Phylogenetic networks are becoming increasingly popular in phylogenetics since they have the ability to describe a wider range of evolutionary events than their tree counterparts. In this paper, we study Markov models on phylogenetic networks and their associated geometry. We restrict our attention to large-cycle networks, networks with a single undirected cycle of length at least four. Using tools from computational algebraic geometry, we show that the semi-directed network topology is generically identifiable for Jukes-Cantor large-cycle network models.
In phylogenetics, it is important for the phylogenetic network model parameters to be identifiable so that the evolutionary histories of a group of species can be consistently inferred. However, as the complexity of the phylogenetic network models grows, the identifiability of network models becomes increasingly difficult to analyze. As an attempt to analyze the identifiability of network models, we check whether two networks are distinguishable. In this paper, we specifically study the distinguishability of phylogenetic network models associated with level-2 networks. Using an algebraic appro…
Phylogenetic networks can represent evolutionary events that cannot be described by phylogenetic trees. These networks are able to incorporate reticulate evolutionary events such as hybridization, introgression, and lateral gene transfer. Recently, network-based Markov models of DNA sequence evolution have been introduced along with model-based methods for reconstructing phylogenetic networks. For these methods to be consistent, the network parameter needs to be identifiable from data generated under the model. Here, we show that the semi-directed network parameter of a triangle-free, level-1 …
We compare the phylogenetic tensors for various trees and networks for two, three and four taxa. If the probability spaces between one tree or network and another are not identical then there will be phylogenetic tensors that could have arisen on one but not the other. We call these two trees or networks distinguishable from each other. We show that for the binary symmetric model there are no two-taxon trees and networks that are distinguishable from each other, however there are three-taxon trees and networks that are distinguishable from each other. We compare the time parameters for the phy…
Phylogenetic networks can model more complicated evolutionary phenomena that trees fail to capture such as horizontal gene transfer and hybridization. The same Markov models that are used to model evolution on trees can also be extended to networks and similar questions, such as the identifiability of the network parameter or the invariants of the model, can be asked. In this paper we focus on finding the invariants of the Cavendar-Farris-Neyman (CFN) model on level-1 phylogenetic networks. We do this by reducing the problem to finding invariants of sunlet networks, which are level-1 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.
Identifiability of phylogenetic networks and quintet concordance factors
University of Edinburgh（爱丁堡大学）；New Jersey Institute of Technology（新泽理工学院）；Texas State University（德克萨斯州立大学）；University of Hawai‘i at Mānoa（夏威夷大学马诺阿分校）；University of Alaska Fairbanks（阿拉斯加大学费尔班克斯分校）
Less rigid than phylogenetic trees, phylogenetic networks allow the description of a wider range of evolutionary events. In this note, we explain how to extend the rank invariants from phylogenetic trees to phylogenetic networks evolving under the general Markov model and the equivariant models.
Invariants for complicated objects such as those arising in phylogenetics, whether they are invariants as matrices, polynomials, or other mathematical structures, are important tools for distinguishing and working with such objects. In this paper, we generalize a complete polynomial invariant on trees to a class of phylogenetic networks called separable networks, which will include orchard networks. Networks are becoming increasingly important for their ability to represent reticulation events, such as hybridization, in evolutionary history. We provide a function from the space of internally m…
The Fair Proportion of a species in a phylogenetic tree is a very simple measure that has been used to assess its value relative to the overall phylogenetic diversity represented by the tree. It has recently been proved by Fuchs and Jin to be equal to the Shapley Value of the coallitional game that sends each subset of species to its rooted Phylogenetic Diversity in the tree. We prove in this paper that this result extends to the natural translations of the Fair Proportion and the rooted Phylogenetic Diversity to rooted phylogenetic networks. We also generalize to rooted phylogenetic networks …
An important and well-studied problem in phylogenetics is to compute a \emph{consensus tree} so as to summarize the common features within a collection of rooted phylogenetic trees, all whose leaf-sets are bijectively labeled by the same set~(X) of species. More recently, however, it has become of interest to find a consensus for a collection of more general, rooted directed acyclic graphs all of whose sink-sets are bijectively labeled by~(X), so called rooted \emph{phylogenetic networks}. These networks are used to analyse the evolution of species that cross with one another, such as plants a…
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.
Phylogenetic networks are rooted, labelled directed acyclic graphs which are commonly used to represent reticulate evolution. There is a close relationship between phylogenetic networks and multi-labelled trees (MUL-trees). Indeed, any phylogenetic network $N$ can be 'unfolded' to obtain a MUL-tree $U(N)$ and, conversely, a MUL-tree $T$ can in certain circumstances be 'folded' to obtain a phylogenetic network $F(T)$ that exhibits $T$. In this paper, we study properties of the operations $U$ and $F$ in more detail. In particular, we introduce the class of stable networks, phylogenetic networks …
A large class of phylogenetic networks can be obtained from trees by the addition of horizontal edges between the tree edges. These networks are called tree based networks. Reticulation-visible networks and child-sibling networks are all tree based. In this work, we present a simply necessary and sufficient condition for tree-based networks and prove that there is a universal tree based network for each set of species such that every phylogenetic tree on the same species is a base of this network. The existence of universal tree based network implies that for any given set of phylogenetic tree…
Phylogenetic trees are widely used to display estimates of how groups of species evolved. Each phylogenetic tree can be seen as a collection of clusters, subgroups of the species that evolved from a common ancestor. When phylogenetic trees are obtained for several data sets (e.g. for different genes), then their clusters are often contradicting. Consequently, the set of all clusters of such a data set cannot be combined into a single phylogenetic tree. Phylogenetic networks are a generalization of phylogenetic trees that can be used to display more complex evolutionary histories, including ret…
Phylogenetics is the study of the evolutionary relationships between organisms. One of the main challenges in the field is to take biological data for a group of organisms and to infer an evolutionary tree, a graph that represents these relationships. Developing practical and efficient methods for inferring phylogenetic trees has lead to a number of interesting mathematical questions across a variety of fields. However, due to hybridization and gene flow, a phylogenetic network may be a better representation of the evolutionary history of some groups of organisms. In this chapter, we introduce…
Maximum parsimony is one of the most frequently-discussed tree reconstruction methods in phylogenetic estimation. However, in recent years it has become more and more apparent that phylogenetic trees are often not sufficient to describe evolution accurately. For instance, processes like hybridization or lateral gene transfer that are commonplace in many groups of organisms and result in mosaic patterns of relationships cannot be represented by a single phylogenetic tree. This is why phylogenetic networks, which can display such events, are becoming of more and more interest in phylogenetic res…
Phylogenetic networks are a generalization of phylogenetic trees that are used to represent reticulate evolution. Unrooted phylogenetic networks form a special class of such networks, which naturally generalize unrooted phylogenetic trees. In this paper we define two operations on unrooted phylogenetic networks, one of which is a generalization of the well-known nearest-neighbor interchange (NNI) operation on phylogenetic trees. We show that any unrooted phylogenetic network can be transformed into any other such network using only these operations. This generalizes the well-known fact that an…
Phylogenetic networks provide a means of describing the evolutionary history of sets of species believed to have undergone hybridization or gene flow during their evolution. The mutation process for a set of such species can be modeled as a Markov process on a phylogenetic network. Previous work has shown that a site-pattern probability distributions from a Jukes-Cantor phylogenetic network model must satisfy certain algebraic invariants. As a corollary, aspects of the phylogenetic network are theoretically identifiable from site-pattern frequencies. In practice, because of the probabilistic n…
Rooted phylogenetic networks provide a way to describe species' relationships when evolution departs from the simple model of a tree. However, networks inferred from genomic data can be highly tangled, making it difficult to discern the main reticulation signals present. In this paper, we describe a natural way to transform any rooted phylogenetic network into a simpler canonical network, which has desirable mathematical and computational properties, and is based only on the 'visible' nodes in the original network. The method has been implemented and we demonstrate its application to some exam…
Recently, considerable effort has been put into developing fast algorithms to reconstruct a rooted phylogenetic network that explains two rooted phylogenetic trees and has a minimum number of hybridization vertices. With the standard approach to tackle this problem being combinatorial, the reconstructed network is rarely unique. From a biological point of view, it is therefore of importance to not only compute one network, but all possible networks. In this paper, we make a first step towards approaching this goal by presenting the first algorithm---called allMAAFs---that calculates all maximu…
Recently much attention has been devoted to the construction of phylogenetic networks which generalize phylogenetic trees in order to accommodate complex evolutionary processes. Here we present an efficient, practical algorithm for reconstructing level-1 phylogenetic networks - a type of network slightly more general than a phylogenetic tree - from triplets. Our algorithm has been made publicly available as the program LEV1ATHAN. It combines ideas from several known theoretical algorithms for phylogenetic tree and network reconstruction with two novel subroutines. Namely, an exponential-time e…
A binary phylogenetic network on a taxon set $X$ is a rooted acyclic digraph in which the degree of each nonleaf node is three and its leaves (i.e.degree-one nodes) are uniquely labeled with the taxa of $X$. It is tree-child if each nonleaf node has at least one child of indegree one. A set of binary phylogenetic trees may or may not be simultaneously displayed in a binary tree-child network. Necessary conditions for multiple phylogenetic trees being simultaneously displayed in a tree-child network are given here. In particular, it is proved that any two phylogenetic trees can always simultane…
The search space of rooted phylogenetic trees is vast and a major research focus of recent decades has been the development of algorithms to effectively navigate this space. However this space is tiny when compared with the space of rooted phylogenetic networks, and navigating this enlarged space remains a poorly understood problem. This, and the difficulty of biologically interpreting such networks, obstructs adoption of networks as tools for modelling reticulation. Here, we argue that the superimposition of biologically motivated constraints, via an SQL-style language, can both stimulate use…
Rooted phylogenetic networks are used to describe evolutionary histories that contain non-treelike evolutionary events such as hybridization and horizontal gene transfer. In some cases, such histories can be described by a phylogenetic base-tree with additional linking arcs, which can for example represent gene transfer events. Such phylogenetic networks are called tree-based. Here, we consider two possible generalizations of this concept to nonbinary networks, which we call tree-based and strictly-tree-based nonbinary phylogenetic networks. We give simple graph-theoretic characterizations of …
On Tree-Network Distinguishability and Full Identifiability of Phylogenetic Networks
Delft Institute of Applied Mathematics, Delft University of Technology（代尔夫特理工大学应用数学研究所）；European Molecular Biology Laboratory, European Bioinformatics Institute (EMBL-EBI)（欧洲分子生物学实验室欧洲生物信息学研究所）
Phylogenetic networks are a generalization of phylogenetic trees that are used in biology to represent reticulate or non-treelike evolution. Recently, several algorithms have been developed which aim to construct phylogenetic networks from biological data using {\em triplets}, i.e. binary phylogenetic trees on 3-element subsets of a given set of species. However, a fundamental problem with this approach is that the triplets displayed by a phylogenetic network do not necessary uniquely determine or {\em encode} the network. Here we propose an alternative approach to encoding and constructing ph…
Phylogenetic networks generalize evolutionary trees, and are commonly used to represent evolutionary histories of species that undergo reticulate evolutionary processes such as hybridization, recombination and lateral gene transfer. Recently, there has been great interest in trying to develop methods to construct rooted phylogenetic networks from triplets, that is rooted trees on three species. However, although triplets determine or encode rooted phylogenetic trees, they do not in general encode rooted phylogenetic networks, which is a potential issue for any such method. Motivated by this fa…
Phylogenetic trees and networks are leaf-labelled graphs that are used to describe evolutionary histories of species. The Tree Containment problem asks whether a given phylogenetic tree is embedded in a given phylogenetic network. Given a phylogenetic network and a cluster of species, the Cluster Containment problem asks whether the given cluster is a cluster of some phylogenetic tree embedded in the network. Both problems are known to be NP-complete in general. In this article, we consider the restriction of these problems to several well-studied classes of phylogenetic networks. We show that…
Semi-directed phylogenetic networks have recently emerged as a class of phylogenetic networks sitting between rooted (directed) and unrooted (undirected) phylogenetic networks as they contain both directed as well as undirected edges. While the spaces of rooted phylogenetic networks and unrooted phylogenetic networks have been analyzed in recent years and various rearrangement moves to traverse these spaces have been introduced, the results do not immediately carry over to semi-directed phylogenetic networks. Here, we propose a simple rearrangement move for semi-directed phylogenetic networks,…
In biodiversity conservation it is often necessary to prioritize the species to conserve. Existing approaches to prioritization, e.g. the Fair Proportion Index and the Shapley Value, are based on phylogenetic trees and rank species according to their contribution to overall phylogenetic diversity. However, in many cases evolution is not treelike and thus, phylogenetic networks have come to the fore as a generalization of phylogenetic trees, allowing for the representation of non-treelike evolutionary events, such as horizontal gene transfer or hybridization. Here, we extend the concepts of phy…
Tree-based phylogenetic networks, which may be roughly defined as leaf-labeled networks built by adding arcs only between the original tree edges, have elegant properties for modeling evolutionary histories. We answer an open question of Francis, Semple, and Steel about the complexity of determining how far a phylogenetic network is from being tree-based, including non-binary phylogenetic networks. We show that finding a phylogenetic tree covering the maximum number of nodes in a phylogenetic network can be be computed in polynomial time via an encoding into a minimum-cost maximum flow problem…
In phylogenetics, phylogenetic trees are rooted binary trees, whereas phylogenetic networks are rooted arbitrary acyclic digraphs. Edges are directed away from the root and leaves are uniquely labeled with taxa in phylogenetic networks. For the purpose of validating evolutionary models, biologists check whether or not a phylogenetic tree is contained in a phylogenetic network on the same taxa. This tree containment problem is known to be NP-complete. A phylogenetic network is reticulation-visible if every reticulation node separates the root of the network from some leaves. We answer an open p…
A fundamental problem in the study of phylogenetic networks is to determine whether or not a given phylogenetic network contains a given phylogenetic tree. We develop a quadratic-time algorithm for this problem for binary nearly-stable phylogenetic networks. We also show that the number of reticulations in a reticulation visible or nearly stable phylogenetic network is bounded from above by a function linear in the number of taxa.
In [2] it was proven that the Cass algorithm is a polynomial-time algorithm for constructing level<=2 networks from clusters. Here we demonstrate, for each k>=0, a polynomial-time algorithm for constructing level-k phylogenetic networks from clusters. Unlike Cass the algorithm scheme given here is only of theoretical interest. It does, however, strengthen the hope that efficient polynomial-time algorithms (and perhaps fixed parameter tractable algorithms) exist for this problem.
The evolutionary relationships among organisms have traditionally been represented using rooted phylogenetic trees. However, due to reticulate processes such as hybridization or lateral gene transfer, evolution cannot always be adequately represented by a phylogenetic tree, and rooted phylogenetic networks that describe such complex processes have been introduced as a generalization of rooted phylogenetic trees. In fact, estimating rooted phylogenetic networks from genomic sequence data and analyzing their structural properties is one of the most important tasks in contemporary phylogenetics. …
Rearrangement operations transform a phylogenetic tree into another one and hence induce a metric on the space of phylogenetic trees. Popular operations for unrooted phylogenetic trees are NNI (nearest neighbour interchange), SPR (subtree prune and regraft), and TBR (tree bisection and reconnection). Recently, these operations have been extended to unrooted phylogenetic networks, which are generalisations of phylogenetic trees that can model reticulated evolutionary relationships. Here, we study global and local properties of spaces of phylogenetic networks under these three operations. In par…
Polyploidization is an evolutionary process by which a species acquires multiple copies of its complete set of chromosomes. The reticulate nature of the signal left behind by it means that phylogenetic networks offer themselves as a framework to reconstruct the evolutionary past of species affected by it. The main strategy for doing this is to first construct a so called multiple-labelled tree and to then somehow derive such a network from it. The following question therefore arises: How much can be said about that past if such a tree is not readily available? By viewing a polyploid dataset as…
This article concerns the following question arising in computational evolutionary biology. For a given subclass of phylogenetic networks, what is the maximum value of 0 <= p <= 1 such that for every input set T of rooted triplets, there exists some network N(T) from the subclass such that at least p|T| of the triplets are consistent with N(T)? Here we prove that the set containing all triplets (the full triplet set) in some sense defines p, and moreover that any network N achieving fraction p' for the full triplet set can be converted in polynomial time into an isomorphic network N'(T) achiev…
Phylogenetic networks extend phylogenetic trees to allow for modeling reticulate evolutionary processes such as hybridization. They take the shape of a rooted, directed, acyclic graph, and when parameterized with evolutionary parameters, such as divergence times and population sizes, they form a generative process of molecular sequence evolution. Early work on computational methods for phylogenetic network inference focused exclusively on reticulations and sought networks with the fewest number of reticulations to fit the data. As processes such as incomplete lineage sorting (ILS) could be at …
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 …
The problem of constructing an optimal rooted phylogenetic network from a set of rooted triplets is an NP-hard problem. In this paper, we present a heuristic algorithm called TripNet which tries to construct an optimal rooted phylogenetic network from an arbitrary set of triplets. We prove some theorems to justify the performance of the algorithm.
Phylogenetic networks are a generalization of phylogenetic trees that allow for the representation of non-treelike evolutionary events, like recombination, hybridization, or lateral gene transfer. In this paper, we present and study a new class of phylogenetic networks, called tree-child phylogenetic networks, where every non-extant species has some descendant through mutation. We provide an injective representation of these networks as multisets of vectors of natural numbers, their path multiplicity vectors, and we use this representation to define a distance on this class and to give an alig…
Phylogenetic Diversity (PD) is a prominent quantitative measure of the biodiversity of a collection of present-day species (taxa). This measure is based on the evolutionary distance among the species in the collection. Loosely speaking, if $\mathcal{T}$ is a rooted phylogenetic tree whose leaf set $X$ represents a set of species and whose edges have real-valued lengths (weights), then the PD score of a subset $S$ of $X$ is the sum of the weights of the edges of the minimal subtree of $\mathcal{T}$ connecting the species in $S$. In this paper, we define several natural variants of the PD score …
Driven by the need for better models that allow one to shed light into the question how life's diversity has evolved, phylogenetic networks have now joined phylogenetic trees in the center of phylogenetics research. Like phylogenetic trees, such networks canonically induce collections of phylogenetic trees, clusters, and triplets, respectively. Thus it is not surprising that many network approaches aim to reconstruct a phylogenetic network from such collections. Related to the well-studied perfect phylogeny problem, the following question is of fundamental importance in this context: When does…
A phylogenetic network is a directed acyclic graph that visualises an evolutionary history containing so-called reticulations such as recombinations, hybridisations or lateral gene transfers. Here we consider the construction of a simplest possible phylogenetic network consistent with an input set T, where T contains at least one phylogenetic tree on three leaves (a triplet) for each combination of three taxa. To quantify the complexity of a network we consider both the total number of reticulations and the number of reticulations per biconnected component, called the level of the network. We …
Phylogenetic networks provide a way to describe and visualize evolutionary histories that have undergone so-called reticulate evolutionary events such as recombination, hybridization or horizontal gene transfer. The level k of a network determines how non-treelike the evolution can be, with level-0 networks being trees. We study the problem of constructing level-k phylogenetic networks from triplets, i.e. phylogenetic trees for three leaves (taxa). We give, for each k, a level-k network that is uniquely defined by its triplets. We demonstrate the applicability of this result by using it to pro…
Phylogenetic networks are becoming of increasing interest to evolutionary biologists due to their ability to capture complex non-treelike evolutionary processes. From a combinatorial point of view, such networks are certain types of rooted directed acyclic graphs whose leaves are labelled by, for example, species. A number of mathematically interesting classes of phylogenetic networks are known. These include the biologically relevant class of stable phylogenetic networks whose members are defined via certain "fold-up" and "un-fold" operations that link them with concepts arising within the th…
Phylogenetic networks are a generalization of phylogenetic trees that allow for the representation of evolutionary events acting at the population level, like recombination between genes, hybridization between lineages, and lateral gene transfer. While most phylogenetics tools implement a wide range of algorithms on phylogenetic trees, there exist only a few applications to work with phylogenetic networks, and there are no open-source libraries either. In order to improve this situation, we have developed a Perl package that relies on the BioPerl bundle and implements many algorithms on phylog…
Phylogenetic networks are a generalization of phylogenetic trees allowing for the representation of non-treelike evolutionary events such as hybridization. Typically, such networks have been analyzed based on their `level', i.e. based on the complexity of their 2-edge-connected components. However, recently the question of how `treelike' a phylogenetic network is has become the center of attention in various studies. This led to the introduction of \emph{tree-based networks}, i.e. networks that can be constructed from a phylogenetic tree, called the \emph{base tree}, by adding additional edges…
In this work, we answer an open problem in the study of phylogenetic networks. Phylogenetic trees are rooted binary trees in which all edges are directed away from the root, whereas phylogenetic networks are rooted acyclic digraphs. For the purpose of evolutionary model validation, biologists often want to know whether or not a phylogenetic tree is contained in a phylogenetic network. The tree containment problem is NP-complete even for very restricted classes of networks such as tree-sibling phylogenetic networks. We prove that this problem is solvable in cubic time for stable phylogenetic ne…
