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. 2025 Aug 18;87(9):132. doi: 10.1007/s11538-025-01506-1

Group-based phylogenetic models on 3-sunlet networks

Shelby Cox 1, Elizabeth Gross 2, Samuel Martin 3,4,
PMCID: PMC12358336  PMID: 40820189

Abstract

Phylogenetic networks describe the evolution of a set of taxa for which reticulate events have occurred at some point in their evolutionary history. Of particular interest is when the evolutionary history between a set of just three taxa has a reticulate event. In molecular phylogenetics, substitution models can model the process of evolution at the genetic level, and the case of three taxa with a reticulate event can be modeled using a substitution model on a semi-directed graph called a 3-sunlet. We investigate a class of substitution models called group-based phylogenetic models on 3-sunlet networks. In particular, we investigate the discrete geometry of the parameter space and how this relates to the dimension of the phylogenetic variety associated to the model. This enables us to give a dimension formula for this variety for general group-based models when the order of the group is odd.

Introduction

Phylogenetic networks are directed graphs that describe the evolution of a set of taxa for which reticulate events have occurred. Such events, which include, horizontal gene transfer and hybridization, are increasingly being discovered to have occurred between taxa, and the development of methods to reconstruct phylogenetic networks from molecular sequence data is an active area of research. It is therefore important that phylogenetic networks and the models that are placed on them are well understood.

In this work, we focus on phylogenetic network-based substitution models. These are latent-variable Markov models where the state space is a set of biological molecules (usually the four nucleic acids {A,G,C,T}), and along each edge in the network, a transition matrix gives the probabilities of each possible substitution occurring along that edge (see Gross and Long (2018), Nakhleh (2011) for further details). In this work we focus on a family of Markov models called group-based models, so called because the state space of the Markov process is identified with a finite abelian group. In molecular phylogenetics, there are several nucleotide substitution models that are group-based models, such as the Jukes-Cantor (JC) model, the Kimura 2-parameter (K2P) model, and the Kimura 3-parameter (K3P) model. In all these cases, the state space of the four nucleic acids {A,G,C,T} is identified with the Klein-four group Z/2Z×Z/2Z.

For Markov models on phylogenetic networks, the joint probabilities of observing particular patterns at the leaves of the network have polynomial parameterizations in terms of the numerical parameters of the model, i.e., the substitution rates along each edge and reticulation edge parameters. This makes them amenable to algebraic study, and, in particular, the space of all possible joint probabilities at the leaves is the intersection of an algebraic variety with the probability simplex.

In this work we are concerned with the dimension of the variety associated to a phylogenetic network and group-based model. The dimension of such varieties, in this case, t-varieties, is an interesting geometric question in its own right, but also has applications to identifiability and phylogenetic network inference. While Gross et al. (2024) establishes the dimension for most group-based phylogenetic network models, the most elusive has been when the network contains 3-cycles. In this work, we focus on the smallest phylogenetic network containing a 3-cycle, which is called a 3-sunlet. While these networks remain the most elusive to understand mathematically, they are perhaps the most important cycles to understand biologically. Indeed, it is assumed that 3-cycles are the most common cycle motif in true phylogenetic networks, since they indicate hybridization or lateral gene transfer between two very closely related taxa, whereas, larger cycles would indicate such reticulation events between less closely related taxa, which, in many cases, is assumed to be rare. Understanding the dimensions of 3-sunlets can help us establish the statistical property of identifiability, as demonstrated in Proposition 3.9. It can also help us understand how 3-sunlet models are geometrically embedded within larger sunlet models, helping interpret residuals when using algebraic methods as in Barton et al. (2022), Martin et al. (2023) or determining the most appropriate penalty term when using Bayesian methods.

For group-based models on phylogenetic trees, after a transformation, the parameterization of the model is monomial Evans and Speed (1993), Székely et al. (1993), and thus the corresponding variety is a toric variety. These models have been well studied (see e.g. Sturmfels and Sullivant (2005)). The parameterization of the model on a sunlet network has a combinatorial interpretation, and, after the same transformation, is described by binomials. Here we look at the 3-cycle case in depth, and establish a dimension result for group-based models for groups of odd order.

Theorem 1.1

Let G be a finite abelian group of odd order +15, and let N be the 3-sunlet network under the general group-based model given by G, with corresponding phylogenetic network variety VNG. Then the affine dimension of VNG is given by

dimVNG=5+1.

We call the quantity 5+1 the expected dimension of the model Gross et al. (2024), and Theorem 1.1 agrees with the conjecture given in Gross et al. (2024). As we will see in Section 4, we believe that this conjecture holds for all finite abelian groups (of order at least 5), but as we discuss in detail in Section 5, the proof strategy that we use here is not easily modified for groups of even order, and thus those cases remain open.

Whilst in analysis of DNA sequences the state space of our models is the four nucleic acids and therefore has order four, odd-order state spaces are common in other analyses. For example, in codon models such as the Goldman-Yang model Goldman and Yang (1994), triplets of nucleotides, called codons, code for amino acids, and are the states of the model. There are 64 possible nucleotide triplets, but often only 61 are used, since three are ‘stop’, codons, which are not modeled. In amino acid models, the 20 amino acids of the standard genetic code are often recoded to a smaller set. One example is Dayhoff six-state recoding Dayhoff et al. (1978), where chemically related amino acids are grouped together to form six states. This grouping is based on the substitution rates in the PAM250 matrix, with amino acids with high substitution rates between them grouped together. However, substitution rates between amino acids can depend on factors such as secondary structure and functional domain Goldman et al. (1998), and so methods have been developed to recode based on replacement patterns from, for example, domain-specific databases Kosio et al. (2004). This introduces the possibility of substitution models with odd-order state spaces.

The paper is organized as follows. In Section 2, we describe phylogenetic networks and the paramaterization map for 3-sunlet networks for the general group-based model. We also outline the tropicalization method from Draisma (2008) that we use to determine a lower bound for the dimension. The method is rooted in tropical geometry and leads to hyperplane arrangements on spaces of weight vectors. We close Section 2 with observations about the chambers of these hyperplane arrangements for 3-sunlet networks. In Section 3, we prove the main theorem of the paper (Theorem 1.1), which gives a formula for the dimension for 3-sunlet networks under general group-based models of odd order greater than or equal to 5. We end the section with a partial identifiability result (Proposition 3.9) for general group-based models of odd order. In Section 4, we investigate the dimension for small finite abelian groups (both even and odd) through computational experiments. In particular, we explore chambers of hyperplane arrangements to highlight the difficulty in finding appropriate weight vectors that can be used to establish dimensions of 3-sunlets. Section 5 closes the paper with a discussion about the challenges involved in understanding 3-sunlets, and more generally, networks with 3-cycles.

Background

A (rooted binary) phylogenetic network is a rooted, acyclic, directed graph where each non-root internal vertex has in-degree one and out-degree two, or in-degree two and out-degree one. We refer to the internal vertices with in-degree one as tree vertices and the internal vertices with in-degree two as reticulation vertices. The leaves of the phylogenetic network (the vertices of in-degree 1 and out-degree 0) are labelled by a set of taxa, for which we will always use the set of the first n positive integers [n]={1,,n}. The two edges directed into a reticulation vertex are called reticulation edges. A phylogenetic network N is said to be level-1 if, in the undirected skeleton of N, no two cycles share an edge. For an example of a level-1 phylogenetic network, see Figure 1. A semi-directed phylogenetic network is a mixed graph that is obtained from a phylogenetic network by suppressing the root vertex and un-directing all non-reticulation edges. Semi-directed networks generalize the notion of unrooted trees, and, for group-based models, if two phylogenetic networks have the same underlying semi-directed topology, then their corresponding varieties are also equal (Gross et al. 2024, Lemma 2.2). Since we are concerned with the dimensions of the corresponding varieties, we will only consider semi-directed phylogenetic networks.

Fig. 1.

Fig. 1

A rooted, level-1 phylogenetic network. This network contains a single 3-cycle and a single 4-cycle. Reticulation edges are drawn with dashed lines

The fundamental building blocks of level-1 semi-directed phylogenetic networks are unrooted trees and k-sunlet networks, which are the minimal semi-directed phylogenetic networks containing a k-cycle. In this paper, we focus on 3-sunlet networks, which are the minimal semi-directed phylogenetic networks containing a triangle (i.e., a 3-cycle). A 3-sunlet can be obtained from the phylogenetic network in Figure 1 by restricting the network to the leaves labelled by taxa 1, 3, and 6 and suppressing vertices of degree 2 (restriction is discussed in more detail towards the end of Section 3). Phylogenetic networks with triangles are thought to be among the most common phylogenetic networks, because hybridization usually occurs between closely related species. Despite this, 3-sunlet networks are the least understood of the sunlet networks.

We place a group-based model of evolution on a level-1 semi-directed phylogenetic network N by arbitrarily assigning direction to all undirected edges (i.e., non-reticulation edges), and choosing a finite abelian group G and a subgroup B of the automorphism group of G, denoted Aut(G). We note that arbitrarily assigning directions to all undirected edges results in the same variety as rooting the semi-directed network even if the chosen edge directions are not consistent with any placement of a root vertex (see the proof of Lemma 2.2 in Gross et al. (2024)). The group G is identified with the state space of the model, and the group B encodes additional constraints that the transition matrices must adhere to. When we choose B={id} we call the model the general group-based model for G. For example, the Kimura 3-parameter (K3P) model is the general group-based model for G=Z/2Z×Z/2Z. The Jukes-Cantor (JC) model is the group-based model with G=Z/2Z×Z/2Z and B=AutG, which we identify with S3, the symmetric group of order 3. In between these two we have the Kimura 2-parameter model (K2P), where B is a subgroup isomorphic to S2. Group-based models have the desirable property that for any phylogenetic tree there exists a Fourier transformation that transforms expressions for the marginal probabilities of observations at the leaves into monomial expressions (see e.g., (Sullivant 2018, Chapter 15) for an overview). For level-1 phylogenetic networks, that same transformation significantly simplifies the expressions for the marginal probabilities, although they are not monomial (Gross and Long 2018, Prop 4.2).

We are interested in identifying the semi-directed phylogenetic network from observed data on the leaves. As noted above, for group-based models, the root of network is not identifiable. For certain group-based models, identifiability results are known (see e.g. Gross and Long (2018), Gross et al. (2021), Hollering and Sullivant (2021), Cummings et al. (2023)), but a general result for all group-based models has yet to be determined. Understanding the dimension of the variety associated to a phylogenetic network and model can assist in determining identifiability. A step in this direction was taken in Gross et al. (2024), and some identifiability results were obtained for arbitrary group-based models. Here, one limiting factor was being unable to determine the dimension of the varieties associated the 3-sunlet network.

The 3-sunlet networks

The 3-sunlet is the semi-directed network topology of a simple 3-leaf phylogenetic network with a single cycle. It poses a particular problem to phylogeneticists, because under the most commonly used 4-state group-based models (JC, K2P, and K3P), the reticulation vertex is not identifiable from data at the leaves of the network (see e.g. (Gross et al. 2021, Lemma 1)). Thus many of the identifiability results obtained for these models require the phylogenetic networks to be ‘triangle free’Gross et al. (2021).

Mathematically, the 3-sunlet is a semi-directed graph whose skeleton consists of a single 3-cycle and one leaf vertex adjacent to each vertex in the cycle. One of the vertices in the cycle is the reticulation vertex, and the two cycle edges adjacent to this vertex are reticulation edges. The reticulation edges are the only directed edges and they are directed towards the reticulation vertex (see Figure 2 for an example). By removing either of the edges e6 or e5 in Figure 2 and undirecting the remaining edge, we obtain an unrooted phylogenetic tree (with a vertex of degree 2), which we denote by T1 and T2 respectively.

Fig. 2.

Fig. 2

(Left) The semi-directed network topology of the 3-sunlet network with taxa labels 1,2, and 3. (Right) A directed 3-sunlet network

In order to simplify our exposition, we begin with the phylogenetic network parameterization in the transformed coordinates. Readers interested in the derivation of this parameterization from the substitution model can consult (Gross et al. 2024, Section 2) for a full explanation. In order to specify the parameterization we must direct the undirected edges of the semi-directed network topology. By (Gross et al. 2024, Lemma 2.2) we may arbitraily choose these directions, and so for the remainder of this work we denote by N the directed 3-sunlet network in Figure 2 (right).

Let G be a finite abelian group and let B be a subgroup of AutG. Let B·G be the set of B-orbits of G, and define +1:=|B·G|. Note that when B={id} we have +1=|G|. A consistent leaf-labelling is a triple g=(g1,g2,g3)G3 satisfying g1+g2+g3=0. For a fixed G there are exactly |G|2 consistent leaf-labellings. We give C6(+1) a basis indexed by B-orbits and edges of N, and denote the basis element corresponding to the B-orbit [g] and edge ei as Eig. We give C2 a basis indexed by consistent leaf-labellings g=(g1,g2,g3). Then the parameterization map ϕN(G,B) (in transformed coordinates) is given by

ϕN(G,B):C6(+1)C2,(ϕN(G,B)(w))g=w11w22w33w4g1+g2w51+w11w22w33w42w61=m1(g)+m2(g).

where wig is the coefficient of Eig in w. Here, the first term m1(g):=w11w22w33w4g1+g2w51 comes from the phylogenetic tree T1 (obtained from N be removing the edge e6). Each superscript gi is given by the edge-labelling of the edge ei in the right hand diagram in Figure 2 (see Gross et al. (2024) for further details). Similarly, the second term m2(g):=w11w22w33w42w61 comes from the phylogenetic tree T2 (obtained from N by removing the edge e5). The phylogenetic variety of N and (GB) is defined as the Zariski closure of the image of ϕN(G,B), denoted

VN(G,B)=imϕN(G,B)¯,

and this is the object that we study. Since the map ϕN(G,B) is homogeneous, the variety VN(G,B) is a projective variety. However, we will mostly remain in affine space and consider the affine cone.

Observe that the map ϕN(G,B) is a morphism of affine varieties. It has comorphism given by

ψN(G,B):C[qg|g1+g2+g3=0]C[aig|i=1,,6,andgB·G],qga11a22a33a4g1+g2a51+a11a22a33a42a61,

somewhere we think of the coordinate ring of C2 as being generated by variables qg for consistent leaf-labellings g=(g1,g2,g3), and we think of the coordinate ring of C6(+1) as being generated by variables aig for i=1,,6 and gB·G. Under this definition, the vanishing ideal of VN(G,B), which we denote IN(G,B), is given by kerψN(G,B). This ideal, and in particular, its generating sets, are an important object of study in mathematical phylogenetics as they can be used for model selection Barton et al. (2022) Cummings et al. (2024) Martin et al. (2023) and to establish identifiability Gross and Long (2018) Hollering and Sullivant (2021) Gross et al. (2021) Cummings et al. (2023). While some polynomials are known for some group-based models such as CFN Cummings et al. (2024) Cummings et al. (2023), and for certain networks, such as the 4-sunlet, polynomials have been calculated for the JC, K2P Martin et al. (2023), and K3P Cummings and Hollering (2026) models, generating sets of IN(G,B) are not known in general.

Determining Dimension

In Section 3, we give a dimension result for the 3-sunlet network and the general group-based model for groups of odd order by following the approach taken in Gross et al. (2024). Here, we introduce the concepts and objects we need. First, for a level-1 phylogenetic network N and group-based model (GB), the variety VN(G,B) is defined as the Zariski closure of the image of a polynomial map (as defined above for the 3-sunlet). Considering the number of free parameters in the domain of the map gives us an upper bound on the dimension.

Lemma 2.1

(Gross et al. 2024, Proposition 4.2) Let N be the 3-sunlet network, G a finite abelian group and B a subgroup of Aut(G) with |G·B|=+1. Then

dimVN(G,B)5+1.

As a result of Lemma 2.1, to prove Theorem 1.1, it is sufficient for us to give a lower bound on the dimension. We do this by exhibiting a Jacobian matrix of sufficient rank of the tropicalization of the parameterization.

Let ϕ=ϕN(G,B):C6(+1)C2 be the parameterization map of the 3-sunlet network under the general group-based model for an abelian group G, and for a consistent leaf-labelling g=(g1,g2,g3), let ϕg be the component of ϕ mapping onto the g-coordinate of C2. Since ϕ is a polynomial map, each ϕg is a polynomial, and we can define

Trop(ϕg):R6(+1)RλminαMλ,α,

where MZ6(+1) denotes the set of exponent vectors corresponding to the monomials in the polynomial expression for ϕg. Then Trop(ϕ):R6(+1)R2 is the map with components Trop(ϕg). Now for λR6(+1) at which Trop(ϕ) is differentiable there exists a matrix Aλ such that Trop(ϕ)(μ)=AλTμ for all μ in an open neighbourhood of λ (in fact AλT is the Jacobian of Trop(ϕ) at λ). Then we have a lower bound on the dimension of the affine variety VNG given by

dimVNGmaxλR6(+1)rankRAλ.

This is a specific case of Corollary 2.3 in Draisma (2008). For full details we recommend the reader consult Draisma (2008) and (Gross et al. 2024, Section 2.3).

In this paper, we study the matrices Aλ, and, in particular, the cone in the space R6(+1) that they induce. We can think about this space as being a kind of ‘tropical dual’ to the parameter space C6(+1), and we adopt the same indexing. That is, the entries of λR6(+1) are indexed by B-orbits and edges of N. Then λ(wig)=λig, where i{1,2,3,4,5,6} and gB·G. Observe that the vector λ defines a monomial order on the polynomial ring C[aig|i=1,,6,andgB·G] (provided we specify another order for resolving ties). For this reason, we call λR6(+1) a weight vector.

For weight vectors λ where Trop(ϕ) is differentiable, each column of Aλ is indexed by a consistent leaf-labelling g=(g1,g2,g3). The g-entry of the parameterization ϕN is given by m1(g)+m2(g), where mi(g) is the g-entry of the parameterization of Ti. For each g and mi=mi(g) we define λ(mi) to be the natural product of the row vector of exponents of mi, with the column vector λ. That is,

λ(m1)=λ(w11w22w33w4g1+g2w51)=λ11+λ22+λ33+λ4g1+g2+λ51 1

and

λ(m2)=λ(w11w22w33w42w61)=λ11+λ22+λ33+λ42+λ61. 2

Thus for a given weight vector λ, we take the column of Aλ indexed by g to be the exponent vector of the monomial mi where λ(mi)<λ(mj) for ij{1,2}; in this case, we will say that λ assigns g to tree Ti. The procedure just described is equivalent to describing an initial ideal of the ideal generated by the image of ψN(G,B), where the initial term for each generator, inλ(ψN(G,B)(qg)), is given by the monomial mi.

In Section 3, we give a solution to the dimension problem for general group-based models on the 3-sunlet network for all finite abelian groups of odd order at least 7 (we handle Z/5Z by computation), by constructing λ such that Aλ has maximal rank.

Defining Hyperplanes

The matrix Aλ is determined by inequalities between linear combinations of the coordinates of the weight vector λ. In this subsection, we construct a hyperplane arrangement HG, which divides Euclidean space into the regions on which Trop(ϕ) is differentiable and Aλ is constant. The hyperplanes themselves correspond to regions on which Trop(ϕ) is not differentiable. By understanding the defining hyperplanes and resulting geometry, we are able to construct a weight vector λ so that Aλ has the maximum possible rank, and therefore gives the best lower bound on the dimension of the variety VN(G,B).

Definition 2.2

A hyperplane arrangement is a collection of hyperplanes H={Hi}iI, HiRn. A connected component of the complement Rn\H is a chamber of H. We denote by C(H) the set of chambers of H.

For the 3-sunlet network, each column of Aλ corresponds to a consistent leaf-labelling g=(g1,g2,g3). The column of Aλ corresponding to g can be one of two vectors, and depends on which inequality the coordinates of λ satisfy. Thus, each consistent leaf-labelling g determines a hyperplane of the arrangement we want to construct for the 3-sunlet network. As above, for i=1,2 we write mi(g) for the monomial corresponding to the ith tree in the g-entry of the parametrization map ϕN. Then, we can define the hyperplane arrangement corresponding to the 3-sunlet as the collection H={Hg|g=(g1,g2,g3) is a consistent leaf-labelling}, where

Hg:={λR6(+1)λ(m1(g))=λ(m2(g))}={λR6(+1)λ11+λ22+λ33+λ4g1+g2+λ51=λ11+λ22+λ33+λ42+λ61}.

In Lemma 2.3, we make several observations about the hyperplanes Hg. Note that the hyperplane arrangement H and the resulting chambers C(H) form the Gröbner fan of the ideal generated by the image of ψN(G,B), denoted imψN(G,B). In this interpretation, the hyperplanes themselves consist of those points λ for which the initial ideal inλ(imψN(G,B)) is not a monomial ideal, and the chambers consist of those points λ for which the initial ideal inλ(imψN(G,B)) is monomial.

We make the following observation about the matrix Aλ.

Lemma 2.3

Let μg:=λ6g-λ5g for all gG. The inequalities determining the matrix Aλ are:

0<μ0orμ0<0,and 3
λ4g1+g2-λ42<μg1orμg1<λ4g1+g2-λ42for allg1,g2Gsuch thatg10. 4

In particular, the determining inequalities depend only on g1 and g2.

Proof

The matrix Aλ is constant on a region R if and only if each column is constant on R, so it follows that the hyperplane arrangement we seek is the union of hyperplane arrangements each of whose regions defines the constant regions of a single column.

The assignment of columns is equivalent to choosing the direction of the inequality in λ(m1)<λ(m2) or λ(m1)>λ(m2). Without loss of generality, assume λ(m1)<λ(m2). Expanding λ(m1) and λ(m2) as in equations (1) and (2) we obtain

λ11+λ22+λ33+λ4g1+g2+λ51<λ11+λ22+λ33+λ42+λ61

Cancelling terms we see that

graphic file with name 11538_2025_1506_Equ47_HTML.gif

Inequality (3) controls the assignment of the |G|=+1 columns with label (0,g2,-g2), for g2G. The inequalities given by (4) each control a single column with label (g1,g2,-g1-g2), with g10. Thus, by crossing the hyperplane μ0=0 exactly +1 of the columns of Aλ change. While, when crossing a hyperplane of the form μg1=λ4g1+g2-λ42, exactly one of the columns of Aλ changes. Observe that we cannot simply choose which inequalities are satisfied and expect to find a weight vector λ that achieves this. That is, some combinations of inequalities cannot be simultaneously satisfied. Thus, for a given G, it is unclear how many chambers lie in the hyperplane arrangement H, but it is at most 2(+1)+1. Lemma 3.2 in the next section gives some restrictions on these assignments for groups of odd order, and we will see further examples in Section 4.

Example 2.4

Let G=Z/2Z. We have four consistent leaf-labellings given by (0, 0, 0), (0, 1, 1), (1, 0, 1), (1, 1, 0). The image of ϕNG is given by

ϕNG(w)000=w10w20w30w40w50+w10w20w30w40w60,ϕNG(w)011=w10w21w31w41w50+w10w21w31w41w60,ϕNG(w)101=w11w20w31w41w51+w11w20w31w40w61,ϕNG(w)110=w11w21w30w40w51+w11w21w30w41w61,

where in each case the first monomial, m1 corresponds to T1, and the second monomial m2 corresponds to T2. Pick a weight vector λ with λ40=3,λ41=1,μ0=1, and μ1=-1. We construct the matrix Aλ. The columns of Aλ are indexed by the 4 consistent leaf-labellings, and the rows are indexed by the 12 parameters wig for i=1,,6 and g=0,1. For columns (0, 0, 0) and (0, 1, 1) we have that 0<μ0=1, so in these cases the monomial from T1 is chosen. For (1, 0, 1) we have that λ41-λ40=-2<μ1=-1 so the monomial from T1 is chosen. For (1, 1, 0) we have that λ40-λ41=2>μ1=-1 so the monomial from T2 is chosen. This gives us the following matrix Aλ

graphic file with name 11538_2025_1506_Equ48_HTML.gif

where the entry corresponding to column g1g2g3 and row wig is the exponent of wig in the monomial from the g1g2g3 entry of the expression for ϕNG(w) corresponding to the tree chosen.

To end this section, we make some observations on the hyperplane arrangement H coming from the 3-sunlet and a group-based model (GB). We will define the rank of a chamber CC(H) as the rank of the corresponding tropical Jacobian matrix Aλ for all λC.

First, observe that when we choose λ so that all columns correspond to T1 or T2 (this is possible by choosing μg either very large or very small for all g), then Aλ is equal to the corresponding tropical Jacobian for the group-based phylogenetic tree model for T1 or T2 (albeit with some extra rows of 0’s corresponding to the parameters w6g or w5g respectively). Since, the matrix Aλ has rank equal to the dimension of the toric variety corresponding to Aλ (see e.g., (Sturmfels 1996, Lemma 4.2)), and these varieties are well studied for trees, in both cases, rankAλ=3+1 (see e.g. (Gross et al. 2024, Lemma 4.1)). This describes two chambers with rank equal to 3+1. In fact, there are two more, as the next proposition demonstrates.

Proposition 2.5

Let C and C be two adjacent chambers in C(H), separated by the hyperplane μ0=0. Then rankC=rankC.

Proof

Let λC and λC be weight vectors, and consider the matrices Aλ and Aλ. Suppose, without loss of generality, that in Aλ the columns indexed by (0,g,-g) with gG correspond to T1, and in Aλ they correspond to T2. Thus, in the columns of Aλ indexed by (0,g,-g), entries in the row indexed by w50 are 1, and entries in the row indexed by w60 are 0. On the other hand, in the columns of Aλ indexed by (0,g,-g), entries in the row indexed by w50 are 0, and entries in the row indexed by w60 are 1. All other entries of Aλ and Aλ are the same. Finally, observe that in the rows indexed by w50 and w60, all entries are 0 outside the columns indexed by (0,g,-g) for gG. Thus, the difference between Aλ and Aλ is a swap of the rows indexed by w50 and w60, and this does not affect the rank.

Applying Proposition 2.5 to the two chambers of rank 3+1 found above, we have four chambers of this rank. These come from column assignments where all columns corresponding to a leaf-labelling (0,g,-g) are assigned to Ti, and all other columns assigned to Tj for i,j{1,2}. It is easy to see that all other chambers have rank strictly greater than this (changing any column corresponding to (g1,g2,g3) with g10 will introduce a 1 into the row w51 or w61 which was previously all 0’s), thus we have exactly four chambers of minimal rank 3+1.

Lemma 2.6

Let H be the hyperplane arrangement given by the 3-sunlet network and group-based model (GB). Then H has exactly four chambers of minimal rank 3+1.

Proposition 2.7

Let H be the hyperplane arrangement given by the 3-sunlet network and group-based model (GB). Given a chamber CC(H) of rank r, the rank of each adjacent chamber is between r-1 and r+1.

Proof

As described above, moving to an adjacent chamber is equivalent to either swapping the assignment of all columns indexed by (0,g,-g) for gG, or swapping the assignment of a single column indexed by (g,h,-g,-h) with g0. In the first case, the rank does not change by Proposition 2.5. In the second case, the rank can change by at most 1.

Proposition 2.5 above shows that every chamber is adjacent to at least one other chamber of equal rank. We would also like to know whether every chamber is adjacent to another of chamber of strictly greater rank, or equivalently, whether the only maximal chambers (with respect to rank) are the globally maximal chambers. As we will see in Section 4.3, the answer to this is no, and there do exist locally maximal chambers.

Dimension for Odd Order Groups

In this section we give the dimension of the 3-sunlet variety for general group-based models where G is a finite abelian group of odd order at least 5. Our results agree with (Gross et al. 2024, Conjecture 7.1). Our method is to find a weight vector λ so that the corresponding tropical Jacobian Aλ has rank greater than or equal to 5+1, where |G|=+1, as detailed in Section 2.

First we set out some notation. Assume |G| is odd and let |G|=2t+1. Choose a subset XG with |X|=t and satisfying the property that if gX then -gX. In particular, 0X. For example, for G equal to the cyclic group of order 2t+1 we could have X={1,2,,t}. We will need the following lemma.

Lemma 3.1

Let G be an abelian group with |G|=2t+1, where tN and t3, i.e. |G| is odd with |G|7. Let XG be a subset with |X|=t such that if gX then -gX. Then there exists an injective function h:XG\X such that hg:=h(g) is not equal to 0,-g, or 2g.

Proof

Let X={g1,,gt} and let ki=-giX. We will give an iterative method of choosing the value of hgi, where at each step we update the set of available choices. To begin with, let K={k1,,kt}, this will be our initial set of available choices. For each i=1,,t-3, choose the value of hgi to be an element of K satisfying hgiki and hgi2gi (should 2gi be in K), and remove the value of hgi from K. This is possible, since at each stage, K contains at least 3 elements.

Now it remains to choose hgt-2,hgt-1 and hgt. In the worst case, we have K={kt-2,kt-1,kt}={2gt-2,2gt-1,2gt}. For each gi we take hgi to be the (possibly unique) kj such that ij and kj2gi.

The next lemma demonstrates that there are relationships among the inequalities in Lemma 2.3.

Lemma 3.2

Fix gG such that g-g, and suppose that we have a weight vector λ such that for some hG the consistent leaf labelling (g,h,-g-h) is assigned to T1 and (g,h,-g-h) is assigned to T2 for all hh. If there exists kG such that (-g,k,g-k) is assigned to T2 then (-g,g+h,-h) is assigned to T2.

Proof

By assumption, since λ assigns (g,h,-g-h) to T1 and (g,h,-g-h) to T2, we have λ4g+h-λ4h<μg and λ4g+h-λ4>μg for all hh (see proof of Lemma 2.3). Thus λ4g+h-λ4h<λ4g+h-λ4 for all hh. Now if k=g+h then the result is tautological, so assume kg+h. Let h=k-g so that (-g,k,g-k)=(-g,g+h,-h). Now if (-g,g+h,-h) is assigned to T2 then λ4-λ4g+h>μ-g. Since λ4-λ4g+h=-(λ4g+h-λ4) and -(λ4g+h-λ4)<-(λ4g+h-λ4h), we have -(λ4g+h-λ4h)=λ4h-λ4g+h>μ-g. Thus, λ(m1((-g,g+h,-h)))>λ(m2((-g,g+h,-h))), and consequently, (-g,g+h,-h) is assigned to T2.

Next we describe a procedure for choosing λ so that the matrix Aλ has rank equal to the expected dimension. This procedure will be illustrated below in Example 3.3. For ease of notation, we will write ν for λ4R2t+1, so that νg=λ4g for all gG. In this notation, for a leaf labeling g=(g1,g2,-g1-g2),

λ(m1)<λ(m2)νg1+g2-νg2<μg1.

This means g=(g1,g2,-g1-g2) is assigned to T1 if and only if νg1+g2-νg2<μg1, and g is assigned to T2 if and only if νg1+g2-νg2>μg1.

Choose ν such that νg0 for all gG, with ν0 large enough and all νg small enough such that for all h,h,kG\{0} we have ν0-νh>νh-νk>νh-ν0. Fix gX. By our choice of ν, we can find μg such that ν0-ν-g>μg and μg>νg+h-νh for all hG with h-g. Then for the consistent leaf-labelling (g,-g,0) we have ν0-ν-g>μg, so (g,-g,0) is assigned to T2. For all h-g we have νg+h-νh<μg so the consistent leaf-labelling (g,h,-g-h) is assigned to T1.

Next, consider -gX. In this case, for all hG the consistent leaf-labelling g=(-g,h,g-h) is assigned to T1 if and only if

ν-g+h-νh<μ-g,

and g is assigned to T2 if and only if

ν-g+h-νh>μ-g.

Choose μ-g=-μg. Then for (-g,0,g) we have

ν-g-ν0=-(ν0-ν-g)<-μg=μ-g,

and thus (-g,0,g) is assigned to T1. Now, consider the inequality μg>νg+h-νh, which holds for all h-g. Write h=-g+k with k0. Then we have μg>νk-ν-g+k and therefore ν-g+k-νk>-μg=μ-g. Thus (-g,k,g-k) is assigned to T2 for all k0.

To summarize, at this point, we have found ν,μg and μ-g to give us the following assignments of the consistent leaf-labellings (g,h,-g-h) and (-g,h,g-h) for all hG:

  • To T1 we have assigned (-g,0,g) and (g,h,-g-h) for all h-g.

  • To T2 we have assigned (g,-g,0) and (-g,h,g-h) for all h0.

We repeat the above procedure for every gX. Then for each of the t pairs {g,-g}G\{0} we have +1 consistent leaf-labellings assigned to T1 and +1 consistent leaf-labellings assigned to T2. Now, for all hG assign (0,h,-h) to T2 so that we have t(+1)+(+1)=(t+1)(2t+1) consistent leaf-labellings assigned to T2 and t(2t+1) consistent leaf-labellings assigned to T1. Note that for t2 we have 2=4t<t(2t+1)<2t(2t-1)=(-1).

We give a small example to illustrate.

Example 3.3

Let G=Z/3Z={0,1,2}, and let X={1}. Pick ν0=10, ν1=0, and ν2=2. When g=1, for the consistent leaf-labellings (1, 2, 0), (1, 1, 1), and (1, 0, 2) respectively we have

ν0-ν2=8>ν2-ν1=2>ν1-ν0=-10.

We choose μ1=7 so that (1, 2, 0) is assigned to T2; and (1, 1, 1) and (1, 0, 2) are assigned to T1.

When g=2, for the consistent leaf-labellings (2, 1, 0), (2, 2, 2) and (2, 0, 1) respectively we have

ν0-ν1=10>ν1-ν2=-2>ν2-ν0=-8.

Setting μ2=-μ1=-7 gives us (2, 1, 0) and (2, 2, 2) assigned to T2 and (2, 0, 1) assigned to T1.

Finally, we choose μ0=-1 so that (0, 0, 0), (0, 1, 2), and (0, 2, 1) are assigned to T2.

Returning to an arbitrary finite abelian group of odd order, we choose ν and μ as above (with μ0 negative) so that we have the assignments of leaf-labellings as described above. We will show that for this choice of λ we have rankAλ5+1. Our strategy is to perform row and column operations on Aλ in order to turn it into a block upper triangular matrix without changing the rank.

We will denote the row of Aλ corresponding to the parameter wig as rig. Our first observation is that for any assignment of consistent leaf-labellings to T1 and T2 we have r1g=r5g+r6g for all gG. The rows r1g therefore do not contribute to the rank of Aλ, so we remove them. Note that this corresponds to the notion of a contracted semi-directed network, which we do not introduce here (see Gross et al. (2024) for further details). Next, perform column swaps so that all columns assigned to T2 are to the left of the columns assigned to T1. Observe that for all columns assigned to T2, the entry corresponding to w2g is equal to the entry corresponding to w4g for all gG (where edge labels are as in Figure 2), and the entry corresponding to w5g is 0 for all gG. In terms of the tree topology, this is because in T2 the vertex between e2 and e4 has degree 2, and because T2 does not contain the edge e5. Perform row swaps so that the corresponding edge order is e2,e3,e6,e4,e5 from top to bottom. Next perform the row operations

r4gr4g-r2g

for all gG. This gives a block upper-triangular matrix of the form

graphic file with name 11538_2025_1506_Equ5_HTML.gif 5

It follows that rankAλrankA+rankB. The submatrix A consists of all columns assigned to T2 and rows corresponding to w2g,w3g, and w6g for all gG. These are all the variables associated to edges in T2 that form a 3-star tree, and so we know rankAdimT2=3+1. In Lemma 3.5, we show that rankA=dimT2=3+1. The submatrix B consists of all columns assigned to T1 and rows r4g-r2g and r5g for all gG. The columns of B are given by the consistent leaf-labellings assigned to T1 which are

S=S1S2={(g,h,-g-h)|gX,h-g}{(-g,0,g)|gX},

so |S1|=2t(t)=2 and |S2|=t=2. In the next example, we illustrate the block triangular matrix in (5) for G=Z/3Z.

Example 3.4

Here we continue Example 3.3. With G=Z/3Z and assignments of consistent leaf-labellings as in the example, after all row operations we have the following matrix.

graphic file with name 11538_2025_1506_Equ49_HTML.gif

Note that in this case the dimension of the space containing the variety is (+1)2=9, which is less than the expected dimension of 5+1=11. Therefore the expected dimension cannot be reached, and indeed, explicit computation shows that the variety has dimension 9. Here we can see explicitly that the submatrices A and B have rank strictly less than as described in Lemmas 3.5 and 3.7.

In the next two lemmas, we give the rank of the submatrix A and a lower bound on the rank of the submatrix B.

Lemma 3.5

Let G be an abelian group with |G|=+1 odd and |G|5, and Aλ as above in (5). Then rankA=3+1.

Proof

The submatrix A consists of columns corresponding to consistent leaf-labellings assigned to T2, and rows corresponding to the variables w2g,w3g, and w6g for all gG. This submatrix also appears in the corresponding matrix of exponents for the 3-star tree with edges e2,e3, and e6 as in Figure 3 (possibly after performing some column swaps).

Fig. 3.

Fig. 3

A 3-star tree with taxa labels and edge labels

Denote by K3 the matrix of exponents corresponding to the general group-based model of G on the 3-star tree. The dimension of the variety corresponding to this model is 3+1 (Gross et al. 2024, Lemma 4.1). Since the parameterization of this model is monomial, K3 has rank equal to the dimension of the model.

Let K(g1,g2,g3) be the column of K3 corresponding to the consistent leaf-labelling (g1,g2,g3), so that we have

K(g1,g2,g3)=E61+E22+E33.

We will show that the columns of K3 corresponding to consistent leaf-labellings that we have assigned to T1 can be written as linear combinations of columns corresponding to consistent leaf-labellings we have assigned to T2, thereby showing that rankA=rankK3=3+1.

First, consider the consistent leaf-labelling (-g,0,g). The reader can check that for any hX with hg we have

K(-g,0,g)=K(0,0,0)+K(-g,h,g-h)+K(-h,h-g,g)-K(-h,h,0)-K(0,h-g,g-h),

and all terms on the right hand side are from consistent leaf-labellings that are assigned to T2. Note that since |G|5, at least one such h exists.

Next consider the consistent leaf-labelling (g,h,-g-h) for gX and h-g. We consider two cases separately:

Case 1: hX. For this case we may write

K(g,h,-g-h)=K(g,-g,0)+K(-g,h,g-h)+K(h,g,-g-h)-K(-g,g,0)-K(h,-g,g-h). 6

Case 2: hX. We break this down into two further cases. First, suppose -g-hX. Then we have

K(g,h,-g-h)=K(g,-g,0)+K(0,h,-h)+K(g+h,0,-g-h)-K(0,0,0)-K(g+h,-g,-h), 7

where we are using that although (g+h,0,-g-h) is assigned to T1, the column K(g+h,0,-g-h) is linearly dependent on columns corresponding to leaf-labellings assigned to T2 by the first part of the proof. Therefore we can substitute the relation from (6) into (7) to obtain K(g,h,-g-h) as a linear combination of columns assigned to T2. On the other hand, if -g-hX then

K(g,h,-g-h)=K(g,-g,0)+K(-g-h,h,g)+K(0,g+h,-g-h)-K(0,-g,g)-K(-g-h,g+h,0).

Remark 3.6

Observe that the relations used in the proof above correspond to the cubic binomials in the ideal for the 3-star tree, as described in Sturmfels and Sullivant (2005).

Lemma 3.7

Let G be a finite abelian group with |G|=+1 odd and |G|7, and Aλ as above in (5). Then rankB2.

Proof

The columns of B correspond to the consistent leaf-labellings we assign to T1, that is, those in the set S, where

S=S1S2={(g,h,-g-h)|gX,h-g}{(-g,0,g)|gX}.

The rows of B correspond to the variables w5g and w4g, where we have performed the row operation r4gr4g-r2g, for all gG. For the leaf-labelling (g1,g2,g3), we have m1(g1,g2,g3)=w11w22w33w4g1+g2w51. Thus, the column of B corresponding to that leaf labelling is the vector in C2+2 given by

B(g1,g2,g3)=E51+E4g1+g2-E42.

To prove the result, it is sufficient to find a subset BS of 2 linearly independent columns.

First consider the columns corresponding to consistent leaf-labellings in S2. Here each column is given by

B(-g,0,g)=E5-g+E4-g-E40,

for gX. Since -gX, the column B(-g,0,g) is the only column of B with a non-zero component in the basis vector E5-g, and is therefore linearly independent of all other columns.

Next we consider columns coming from consistent leaf-labellings in S1. For each gX consider the leaf-labellings (g,0,-g),(g,g,-2g), and (g,hg,-g-hg), where for each gX the element hg is chosen from G\(X{0}) with the conditions that hg-g,2g, and if gg then hghg (this is possible by Lemma 3.1). We claim that the set

B={B(-g,0,g)|gX}{B(g,0,-g),B(g,g,-2g),B(g,hg,-g-hg)|gX}

is linearly independent. Let

Vg=spanC{B(g,0,-g),B(g,g,-2g),B(g,hg,-g-hg)}.

By the choice of hg, dimVg=3 for all gX. Thus, to prove the claim, it is sufficient to show that VgVg={0} for all g,gX with gg.

First, by considering the basis vectors E5g and E5, we must have that VgVgW where W=spanC{E4h|hG}, so that VgVg=(VgW)(VgW). Now VgW is spanned by the vectors B(g,0,-g)-B(g,g,-2g) and B(g,0,-g)-B(g,hg,-g-hg), where

B(g,0,-g)-B(g,g,-2g)=2E4g-E42g-E40,B(g,0,-g)-B(g,hg,-g-hg)=E4g-E4g+hg+E4g-E40.

An arbitrary element of vVgW is then given by

v=α(B(g,0,-g)-B(g,g,-2g))+β(B(g,0,-g)-B(g,hg,-g-hg))=(2α+β)E4g-αE42g+βE4g-βE4g+hg-(α+β)E40,

for α,βC. Observe that from the choice of hg, the basis vectors appearing in this expression are distinct. Now suppose that for gX with gg, we have another such element vVgW with coefficients α and β. We will show that if v=v then v=v=0 so that (VgW)(VgW)={0} as claimed. We have

v=(2α+β)E4g-αE42g+βE4g-βE4g+hg-(α+β)E40, 8
v=(2α+β)E4-αE42g+βE4g-βE4g+hg-(α+β)E40. 9

Now suppose v=v. By examining the coefficients of E40 we must have

α+β=α+β. 10

Next consider the coefficient of E4. Either E4 appears in the expression for v or 2α+β=0. For the former, since gg,0 and gX so ghg, we must have either g=2g or g=g+hg. We consider these three cases separately.

Case 1: g=2g. By equating coefficients of E4=E42g we have 2α+β=-α, and substituting in to equation (10) gives β=3α+2β. Next consider the coefficient of E4g. Either this is zero (i.e. β=0) or E4g appears in the expression for v.

If β0 then since hgg,hg,0, or g=2g, we must have hg=g+hg, and therefore β=-β, from which it follows by Equation (10) that α+2β=α. Now since the coefficient of E4g+hg is not zero we must have either g+hg=g or hg. If g+hg=hg then substituting in hg=g+hg gives g+g+hg=hg and therefore g=-g, a contradiction. We therefore must have g+hg=g so then -β=2α+β, i.e., α=0. But now we have simultaneous equations 2β=α and 3β=α, so α=β=0.

Now, if β=0 then we have α=-2α and β=3α. Substituting these values into equation (8) and setting v=v gives

-αE4g+2αE42g+3αE4g-3αE4g+hg=2αE4-αE42g,

so we conclude that α=0, and therefore v=v=0.

Case 2: g=g+hg. By equating coefficients of E4=E4g+hg we have 2α+β=-β. Next consider the coefficient of E4g. As before, either this is zero or E4g appears in the expression for v.

If β0 then we must have hg=2g, so that β=-α and therefore β=α+2β by Equation (10), and then 3α+3β=0. Now consider the coefficient of E4g+hg. Either g+hg=g or g+hg=hg. If g+hg=g then g+hg+hg=g so hg = -hg, but this contradicts how hg and hg were chosen; both hg and hgG\(X{0}), but then by definition of X we must have -hg and -hgX. Thus we must have g+hg=hg, and so β=-β. Now we have β=-α=-β so by equation 10 we have 0=α+3β. Solving this simultaneously with 3α+3β=0 gives v=v=0.

If β=0 we have 2α=-β and so by Equation (10) α=3α. Substituting into equation (8) and setting v=v gives us α=0 as in case 1, and therefore v=v=0.

Case 3: 2α+β=0. In this case we have β=-2α so α=-(α+β). This gives us

v=(α+β)E42g+2(α+β)E4g-2(α+β)E4g+hg-(α+β)E40.

In particular, if v0 then α+β0 so that v has 4 linearly independent non-zero terms. We have

v=(2α+β)E4g-αE42g+βE4g-βE4g+hg-(α+β)E40,

so if v=v we must have exactly four non-zero terms in the expression for v. This is only possible if α=0 or 2α+β=0. If α=0 then we have

v=βE4g+βE4g-βE4g+hg-βE40,v=βE42g+2βE4g-2βE4g+hg-βE40.

If v=v then by equating coefficients we must have β=0 and thus v=v=0. On the other hand, if 2α+β=0 we get

v=-αE42g-2αE4g+2αE4g+hg+αE40,v=-αE42g-2αE4g+2αE4g+hg+αE40.

Examining the coefficient of E42g we see that since gg and g0 we must have -α=±2α, so α=0 and v=v=0.

Putting together the results of this section and (Gross et al. 2024, Proposition 4.2), we get Theorem 1.1, which we restate here for convenience. By direct calculation (see Table 4), we have that for G=Z/5Z, the dimension of VNG is 21, the expected dimension, so we include this result in the statement.

Table 4.

The number of samples for each Aλ rank when G=Z/5Z

rank 13 14 15 16 17 18 19 20 21
# chambers 4 80 560 2,160 5,228 11,520 27,960 41,360 24,480
# chambers .004% .071% .494% 1.906% 4.612% 10.163% 24.667% 36.488% 21.596%

Theorem 1.1

Let G be a finite abelian group of odd order +15, and let N be the 3-sunlet network under the general group-based model given by G, with corresponding phylogenetic network variety VNG. Then the affine dimension of VNG is given by

dimVNG=5+1.

Proof

For G=Z/5Z, the result is acheived computationally by using random sampling to find a weight vector λ for which Aλ has rank 5+1=21 (Table 4). For all other G, choose λ as described in this section. Through row operations we can transform the matrix Aλ into a block upper triangular matrix of the form

Aλ=A0B,

so that rankAλrankA+rankB. By Lemma 3.5, rankA=3+1, and, by Lemma 3.7, rankB2. Thus, we obtain rankAλ5+1, and so the affine dimension of VNG is at least 5+1. On the other hand, Lemma 2.1 says that dimVNG is at most 5+1.

As discussed at the beginning of this work, the 3-sunlet was the only sunlet where a dimension formula was not given in Gross et al. (2024). Since level-1 phylogenetic networks can be broken down into trees and sunlet networks, for the case when G is an abelian group of odd order at least 5 and N is a level-1 phylogenetic network, we can now give a full dimension result.

Theorem 3.8

Let N be a level-1 phylgenetic network with n leaves, m edges, and c cycles. Let G be a finite abelian group of odd order +15. Then the variety corresponding to N under the general group-based model for G, denoted VNG, has dimension (m-c)+1.

Proof

Following (Gross et al. 2024, Theorem 1.1), we prove the result by induction on the number of cut edges of N. If N has no cut edges, then it is either the 3-star tree or a sunlet network. If N is the 3-star tree, then it has no cycles and dimVNG=3+1 (Gross et al. 2024, Lemma 4.1). If N is a 3-sunlet network then it has a single cycle and 6 edges, and dimVNG=5+1 by Theorem 1.1. If N is an n-sunlet network with n>3 then it has a single cycle and 2n edges, and dimVNG=(2n-1)+1 (Gross et al. 2024, Theorem 4.7). Thus in all cases the result holds.

For the induction step, suppose that N is a level-1 phylogenetic network with a cut edge e, m edges, and c cycles. Let N1 and N2 be the networks obtained by cutting N at e and observe that both N1 and N2 have strictly fewer cut edges than N. For i=1,2 let mi and ci be the number of edges and cycles respectively of Ni, then by induction we have dimVNiG=(mi-ci)+1. Next we have that the ideal ING defining VNiG is given by the toric fiber product of IN1G and IN2G (Cummings et al. 2024, Remark 3.3). Using (Gross et al. 2024, Corollary 3.4) we obtain

dimVNG=dimVN1G+dimVN2G-(+1)=(m1-c1)+1+(m2-c2)+1-(+1)=(m1+m2-1-(c1+c2))+1=(m-c)+1.

The dimension results in Gross et al. (2024) were used to prove identifiability statements for level-1 phylogenetic networks under group-based models. Now that we have the dimension result for 3-sunlet networks, we can remove the ‘triangle-free’ restriction placed on those statements for the general-group based model for G, when G is a finite abelian group of odd order at least 5. The proof of the following result is identical to (Gross et al. 2024, Proposition 6.9). First, recall the following two definitions. We say that two phylogenetic networks N1 and N2 are distinguishable over G if VN1GVN2G and VN2GVN1G. Given a level-1 phylogenetic network N on n leaves and a subset A of the leaf-set [n], the network restricted to A, denoted N|A, is the level-1 phylogenetic network obtained from N by removing all edges and vertices that do not lie on any path between two leaves in A, and suppressing any resulting vertices of degree 2 (except the root vertex). See (Gross and Long 2018, Definition 4.1) for a full definition.

Proposition 3.9

Let N1 and N2 be two level-1 phylogenetic networks on n leaves and both with exactly c cycles. Let G be a finite abelian group of odd order 5. If there exists a subset A[n] such that either

  1. N1|A and N2|A are level-1 phylogenetic networks with distinct numbers of cycles; or

  2. N1|A is a tree and N2|A is a level-1 phylogenetic network (i.e. with at least 1 cycle); or

  3. N1|A and N2|A are distinct trees;

then N1 and N2 are distinguishable over G.

Experimental Results

Theorem 1.1 confirms that for odd order groups, the general group-based model on a 3-sunlet has the expected dimension 5+1 (except G=Z/3Z). In this section, we investigate the dimension for small finite abelian groups, and whilst an analogous construction of λ for even order groups does not give us a maximal rank Aλ, through experiments we find that the expected dimension is obtained in all cases once G is sufficiently large. The code we used to perform these calculations was written in Julia Bezanson et al. (2017) and available to download at https://github.com/shelbycox/3-Sunlet.

Sampling Methods

We use the hyperplane description from Section 2.3 to compute the possible matrices Aλ (and their ranks) that can appear for the groups Z/3Z, Z/4Z, Z/2Z×Z/2Z, and Z/5Z. For each of the possible 2(+1)+1 regions, we use OSCAR Oscar (2024) to test whether the region is full dimensional. We then use random sampling of points p=(μ,λ4) in (-.5,.5)2(+1) to obtain exactly one point in each region. For other small groups mentioned in this section, we used only random sampling with 232 samples for each group to obtain the results. In each case, we retain at most one point for each region.

Chamber counts for small groups

Tables 1, 2, 3, 4 and 5 are the result of the computations described above for the groups Z/3Z, Z/4Z, Z/2Z×Z/2Z, Z/5Z, and Z/6Z respectively. For the first four groups listed, we confirmed that these are exactly the regions of the hyperplane arrangement. For Z/6Z, the data in the table is the result of sampling 232 points and then retaining no more than one point per region. In addition, we include an illustration of relationships between chambers for Z/3Z in Figure 4. We make some observations.

Table 1.

The number of samples for each Aλ rank when G=Z/3Z

rank 7 8 9
# chambers 4 24 64
% chambers 4.3% 26.1% 69.6%

Table 2.

The number of samples for each Aλ rank when G=Z/4Z

rank 10 11 12 13 14 15 16
# chambers 4 48 180 496 864 624 112
% chambers .2% 2.06% 7.7% 21.3% 37.1% 26.8% 4.8%

Table 3.

The number of samples for each Aλ rank when G=Z/2Z×Z/2Z

rank 10 11 12 13 14 15 16
# chambers 4 48 156 584 1056 480 0
% chambers .2% 2.06% 6.7% 25.1% 45.4% 20.6% 0%

Table 5.

The number of samples for each Aλ rank when G=Z/6Z

rank 16 17 18 19 20 21
# chambers 4 120 1296 7,180 26,576 79,156
% chambers .00004% .00129% .01391% .07707% .28528% .84969%
rank 22 23 24 25 26
# chambers 229,069 742,458 2,148,510 3,606,334 2,475,117
% chambers 2.45892% 7.96986% 23.06303% 38.71193% 26.56897%

Fig. 4.

Fig. 4

Poset of H(Z/3Z), graded by distance (right to left) from the starting chamber, T1 (at the bottom of the poset or far right of the diagram). The number in each node is the rank of the corresponding chamber. Key: blue/square - rankAλ=7, red/triangle - rankAλ=8, yellow/circle - rankAλ=9

Remark 4.1

As observed in Lemma 2.6, in all cases we have exactly 4 chambers of lowest rank, which is equal to the dimension of the 3-star tree, 3+1. Two of the chambers correspond to when all leaf-labellings are assigned to either T1 or T2. Weight vectors λ with λ5g sufficiently large and λ6g=0 for all gG, or vice-versa, lie in these chambers. The remaining two chambers correspond to when all leaf-labellings (0,g,-g) are assigned to T1 and all others assigned to T2; and when all leaf-labellings (0,g,-g) are assigned to T2 and all others assigned to T1. These chambers can be reached by taking the previous weight vector λ and swapping the values of λ50 and λ60.

For G=Z/3Z, these chambers can be seen in Figure 4, drawn in squares and highlighted in blue at the very top and very bottom of the diagram. The shortest path in the poset between T1 and T2 has length 7, meaning that 7 hyperplanes need to be crossed to reach one from the other.

Remark 4.2

When G=Z/3Z, most chambers have rank 9, which is the maximum possible, because the space containing the corresponding variety is C9. However, for Z/4Z the maximum rank, 16, is achieved by only 4.8% of chambers. For Z/2Z×Z/2Z the maximum rank is 15, which is achieved by only 20.6% of chambers. For both Z/4Z and Z/2Z×Z/2Z rank 14 chambers are observed the most (37.1% and 45.4%).

Remark 4.3

From Table 2 and Table 3, we observe that the number of chambers in the hyperplane arrangement is the same for Z/4Z and Z/2Z×Z/2Z. We speculate that this is true in general for groups of the same order. However, we also observe that the distribution of ranks for these chambers differs. Thus, the distribution of ranks can depend on the structure of G, and not just |G|.

Locally maximal chambers

In our investigations, we were curious whether a greedy algorithm could be used to find an appropriate λ by moving from chamber to chamber. For such a method to work, there should be no locally maximal chambers that do not achieve the maximum rank. A locally maximal chamber in terms of rank means that all adjacent chambers have rank less than or equal to the rank of the chamber. Here, we describe our search for locally maximal chambers that do not achieve maximum rank. To do this computationally, we cycle through every chamber in the hyperplane arrangement found by random sampling (as in Section 4.2) and check whether the rank of this chamber is less than the global maximum, but greater than the rank of all adjacent chambers. The code is available in the file locallyMaximalChambers.jl.

The number of such chambers for small groups is given in Table 6. For groups with |G|5, the rank of the locally maximal chambers of deficient rank is always one less than the dimension of the model. For G=Z/6Z, this appears to not be the case. For this group, our code did not complete, but had still found 625 locally maximal chambers of ranks 23, 24, and 25 before it was terminated. However, since we were not able to verify that we had found all chambers in the hyperplane arrangement through random sampling, it may be the case that some of the locally maximal chambers found are not maximal at all.

Table 6.

The number of locally maximal chambers of deficient rank for each group. Locally maximal chambers of deficient rank are chambers which have non-maximal rank, and for which all adjacent chambers have rank lesser than or equal to the rank of the chamber

Group Z/3Z Z/4Z Z/2Z×Z/2Z Z/5Z Z/6Z
Locally maximal chambers 0 128 0 1840 625
% 0% 5.50% 0% 1.62% 0.006%

Weight vector for groups of even order

In Section 3, we construct a weight vector λ where Aλ is the maximum rank possible for groups of odd order. Here, for even order groups, we construct an analogous vector λ by following the construction in Section 3, but including elements of order 2 in the set X. In this case, we find that the rank of Aλ does not always achieve the empirically maximum rank. That is, random sampling sometimes finds λ with Aλ having greater rank than Aλ.

Observe that in all cases, through random sampling we are able to find a weight vector where Aλ has the maximum possible rank according to Lemma 2.1. This means that in all cases in Table 7, the dimension of the variety is equal to the expected dimension of 5+1.

Table 7.

The third column of the table (Empirical maximum) lists the maximum rank of Aλ found by randomly sampling λ. The fourth column (Modified Section 3 construction) lists the rank of Aλ, when λ is the vector described at the beginning of this section. The last column records the difference between the third and fourth columns.

rank(Aλ)
Modified Section 3
Group |G| Empirical maximum construction Gap
Z/4Z 4 16 15 1
Z/2Z2 15 13 2
Z/6Z 6 26 26
Z/8Z 36 36
Z/4Z×Z/2Z 8 36
Z/2Z3 29 7
Z/10Z 10 46 46
Z/12Z 12 56 56
Z/3Z×(Z/2Z)2 56
Z/14Z 14 66 66
Z/16Z 16 76 76
Z/8Z×Z/2Z 76
Z/4Z×Z/4Z 76
Z/4Z×Z/2Z2 76
Z/2Z4 61 15
Z/18Z 18 86 86
Z/3Z2×Z/2Z 86
Z/2Z5 32 156 125 31

We see that for λ constructed according to the methods in Section 3, Aλ often achieves the maximum empirical rank, and it is only for powers of Z/2Z that this construction does not work. Furthermore, in these cases, the difference between the rank of Aλ and the maximum rank is equal to |G|-1. In Appendix A, we provide an example comparing the constructed λ and the empirically maximal λ for G=Z/4Z. Specifically, we find a weight vector λ for which the corresponding matrix, Aλ, achieves the maximum rank of 16, and compare it to the matrix Aλ obtained from a construction analogous to that in Section 3. This gives good evidence that it is always possible to achieve the maximum possible rank (i.e., 5+1) as the rank of Aλ, when |G|5.

Discussion

In this paper, we give a dimension formula for varieties associated to a 3-sunlet phylogenetic network and general group-based model, where the group G is a finite abelian group of odd order. To do this, we use ideas from tropical geometry and linear algebra. Our proof relies on the fact that for odd order groups, there are no elements of order 2, and thus the non-identity elements can be partitioned into two sets of equal size (one of which we refer to as X in Section 3), each containing mutually inverse elements. Thus, our proof does not obviously generalise to even-order groups, and a full understanding of the dimension of these models remains open. In Section 4.4 we construct weight vectors for even-order groups by assigning self-inverse elements to X and following the construction in Section 3. However, as shown in Table 7, for those groups that are products of Z/2Z, the rank of the corresponding Aλ is less than the dimension of the model. Interestingly, the difference is |G|-1.

We have not yet explored models for which the subgroup BAutG is non-trivial. In Gross et al. (2024), a dimension formula is given for triangle-free phylogenetic networks for all group-based models (i.e., for all such subgroups B). This is achieved by choosing a weight vector that, for each edge in the network, is constant on the coordinates associated to the B-orbits on that edge. Since the structure of Aut(G) varies with G, this is only possible if we consider the two orbits {0} and G\{0}. However, in our experiments we found that for the 3-sunlet, the weight vectors λ with Aλ of maximal rank were not constant on these orbits, suggesting that a case-by-case analysis may be required. We identified equivalent weight vectors λ (i.e. those for which Aλ is the same) with chambers in a hyperplane arrangement. Each chamber can be interpreted as a toric degeneration of the variety. Two of the chambers correspond to the two distinct phylogenetic trees displayed by a 3-sunlet. These have the lowest rank among all chambers, and the rank is equal to the dimension of the variety associated to the group-based model on the corresponding tree. The remaining chambers correspond to a mixture of the Fourier coordinates for these two trees, and so there is no clear phylogenetic intepretation of these chambers.

The investigations in this paper highlight the intricate challenges involved in understanding 3-cycles. In the sampling experiments in Section 4, we were able to examine the ranks of all chambers in the hyperplane arrangement. However, it is only when |G|5 that the expected dimension of the variety (5+1) is less than the dimension of the ambient space (|G|2), so the cases with |G|4 are exceptional. We observe that as the group gets larger, there are proportionally more chambers of maximal rank. However, due to the large growth in the number of chambers as the size of the group grows, we were not able to determine if this is a general pattern.

A first approach at obtaining the result for all finite abelian groups may be to try to adapt the weight vector λ. Guidance on appropriate adaptations could be found by understanding how changes in the vector correspond to moving between chambers. Indeed, a good understanding of the hyperplane arrangement may make it possible to devise an algorithm to search for a weight vector λ for which Aλ has maximal rank. However, as we have shown in Section 4.3, this is also not straightforward. In this section, we found that for at least some groups there are locally maximal chambers, and therefore a greedy algorithm starting at a lowest rank chamber and moving to chambers of strictly larger rank may not terminate on a globally maximal chamber.

To further understand the varieties associated to 3-sunlet networks and group-based models, we would like to be able to describe generating sets of the corresponding ideals. Polynomials in these ideals are called phylogenetic invariants, and are useful for determining model identifiability, model selection, and even topology inference from sequence data. However, calculating generating sets is challenging. Using Macaulay2Grayson and Stillman (2022) and elimination theory, we attempted to find a Gröbner basis for the ideal corresponding to a 3-sunlet under the general group-based model with G=Z/5Z – the smallest odd-order group for which we have a dimension result and for which the variety does not fill the whole space. After 50 days on an HPC these computations were still running. We also used the MultigradedImplicitization package Cummings and Hollering (2026) to find generators of fixed total degree. The computations completed for degrees up to and including 7, and in each case there were no generators. After 45 days the computations for degree 8 were still running. In this case we have a polynomial ring with n=25 generators, so the dimension of the space spanned by monomials of degree d=8 is large: n+d-1d=328=10518300. This highlights the difficulty in calculating generators even for small models. Further work is necessary for this to be achieveable.

Acknowledgements

SC was supported by the National Science Foundation Graduate Research Fellowship under Grant No. DGE-1841052, and by the National Science Foundation under Grant No. 1855135 during the writing of this paper. SM was supported by the Biotechnology and Biological Sciences Research Council (BBSRC), part of UK Research and Innovation, through the Core Capability Grant BB/CCG1720/1 at the Earlham Institute and is grateful for HPC support from NBI’s Research Computing group. SM is grateful for further funding from BBSRC (grant number BB/X005186/1) which also supported this work. EG was supported by the National Science Foundation grant DMS-1945584. This project was initiated at the “Algebra of Phylogenetic Networks Workshop" held at the University of Hawai‘i at Mānoa and supported by National Science Foundation grant DMS-1945584. Additional parts of this research were performed while EG and SC were visiting the Institute for Mathematical and Statistical Innovation (IMSI) for the semester-long program on “Algebraic Statistics and Our Changing World," IMSI is supported by the National Science Foundation (Grant No. DMS-1929348).

Appendix A. Example Weights and Matrices

A.1 Example when G=Z/4Z.

Below, we study the λ-construction from Section 3, adapted for even order groups (so all order two elements are in X), for G=Z/4Z. We denote this weight vector by λguess and denote the corresponding matrix by Aguess. As noted in Table 7, rkAguess=15, which is not the empirical maximum. Through random sampling, we find in Table 2 that there are 112 chambers of the Z/4Z-sunlet arrangement whose corresponding matrices achieve the maximal rank. We pick a weight vector, λmax, in one of these chambers so that the corresponding matrix, Amax, has maximal rank and differs from Aguess in exactly one column. For the λmax chosen here, the two matrices differ only in the column [[2], [1], [1]].

graphic file with name 11538_2025_1506_Equ50_HTML.gif

Funding

Open Access funding enabled and organized by Projekt DEAL.

Footnotes

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