between what is the same species and not, although this boundary does not necessarily lie in time domain. Inability to interbreed is a vague definition, since it may be due to very different reasons: phisiological, genetic, assortative mating, etc.

Operational concepts
In some domains we could choose to define species based on a criteria of convenience. E.g., considering an ecological community one may define as species the organisms that cannot interbreed, but also those that do not share the same habitat. While such an operational definition may work well for practical studies, it is not unique and subject to change from one study to another.

Are species an emergent property or an ensemble of quantitative differences? Can we give a robust definition of species/speciation? Can we define a domain of applicability of such a definition? (e.g., defining the forms of life for which species can be defined)

Background
When speaking of emergent properties, I tried to skip some techncialities that led me to this question, which is why this part of my question may appears somewhat vague. So I would like to add more details here.

Phil Anderson in his article Broken symmetry, emergent properties, dissipative structures, life: are they related? formulates emergence in mathematically rigorous terms as a kind of phase transition in a non-equilibrium system. Phase transitions are characterized by summetry breaking, i.e., appearance of a completely new property in a system. What I termed previously robustness is, in Anderson's language, called phase rigidity.

The question is thus: whether speciation is a kind of such phase transition (as claimed, e.g., in the Melamede's article, already cited earlier) OR whether it is merely incremental accumulation of changes. (However, in such a formulation question is unlikely to be answered either in physics or biology communities.)

For more context: in his article Anderson criticizes Ilya Prigogine's school on a few technical points, but it is Prigogine and his co-workers, who are at the origin of such thinking. In particular, they have multiple publications on the subject of such disspative structures, and non-equilibrum thermodynamics more generally. Many of their articles are available by googling.

Update
After some discussions in the comments: the crux of the question is whether (or tow hat extent) speciation occurs via abrupt or incremental changes.

  • Organisms differing by abrupt acquisitions, such as presence of a nuclear membrane or an extra chromosome, are unambiguously assigned to different species (and cannot breed or reproduce viable offspring).
  • On the other hand, most ambiguous cases seem to be those where the "species" differ only by a incremental changes (such as the number of nucleotide substitutions/insertions/deletions).

Ring species could be a possible test case: if the species at the beginning and the end of the ring cannot breed, do they always differ only by incremental changes? If one species differed by a number of chromosomes, it likely wouldn't be able to breed with any of the others.

Update 2
Here is a useful answer: species can be defined from many different viewpoints.

Roger V.
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  • Ew "closely related species were able to breed between themselves in past", it's also the case in the present. See for instance the development of the "coywolf" in eastern North America: https://en.wikipedia.org/wiki/Coywolf – jamesqf May 05 '21 at 18:50
  • @jamesqf sure, horse and donkey can breed, but two breeds of dogs sometimes can't. This underscores the insufficiency of this definition. But the point I was making was dufferent - if you have suggestion of more precise formulation, I am listening. – Roger V. May 05 '21 at 19:52
  • @Vadim often the difference with dogs is physical rather than "can't". Imagine a female chihuahua trying to breed with a male great dane, or vice versa. I don't think the genitalia would fit, nor would the pups in the first instance. – bob1 May 05 '21 at 20:35
  • @bob1 physically can't :) the point is that any two nearest relatives could cross at some "point". It is the chimpansee-girilla-human problem - which of the three are closest relatives? – Roger V. May 05 '21 at 21:21
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    @Vadim my point was that they can if you take the sperm from one and use it to fertilize the other, it's just they they don't (not for lack of trying I am told) because things don't fit, or can't reach... They are definitely not separate species. Horse and donkey can interbreed but offspring are usually sterile - mule and hinny respectively, so definite separate species. But the boundaries between other organisms aren't so clear. – bob1 May 05 '21 at 21:40
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    @bob1: Yes, just about everyone regards dog breeds as varieties of a single species, yet the "don't fit" problems ensure that some breed pairings don't produce offspring that are viable without human assistance. (Indeed, I've read that some show breeds like bulldogs need human assistance.) OTOH, dogs, wolves, and coyotes are regarded as different species, yet can interbreed and produce viable offspring. – jamesqf May 06 '21 at 04:27
  • @bob1 my understanding is that horse and donkey have different number of chromosomes, whereas dogs differ by a number of point mutation (nucleotide substitutions/insertions/deletions). To retake the main theme: this is an example of abrupt vs. incremental changes. Are any known ring species more than dogs in this respect? – Roger V. May 06 '21 at 06:22
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    Another example of different species that freely interbreed are trout. Particularly within the US Great Basin, where the drier climate after the Pleistocene created a lot of isolated stream/lake systems, which often evolved their own trout species. Then eastern brook & rainbow trout were introduced, and freely bred with the native species, and with each other, producing viable hybrids such as the tiger trout: http://www.ndow.org/Species/Fish/Tiger_Trout/ – jamesqf May 06 '21 at 16:53
  • @jamesqf by what criteria are these "species" are then judged to be different species? – Roger V. May 06 '21 at 17:04
  • @Vadim: 1) Taxonomists and people who discover (or want to have discovered) new species give them names. In that context, see the arguments between "lumpers" (who think the minor differences between some of those species should better be called varieties) and "splitters" (who think minor differences justify calling them new species). 2) In the US, the EPA, Forest Service, various environmental groups &c who use the "splitter" side of the argument as justification for policies protecting the supposed species. – jamesqf May 07 '21 at 00:26

1 Answers1

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Can we give a robust definition of species?

  • No.

Species constantly evolve, diverge, converge, interbreed, and mix and shuffle and trade and spread genes. To draw a box at any particular point in time around a population or lineage, and to say, "all contained within are members of such-and-such species" fails to appreciate that this would only work for that particular case, and arbitrarily so; it is obviously futile to have a sound microtaxonomy that would be generalizable to all animals, or microbes, or plants, or viruses for that matter. What consensus there might be for (e.g.) beetles is a poor yardstick for judging whether two alike bacteria ought to be lumped together as a species. Also, life, genes, individuals and populations are dynamic over time, so whatever box we draw will by definition require adjustment, and you can see that this adjustment will have to be arbitrary in many respects. This is an old problem, and it is more philosophical and ever-debatable, rather than something to be settled by empirical science triumphantly. From Nicholson's 1872 A manual of zoology (two centuries ago now):

"No term is more difficult to define than "species," and on no point are zoologists more divided than as to what should be understood by this word."

And to echo all that has been said from Piglucci (2003) Bioessays:

"First, the species problem is not primarily an empirical one, but it is rather fraught with philosophical questions that require — but cannot be settled by — empirical evidence."

Which also summarizes a few major biological definitions, some of which do not align:

Table of species concepts

And also a few philosophical issues that are worth considering when discussing the species problem:

enter image description here

This is not to say that conventions don't exist, or that there is a lack of an operational and specific definition of a species in each respective field; however, there is no overarching definition that applies robustly, hence the answer at the top!

David
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S Pr
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  • Also, search around on this SE for similar Q&As. I find no duplicates but some similar ones. – S Pr May 04 '21 at 12:00
  • Thank you for the answer. It provides many useful insights. However, what I really would like to understand is when quantitative differences between two organisms transform into qualitative ones... or whether such a transformation never takes place and the differences are always quantitative. – Roger V. May 04 '21 at 13:36
  • @Vadim Can you not think of species that have qualitative differences between them? Even individuals of the same species can have qualitative differences. Maybe you have a different idea of what that means? – Bryan Krause May 04 '21 at 15:42
  • @BryanKrause Good point. Perhaps, it is more precise to ask what quality makes organisms belong to the same species or not. I think not being able to breed is a wrong quality, but the idea is correct: the quality is either there or not. If we can show that such a quality cannot exist, it is in itself a strong claim. – Roger V. May 04 '21 at 16:40
  • @Vadim I think SPr's answer and some other Q&A here make it pretty clear there is no single definition, and can't be one that is agreed upon. – Bryan Krause May 04 '21 at 16:45
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    Remi's answer here is excellent: https://biology.stackexchange.com/a/39669/27148 – Bryan Krause May 04 '21 at 16:46
  • @BryanKrause the linked answer is indeed very complete. However, it addresses only one line of thinking - the one that I outlined in the first part of my question. And this line of thinking, it seems, is in contradiction with how we think about the emergence of life, which impliues a series of abrupt qualitative changes. – Roger V. May 04 '21 at 17:00
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    @Vadim What does "emergence of life" have to do with "species"? Who is the "we" that implies abrupt qualitative change? I think Remi's answer there makes clear that "species" is just a classification we draw for convenience when what we are really dealing with is "lineage" which can't be separated except by (relatively) arbitrary means. – Bryan Krause May 04 '21 at 17:02
  • @BryanKrause I cited Phil Anderson and Ilya Prigogine in my question. Although neither might have mentioned species - this specific question is based on the quoted Melamede's article, where speciation is listed as one of the abrupt transitions between the emergence of cell and the evolution of body systems. – Roger V. May 04 '21 at 17:09
  • @Vadim I can't access that article and don't see any quote from it. I'm guessing something is getting lost in the details. I would not describe speciation as "abrupt" except in special cases or using special definitions of species (or a geological meaning of "abrupt", perhaps). – Bryan Krause May 04 '21 at 17:15
  • @BryanKrause I suppose that it can be found for download by googling - (but I wouldn't post link for downloading copyrighted material). Anyhow, I see your point of view, and I tend to agree with it... but it still seems as a grey area to me. – Roger V. May 04 '21 at 17:20
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    @Vadim Dug a bit further and did find the article.. so, okay, this seems to me like a lot of word salad and not a lot of substance, at least with a casual glance. The author also seems to be mixing the words "species" referring to biological species and "species" in the context of "chemical species", the more generic "kind/sort", which has little relationship to the concept in biology. I wouldn't recommend taking it too seriously. In any event, I don't see any reference to anything "abrupt", if there's a particular quote that is relevant you should include it in your question. – Bryan Krause May 04 '21 at 17:26
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    @Vadim - check out ring species as an example of the difficulties in determining what is a species and where. The problem is that you are approaching it in a computational manner, where things have exact definitions. Species do not have exact definitions precisely for the reasons S Pr outlined. Biology is messy! An example of the difficulty is cancer research, where cell lines evolve in the lab so that results from one lab are not reproducible in another, even though both are working on the same cell line. – bob1 May 05 '21 at 20:43
  • @bob1 I am not looking for exact universal definition of species. The point of the question is whether species a result of small incremental changes or an abrupt jump - like appearance of a cellular membrane. Depending on which corner you look from (taxonomy or abiogenesis) the answer is different (in my view). – Roger V. May 05 '21 at 21:19
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    @Vadim - both is the answer. Abrupt jumps are seen in some circumstances, incremental in others. – bob1 May 05 '21 at 21:44
  • @Vadim You say in my view, species definition depends whether one considers it from a taxonomical vs. abiogenetic point of view. I think we would all disagree with you. Defining species has exactly the same problems regardless of whether you believe that at some point (i) life was designed to evolve, (ii) created and subsequently evolved, or (iii) evolved out of inorganic material. Evolution exists and that makes defining species boundaries arbitrary, since all life, populations and lineages are in constant flux over time. That's it. See, whether it's abrupt or not is irrelevant. – S Pr May 07 '21 at 11:33
  • @SPr Perhaps abiogenic is too general a word for the veiw that I expressed, but I think I have described clearly enough the two contradictory viewpoints in my question. We have a good reason to belive that there is abrupt transition between being alive or not, between being cellular organism or not, between having a nucleus or not, etc. Any division in species that is not grounded in evolution is indeed arbitrary. And I don't see any problem with populations and lineages being in flux - so the species change, some disappear, some split into more species. – Roger V. May 07 '21 at 11:55