arXiv subjects

Gabriel Cardona

Publications and source records attributed to Gabriel Cardona.

At least 19 recordsLinked to original sources

arXiv2026

Counting Spinal Tree-Child Networks via Word Encodings and Generating Functions

We study the enumeration of spinal tree-child phylogenetic networks, a rigid family of tree-child networks in which all internal vertices lie on a single root--to--leaf path. We provide two complementary combinatorial frameworks. First, we introduce a word model: unlabeled spinal networks correspond to a suitable class of restricted words with fixed multiplicities, taken modulo a simple relabeling equivalence, which yields an explicit closed enumeration. Second, we develop a symbolic-method approach based on a marked version of trees that admits a clean recursive specification; its boxed-product translation leads to a solvable bivariate generating function and a direct derivation of the coefficients.

math.CO↗

arXiv2023

Generation of orchard and tree-child networks

Phylogenetic networks are an extension of phylogenetic trees that allow for the representation of reticulate evolution events. One of the classes of networks that has gained the attention of the scientific community over the last years is the class of orchard networks, that generalizes tree-child networks, one of the most studied classes of networks. In this paper we focus on the combinatorial and algorithmic problem of the generation of orchard networks, and also of tree-child networks. To this end, we use that these networks are defined as those that can be recovered by a reversing a certain reduction process. Then, we show how to choose a ``minimum'' reduction process among all that can be applied to a network, and hence we get a unique representation of the network that, in fact, can be given in terms of sequences of pairs of integers, whose length is related to the number of leaves and reticulations of the network. Therefore, the generation of networks is reduced to the generation of such sequences of pairs. Our main result is a recursive method for the efficient generation of all minimum sequences, and hence of all orchard (or tree-child) networks with a given number of leaves and reticulations. An implementation in C of the algorithms described in this paper, along with some computational experiments, can be downloaded from the public repository https://github.com/gerardet46/OrchardGenerator. Using this implementation, we have computed the number of orchard networks with at most 6 leaves and 8 reticulations.

q-bio.PE↗

arXiv2023

Comparison of orchard networks using their extended $\mu$-representation

Phylogenetic networks generalize phylogenetic trees in order to model reticulation events. Although the comparison of phylogenetic trees is well studied, and there are multiple ways to do it in an efficient way, the situation is much different for phylogenetic networks. Some classes of phylogenetic networks, mainly tree-child networks, are known to be classified efficiently by their $\mu$-representation, which essentially counts, for every node, the number of paths to each leaf. In this paper, we introduce the extended $\mu$-representation of networks, where the number of paths to reticulations is also taken into account. This modification allows us to distinguish orchard networks and to define a sound metric on the space of such networks that can, moreover, be computed efficiently. The class of orchard networks, as well as being one of the classes with biological significance (one such network can be interpreted as a tree with extra arcs involving coexisting organisms), is one of the most generic ones (in mathematical terms) for which such a representation can (conjecturally) exist, since a slight relaxation of the definition leads to a problem that is Graph Isomorphism Complete.

q-bio.PE↗

arXiv2020

Revisiting Shao and Sokal's $B_2$ index of phylogenetic balance

Measures of phylogenetic balance, such as the Colless and Sackin indices, play an important role in phylogenetics. Unfortunately, these indices are specifically designed for phylogenetic trees, and do not extend naturally to phylogenetic networks (which are increasingly used to describe reticulate evolution). This led us to consider a lesser-known balance index, whose definition is based on a probabilistic interpretation that is equally applicable to trees and to networks. This index, known as the $B_2$ index, was first proposed by Shao and Sokal in 1990. Surprisingly, it does not seem to have been studied mathematically since. Likewise, it is used only sporadically in the biological literature, where it tends to be viewed as arcane. In this paper, we study mathematical properties of $B_2$ such as its expectation and variance under the most common models of random trees and its extremal values over various classes of phylogenetic networks. We also assess its relevance in biological applications, and find it to be comparable to that of the Colless and Sackin indices. Altogether, our results call for a reevaluation of the status of this somewhat forgotten measure of phylogenetic balance.

q-bio.PE↗

arXiv2020

Counting Tree-Child Networks and Their Subclasses

Galled trees are studied as a recombination model in population genetics. This class of phylogenetic networks is generalized into tree-child, galled and reticulation-visible network classes by relaxing a structural condition imposed on galled trees. We count tree-child networks through enumerating their component graphs. Explicit counting formulas are also given for galled trees through their relationship to ordered trees, phylogenetic networks with few reticulations and phylogenetic networks in which the child of each reticulation is a leaf.

q-bio.PE↗

arXiv2019

Generation of Tree-Child phylogenetic networks

Phylogenetic networks generalize phylogenetic trees by allowing the modelization of events of reticulate evolution. Among the different kinds of phylogenetic networks that have been proposed in the literature, the subclass of binary tree-child networks is one of the most studied ones. However, very little is known about the combinatorial structure of these networks. In this paper we address the problem of generating all possible binary tree-child networks with a given number of leaves in an efficient way via reduction/augmentation operations that extend and generalize analogous operations for phylogenetic trees and are biologically relevant. Since our solution is recursive, this also provides us with a recurrence relation giving an upper bound on the number of such networks.

cs.DS↗

arXiv2013

The expected value of the squared euclidean cophenetic metric under the Yule and the uniform models

The cophenetic metrics $d_{φ,p}$, for $p\in {0}\cup[1,\infty[$, are a recent addition to the kit of available distances for the comparison of phylogenetic trees. Based on a fifty years old idea of Sokal and Rohlf, these metrics compare phylogenetic trees on a same set of taxa by encoding them by means of their vectors of cophenetic values of pairs of taxa and depths of single taxa, and then computing the $L^p$ norm of the difference of the corresponding vectors. In this paper we compute the expected value of the square of $d_{φ,2}$ on the space of fully resolved rooted phylogenetic trees with $n$ leaves, under the Yule and the uniform probability distributions.

q-bio.PE↗

arXiv2013

Cophenetic metrics for phylogenetic trees, after Sokal and Rohlf

Phylogenetic tree comparison metrics are an important tool in the study of evolution, and hence the definition of such metrics is an interesting problem in phylogenetics. In a paper in Taxon fifty years ago, Sokal and Rohlf proposed to measure quantitatively the difference between a pair of phylogenetic trees by first encoding them by means of their half-matrices of cophenetic values, and then comparing these matrices. This idea has been used several times since then to define dissimilarity measures between phylogenetic trees but, to our knowledge, no proper metric on weighted phylogenetic trees with nested taxa based on this idea has been formally defined and studied yet. Actually, the cophenetic values of pairs of different taxa alone are not enough to single out phylogenetic trees with weighted arcs or nested taxa. In this paper we define a family of cophenetic metrics that compare phylogenetic trees on a same set of taxa by encoding them by means of their vectors of cophenetic values of pairs of taxa and depths of single taxa, and then computing the $L^p$ norm of the difference of the corresponding vectors. Then, we study, either analytically or numerically, some of their basic properties: neighbors, diameter, distribution, and their rank correlation with each other and with other metrics.

q-bio.PE↗

arXiv2012

Exact formulas for the variance of several balance indices under the Yule model

One of the main applications of balance indices is in tests of null models of evolutionary processes. The knowledge of an exact formula for a statistic of a balance index, holding for any number n of leaves, is necessary in order to use this statistic in tests of this kind involving trees of any size. In this paper we obtain exact formulas for the variance under the Yule model of the Sackin index, the Colless index and the total cophenetic index of binary rooted phylogenetic trees with n leaves. We also obtain the covariance of the Sackin and the total cophenetic index.

q-bio.PE↗

arXiv2012

The expected value under the Yule model of the squared path-difference distance

The path-difference metric is one of the oldest and most popular distances for the comparison of phylogenetic trees, but its statistical properties are still quite unknown. In this paper we compute the expected value under the Yule model of evolution of its square on the space of fully resolved rooted phylogenetic trees with n leaves. This complements previous work by Steel-Penny and Mir-Rosselló, who computed this mean value for fully resolved unrooted and rooted phylogenetic trees, respectively, under the uniform distribution.

q-bio.PE↗

arXiv2010

A metric for galled networks

Galled networks, directed acyclic graphs that model evolutionary histories with reticulation cycles containing only tree nodes, have become very popular due to both their biological significance and the existence of polynomial time algorithms for their reconstruction. In this paper we prove that Nakhleh's $m$ measure is a metric for this class of phylogenetic networks and hence it can be safely used to evaluate galled network reconstruction methods.

q-bio.PE↗

arXiv2009

Comparison of Galled Trees

Galled trees, directed acyclic graphs that model evolutionary histories with isolated hybridization events, have become very popular due to both their biological significance and the existence of polynomial time algorithms for their reconstruction. In this paper we establish to which extent several distance measures for the comparison of evolutionary networks are metrics for galled trees, and hence when they can be safely used to evaluate galled tree reconstruction methods.

q-bio.PE↗

arXiv2009

The comparison of tree-sibling time consistent phylogenetic networks is graph isomorphism-complete

In a previous work, we gave a metric on the class of semibinary tree-sibling time consistent phylogenetic networks that is computable in polynomial time; in particular, the problem of deciding if two networks of this kind are isomorphic is in P. In this paper, we show that if we remove the semibinarity condition above, then the problem becomes much harder. More precisely, we proof that the isomorphism problem for generic tree-sibling time consistent phylogenetic networks is polynomially equivalent to the graph isomorphism problem. Since the latter is believed to be neither in P nor NP-complete, the chances are that it is impossible to define a metric on the class of all tree-sibling time consistent phylogenetic networks that can be computed in polynomial time.

q-bio.PE↗

arXiv2008

On Nakhleh's latest metric for phylogenetic networks

We prove that Nakhleh's latest dissimilarity measure for phylogenetic networks is a metric on the classes of tree-child phylogenetic networks, of semi-binary time consistent tree-sibling phylogenetic networks, and of multi-labeled phylogenetic trees. We also prove that it distinguishes phylogenetic networks with different reduced versions. In this way, it becomes the dissimilarity measure for phylogenetic networks with the strongest separation power available so far.

q-bio.PE↗

arXiv2008

Path lengths in tree-child time consistent hybridization networks

Hybridization networks are representations of evolutionary histories that allow for the inclusion of reticulate events like recombinations, hybridizations, or lateral gene transfers. The recent growth in the number of hybridization network reconstruction algorithms has led to an increasing interest in the definition of metrics for their comparison that can be used to assess the accuracy or robustness of these methods. In this paper we establish some basic results that make it possible the generalization to tree-child time consistent (TCTC) hybridization networks of some of the oldest known metrics for phylogenetic trees: those based on the comparison of the vectors of path lengths between leaves. More specifically, we associate to each hybridization network a suitably defined vector of `splitted' path lengths between its leaves, and we prove that if two TCTC hybridization networks have the same such vectors, then they must be isomorphic. Thus, comparing these vectors by means of a metric for real-valued vectors defines a metric for TCTC hybridization networks. We also consider the case of fully resolved hybridization networks, where we prove that simpler, `non-splitted' vectors can be used.

q-bio.PE↗

arXiv2008

Nodal distances for rooted phylogenetic trees

Dissimilarity measures for (possibly weighted) phylogenetic trees based on the comparison of their vectors of path lengths between pairs of taxa, have been present in the systematics literature since the early seventies. But, as far as rooted phylogenetic trees goes, these vectors can only separate non-weighted binary trees, and therefore these dissimilarity measures are metrics only on this class. In this paper we overcome this problem, by splitting in a suitable way each path length between two taxa into two lengths. We prove that the resulting splitted path lengths matrices single out arbitrary rooted phylogenetic trees with nested taxa and arcs weighted in the set of positive real numbers. This allows the definition of metrics on this general class by comparing these matrices by means of metrics in spaces of real-valued $n\times n$ matrices. We conclude this paper by establishing some basic facts about the metrics for non-weighted phylogenetic trees defined in this way using $L^p$ metrics on these spaces of matrices.

q-bio.PE↗

arXiv2008

A Distance Metric for Tree-Sibling Time Consistent Phylogenetic Networks

The presence of reticulate evolutionary events in phylogenies turn phylogenetic trees into phylogenetic networks. These events imply in particular that there may exist multiple evolutionary paths from a non-extant species to an extant one, and this multiplicity makes the comparison of phylogenetic networks much more difficult than the comparison of phylogenetic trees. In fact, all attempts to define a sound distance measure on the class of all phylogenetic networks have failed so far. Thus, the only practical solutions have been either the use of rough estimates of similarity (based on comparison of the trees embedded in the networks), or narrowing the class of phylogenetic networks to a certain class where such a distance is known and can be efficiently computed. The first approach has the problem that one may identify two networks as equivalent, when they are not; the second one has the drawback that there may not exist algorithms to reconstruct such networks from biological sequences. We present in this paper a distance measure on the class of tree-sibling time consistent phylogenetic networks, which generalize tree-child time consistent phylogenetic networks, and thus also galled-trees. The practical interest of this distance measure is twofold: it can be computed in polynomial time by means of simple algorithms, and there also exist polynomial-time algorithms for reconstructing networks of this class from DNA sequence data. The Perl package Bio::PhyloNetwork, included in the BioPerl bundle, implements many algorithms on phylogenetic networks, including the computation of the distance presented in this paper.

q-bio.PE↗

arXiv2008

Two metrics for general phylogenetic networks

We prove that Nakhleh's latest dissimilarity measure for phylogenetic networks separates distinguishable phylogenetic networks, and that a slight modification of it provides a true distance on the class of all phylogenetic networks.

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