Skip to main content
Log in

Infinite Graphs in Systematic Biology, with an Application to the Species Problem

  • Regular Article
  • Published:
Acta Biotheoretica Aims and scope Submit manuscript

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?”

This is a preview of subscription content, log in via an institution to check access.

Access this article

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5
Fig. 6
Fig. 7

Similar content being viewed by others

Notes

  1. Baum and Shaw (1995, p. 294) and Velasco (2008, p. 868) point out the problem of big cleavages and small cleavages, and the resulting vagueness. This vagueness disappears under infinitary assumptions.

  2. Unless something unnatural occurred, for example, all the living specimens of each species joining to co-parent a single sterile child.

  3. By A3, the biosphere is countable; it can be shown that only the countable axiom of choice is needed to guarantee existence of inspecies.

  4. 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).

  5. 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.”

  6. 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  xymale (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.

  7. 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).

  8. 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.

References

  • Alexander S (preprint) Biologically unavoidable sequences. Submitted. http://arxiv.org/abs/1212.0186

  • Baum DA (2009) Species as ranked taxa. Syst Biol 58:74–86

    Article  Google Scholar 

  • Baum DA, Shaw KL (1995) Genealogical perspectives on the species problem. In: Hoch PC, Stephenson AC (eds) Experimental and molecular approaches to plant biosystematics. Missouri Botanical Garden, St Louis, pp 289–303

    Google Scholar 

  • Cohen J (2004) Mathematics is biology’s next microscope, only better; biology Is mathematics’ next physics, only better. PLoS Biol 2(12). Available: http://www.plosbiology.org/article/info

  • Darwin C (1872) On the origin of species, 6th edn. John Murray, London

  • Darwin F (1898) The Knight–Darwin law. Nature 58:630–632

    Article  Google Scholar 

  • Diamond J (1997) Guns, germs, and steel. W. W. Norton & Company, New York

    Google Scholar 

  • Dress A, Moulton V, Steel M, Wu T (2010) Species, clusters and the ‘Tree of life’: a graph-theoretic perspective. J Theor Biol 265: 535–542. Available: http://www.math.canterbury.ac.nz/m.steel/Non_UC/files/research/species_jtb.pdf

    Google Scholar 

  • Gowers T (2008) How to use Zorn’s lemma. Available: http://gowers.wordpress.com/2008/08/12/how-to-use-zorns-lemma/

  • Hennig W (1966, reprinted 1979). Phylogenetic systematics. Davis D, Zangerl R, translators. University of Illinois Press, Urbana

  • Hyde D (2011) Sorites Paradox. In: Zalta E (ed) The stanford encyclopedia of philosophy (Winter 2011 Edition). Available: http://plato.stanford.edu/archives/win2011/entries/sorites-paradox/

  • Johnston N (2010) The B36/S125 “2 × 2” Life-like cellular automaton. In: Adamatzky A (ed) Game of life cellular automata. Springer, New York. Preprint available: http://njohns01home.webfactional.com/wp-content/uploads/2010/01/2x2.pdf

  • Kőnig D (1936) Theorie der Endlichen und Unendlichen Graphen. Akademische Verlagsgesellschaft, Leipzig

    Google Scholar 

  • Kornet D (1993) Permanent splits as speciation events: a formal reconstruction of the internodal species concept. J Theor Biol 164:407–435

    Article  Google Scholar 

  • Kornet D, Metz J, Schellinx H (1995) Internodons as equivalence classes in genealogical networks: building-blocks for a rigorous species concept. J Math Biol 34:110–122

    Article  Google Scholar 

  • Kornet D, McAllister J (2005) The composite species concept: a rigorous basis for cladistic practice. In: Reydon T, Hemerik L (eds) Current themes in theoretical biology: a Dutch perspective. Springer, Dordrecht, pp 95–127

    Chapter  Google Scholar 

  • Nixon K, Wheeler Q (1990) Another way of looking at the species problem: a reply to de Queiroz and Donoghue. Cladistics 6:77–81

    Article  Google Scholar 

  • de Queiroz K (2007) Species concepts and species delimitation. Syst Biol 56:879–886

    Article  Google Scholar 

  • de Queiroz K, Donoghue J (1988) Phylogenetic systematics and the species problem. Cladistics 4:317–338

    Article  Google Scholar 

  • Steel M (2007) Tools to construct and study big trees: a mathematical perspective. In: Hodkinson T, Parenell J, Waldren S (eds) Reconstructing the tree of life: taxonomy and systematics of species rich Taxa. CRC Press (Taylor and Francis), New York, pp 97–112. Preprint available: http://www.math.canterbury.ac.nz/m.steel/Non_UC/files/research/tools.pdf

  • Sturmfels B (2005) Can biology lead to new theorems? Annual report of the Clay Mathematics Institute. Available: http://math.berkeley.edu/bernd/ClayBiology.pdf

  • Velasco J (2008) The internodal species concept: a response to ‘The tree, the network, and the species’. Biol J Linnean Soc 93:865–869

    Article  Google Scholar 

  • Zorn M (1935) A remark on method in transfinite algebra. Bull Am Math Soc 41:667–670

    Article  Google Scholar 

Download references

Acknowledgments

We thank Timothy J. Carlson, Bora Bosna, Mike Fenwick, and especially the referees for much productive feedback and discussion.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Samuel A. Alexander.

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

$$ u{\bf INT}v :\Leftrightarrow u({\bf PC}_{\geq u})v \wedge \forall r[\{u({\bf PC}_{\geq u})r \wedge \left( t(r)\leq t(v) \right)\} \Rightarrow ({{\mathbb{DYN}}}(u)\cap \geqq(r) = {{\mathbb{DYN}}}(r))]. $$

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.

Rights and permissions

Reprints and permissions

About this article

Cite this article

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

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s10441-012-9168-y

Keywords

Navigation