Abstract
We argue that C. Darwin and more recently W. Hennig worked at times under the simplifying assumption of an eternal biosphere. So motivated, we explicitly consider the consequences which follow mathematically from this assumption, and the infinite graphs it leads to. This assumption admits certain clusters of organisms which have some ideal theoretical properties of species, shining some light onto the species problem. We prove a dualization of a law of T. A. Knight and C. Darwin, and sketch a decomposition result involving the internodons of D. Kornet, J. Metz and H. Schellinx. A further goal of this paper is to respond to B. Sturmfels’ question, “Can biology lead to new theorems?”
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Notes
Unless something unnatural occurred, for example, all the living specimens of each species joining to co-parent a single sterile child.
By A3, the biosphere is countable; it can be shown that only the countable axiom of choice is needed to guarantee existence of inspecies.
This work reflects a stepwise formalization of Hennig’s internodal species: unavoidability of the implied permanency of cleavages (Kornet 1993), formal implementation (Kornet et al. 1995); lowering species’ status, because of implied short lifespans, to building blocks (internodons) (Kornet et al. 1995); remedial grouping by secondary morphological criteria into composite species (Kornet and McAllister 2005). The entire project was first informally printed as a PhD thesis (Reconstructing Species; Demarcations in Genealogical Networks, 1993, Leiden University).
To quote Kornet et al. (1995), internodons are “parts of a genealogical network of individual organisms between two successive permanent splits or between a permanent split and an extinction event.”
As candidates one may think of organisms with complex haploid/diploid life cycles such as social animals (ants, bees, termites, weevils) or the creative algae, fungi, mosses. But simpler still would be a genuine sexual network with an additional variant male type with a y′ chromosome. Say, this variant male’s sperm cells carrying the y′ chromosome always outcompete its own sperm cells carrying the x chromosome. Then, per generation we have one (non-variant) genuine xy male (generating x and y sperm cells), one (non-variant) genuine xx female (generating x egg cells only), and one variant xy′ male (generating y′ sperm cells that always outcompete its x sperm cells). In this way a variant can only co-parent (with the genuine female) an xy′ variant son.
Thus, the undirdescendants of u are precisely the members of the gross dynasty of u (minus u and its same-exact-age peers), in the language of Kornet et al. (1995).
To be precise, we are assuming that every individual has the SD property, which equates the equivalence relations INT and INTSD, as well as the sets \({{\mathbb{DYN}}}\) and \({\mathbb{GDYN}}\), of the 1995 paper.
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Acknowledgments
We thank Timothy J. Carlson, Bora Bosna, Mike Fenwick, and especially the referees for much productive feedback and discussion.
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Proof of Theorem 12
Proof of Theorem 12
In this appendix we will prove Theorem 12 (hereafter we assume its hypotheses). First we review the definitions in Kornet et al. (1995). As we state them they are not equivalent; they are altered to fit the hypotheses of Theorem 12, allowing us to simplify them.Footnote 8
Definition 5
For any \(u\in G\), write \({{\mathbb{DYN}}(u)}\) for the set \(\{x\in G : u({\bf PC}_{\geq u})x\}\) and write ≧ (u) for \(\{x\in G : t(x)>t(u)\}. \) If \(u,v\in G\) and t(u) < t(v), define
If t(v) < t(u) then u INT v :⇔ v INT u, and if t(v) = t(u) then u = v by our assumption that no individuals are born simultaneously, and we define u INT u.
The main theorem of Kornet et al. (1995) implies INT is an equivalence relation.
Definition 6
An internodon is an INT-equivalence class.
We could now dive directly into the proof of Theorem 12 but we prefer to factor out a sufficient condition which might be useful in the future for proving that other species notions decompose into internodons (this condition, too, will be generalized in future work).
Proposition 13
(Sufficient conditions for internodons-decomposition) (Assuming the hypotheses of Theorem 12.) Let \(S\subseteq V. \) If S is undirancestrally closed, and for each \(u\in S\) and \({t\in {\mathbb{R}}}\) there is some \(u'\in S\) born after t with \(u({\bf PC}_{\geq u})\) u′; then S is a union of internodons.
Proof
It suffices to let \(u\in S, v\in G\), and show that if u INT v then \(v\in S\).
Case 1: v is born before u. Since v INT u, in particular \(v({\bf PC}_{\geq v})\) u. Thus v is an undirancestor of u, putting \(v\in S\) by undirancestral closure.
Case 2: v is born after u. Since u INT v, by definition of INT we have \(u({\bf PC}_{\geq u})\) v and (*) for every r such that \(u({\bf PC}_{\geq u})\) r and t(r) ≤ t(v), we have \({{\mathbb{DYN}}(u)\,\cap \geqq(r)={\mathbb{DYN}}(r)}\).
By the proposition’s hypothesis, there is some \(u'\in S\), born after v, such that \(u({\bf PC}_{\geq u})\) u′. Letting r = v in (*), we see (since \(u({\bf PC}_{\geq u})\) v and t(v) ≤ t(v)) that \({{\mathbb{DYN}}(u)\cap\geqq(v)={\mathbb{DYN}}(v). }\) Since \({u'\in {\mathbb{DYN}}(u)\cap\geqq(v)}\), this shows \({u'\in{\mathbb{DYN}}(v)}\), that is, \(v({\bf PC}_{\geq v})\) u′. Thus v is an undirancestor of u′, so \(v\in S\) by undirancestral closure of S. \(\square\)
Proof of Theorem 12
Let S be an inspecies (type II). The first hypothesis of Proposition 13 holds since S is undirancestrally closed by definition. The second hypothesis of Proposition 13 is immediate by Proposition 6. \(\square\)
The reader will have noticed that despite imposing such onerous hypotheses on Theorem 12, we barely seem to have actually used those hypotheses. Their key use was in simplifying the definition of internodons.
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Alexander, S.A. Infinite Graphs in Systematic Biology, with an Application to the Species Problem. Acta Biotheor 61, 181–201 (2013). https://doi.org/10.1007/s10441-012-9168-y
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DOI: https://doi.org/10.1007/s10441-012-9168-y