In this paper, we investigate a conjecture by von Haeseler concerning the Maximum Parsimony method for phylogenetic estimation, which was published by the Newton Institute in Cambridge on a list of open phylogenetic problems in 2007. This conjecture deals with the question whether Maximum Parsimony trees are hereditary. The conjecture suggests that a Maximum Parsimony tree for a particular (DNA) alignment necessarily has subtrees of all possible sizes which are most parsimonious for the corresponding subalignments. We answer the conjecture affirmatively for binary alignments on five taxa but also show how to construct examples for which Maximum Parsimony trees are not hereditary. Apart from showing that a most parsimonious tree cannot generally be reduced to a most parsimonious tree on fewer taxa, we also show that compatible most parsimonious quartets do not have to provide a most parsimonious supertree. Last, we show that our results can be generalized to Maximum Likelihood for certain nucleotide substitution models.
Phylogenetic (i.e. leaf-labeled) trees play a fundamental role in evolutionary research. A typical problem is to reconstruct such trees from data like DNA alignments (whose columns are often referred to as characters), and a simple optimization criterion for such reconstructions is maximum parsimony. It is generally assumed that this criterion works well for data in which state changes are rare. In the present manuscript, we prove that each phylogenetic tree $T$ with $n\geq 20 k$ leaves is uniquely defined by the set $A_k(T)$, which consists of all characters with parsimony score $k$ on $T$. T…
One of the main aims of phylogenetics is the reconstruction of the correct evolutionary tree when data concerning the underlying species set are given. These data typically come in the form of DNA, RNA or protein alignments, which consist of various characters (also often referred to as sites). Often, however, tree reconstruction methods based on criteria like maximum parsimony may fail to provide a unique tree for a given dataset, or, even worse, reconstruct the `wrong' tree (i.e. a tree that differs from the one that generated the data). On the other hand it has long been known that if the a…
Within the field of phylogenetics there is great interest in distance measures to quantify the dissimilarity of two trees. Here, based on an idea of Bruen and Bryant, we propose and analyze a new distance measure: the Maximum Parsimony (MP) distance. This is based on the difference of the parsimony scores of a single character on both trees under consideration, and the goal is to find the character which maximizes this difference. In this article we show that this new distance is a metric and provides a lower bound to the well-known Subtree Prune and Regraft (SPR) distance. We also show that t…
To the known fact that Parsimony method sometimes fails on the problem of inferring species trees from gene trees, here we proved that no mater of what topology the true 9-taxon and greater species tree is the only thing one needs to break down Parsimony is to have in this species tree three consecutive inner edges not going through the root but perhaps ending on it with lengths T1, T2, T3 of some proportions. Obviously, the probability to meet these lengths is growing in general with the size of species tree. Therefore, Parsimony may be applied only when the described lengths of edges cannot …
The parsimony score of a character on a tree equals the number of state changes required to fit that character onto the tree. We show that for unordered, reversible characters this score equals the number of tree rearrangements required to fit the tree onto the character. We discuss implications of this connection for the debate over the use of consensus trees or total evidence, and show how it provides a link between incongruence of characters and recombination.
One of the main aims in phylogenetics is the estimation of ancestral sequences based on present-day data like, for instance, DNA alignments. One way to estimate the data of the last common ancestor of a given set of species is to first reconstruct a phylogenetic tree with some tree inference method and then to use some method of ancestral state inference based on that tree. One of the best-known methods both for tree inference as well as for ancestral sequence inference is Maximum Parsimony (MP). In this manuscript, we focus on this method and on ancestral state inference for fully bifurcating…
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 used to display the relationship of different species whose evolution is not treelike, which is the case, for instance, in the presence of hybridization events or horizontal gene transfers. Tree inference methods such as Maximum Parsimony need to be modified in order to be applicable to networks. In this paper, we discuss two different definitions of Maximum Parsimony on networks, "hardwired" and "softwired", and examine the complexity of computing them given a network topology and a character. By exploiting a link with the problem Multicut, we show that computing the…
We examine a mathematical question concerning the reconstruction accuracy of the Fitch algorithm for reconstructing the ancestral sequence of the most recent common ancestor given a phylogenetic tree and sequence data for all taxa under consideration. In particular, for the symmetric 4-state substitution model which is also known as Jukes-Cantor model, we answer affirmatively a conjecture of Li, Steel and Zhang which states that for any ultrametric phylogenetic tree and a symmetric model, the Fitch parsimony method using all terminal taxa is more accurate, or at least as accurate, for ancestra…
We consider the problem of inferring an ancestral state from observations at the leaves of a tree, assuming the state evolves along the tree according to a two-state symmetric Markov process. We establish a general branching rate condition under which maximum parsimony, a common reconstruction method requiring only the knowledge of the tree, succeeds better than random guessing uniformly in the depth of the tree. We thereby generalize previous results of (Zhang et al., 2010) and (Gascuel and Steel, 2010). Our results apply to both deterministic and i.i.d. edge weights.
In phylogenetic studies, biologists often wish to estimate the ancestral discrete character state at an interior vertex $v$ of an evolutionary tree $T$ from the states that are observed at the leaves of the tree. A simple and fast estimation method --- maximum parsimony --- takes the ancestral state at $v$ to be any state that minimises the number of state changes in $T$ required to explain its evolution on $T$. In this paper, we investigate the reconstruction accuracy of this estimation method further, under a simple symmetric model of state change, and obtain a number of new results, both fo…
One of the goals of phylogenetic research is to find the species tree describing the evolutionary history of a set of species. But the trees derived from geneti data with the help of tree inference methods are gene trees that need not coincide with the species tree. This can for example happen when so-called deep coalescence events take place. It is also known that species trees can differ from their most likely gene trees. Therefore, as a means to find the species tree, it has been suggested to use subtrees of the gene trees, for example triples, and to puzzle them together in order to find t…
Given two phylogenetic trees on the same set of taxa X, the maximum parsimony distance d_MP is defined as the maximum, ranging over all characters c on X, of the absolute difference in parsimony score induced by c on the two trees. In this note we prove that for binary trees there exists a character achieving this maximum that is convex on one of the trees (i.e. the parsimony score induced on that tree is equal to the number of states in the character minus 1) and such that the number of states in the character is at most 7d_MP - 5. This is the first non-trivial bound on the number of states r…
Tuffley and Steel (1997) proved that Maximum Likelihood and Maximum Parsimony methods in phylogenetics are equivalent for sequences of characters under a simple symmetric model of substitution with no common mechanism. This result has been widely cited ever since. We show that small changes to the model assumptions suffice to make the two methods inequivalent. In particular, we analyze the case of bounded substitution probabilities as well as the molecular clock assumption. We show that in these cases, even under no common mechanism, Maximum Parsimony and Maximum Likelihood might make conflict…
As an alternative to parsimony analyses, stochastic models have been proposed (Lewis, 2001), (Nylander, et al., 2004) for morphological characters, so that maximum likelihood or Bayesian analyses may be used for phylogenetic inference. A key feature of these models is that they account for ascertainment bias, in that only varying, or parsimony-informative characters are observed. However, statistical consistency of such model-based inference requires that the model parameters be identifiable from the joint distribution they entail, and this issue has not been addressed. Here we prove that para…
In phylogenetics, distances are often used to measure the incongruence between a pair of phylogenetic trees that are reconstructed by different methods or using different regions of genome. Motivated by the maximum parsimony principle in tree inference, we recently introduced the maximum parsimony (MP) distance, which enjoys various attractive properties due to its connection with several other well-known tree distances, such as TBR and SPR. Here we show that computing the MP distance between two trees, a NP-hard problem in general, is fixed parameter tractable in terms of the TBR distance bet…
One of the main aims of phylogenetics is to reconstruct the \enquote{Tree of Life}. In this respect, different methods and criteria are used to analyze DNA sequences of different species and to compare them in order to derive the evolutionary relationships of these species. Maximum Parsimony is one such criterion for tree reconstruction and, it is the one which we will use in this paper. However, it is well-known that tree reconstruction methods can lead to wrong relationship estimates. One typical problem of Maximum Parsimony is long branch attraction, which can lead to statistical inconsiste…
Recently, the perfect phylogeny model with persistent characters has attracted great attention in the literature. It is based on the assumption that complex traits or characters can only be gained once and lost once in the course of evolution. Here, we consider a generalization of this model, namely Dollo parsimony, that allows for multiple character losses. More precisely, we take a combinatorial perspective on the notion of Dollo-$k$ characters, i.e. traits that are gained at most once and lost precisely $k$ times throughout evolution. We first introduce an algorithm based on the notion of s…
Phylogenetic methods typically rely on an appropriate model of how data evolved in order to infer an accurate phylogenetic tree. For molecular data, standard statistical methods have provided an effective strategy for extracting phylogenetic information from aligned sequence data when each site (character) is subject to a common process. However, for other types of data (e.g. morphological data), characters can be too ambiguous, homoplastic or saturated to develop models that are effective at capturing the underlying process of change. To address this, we examine the properties of a classic bu…
In phylogenetic analysis, for non-molecular data, particularly morphology, parsimony optimization is the most commonly employed approach. In the past and present application of the parsimony principle, extra step numbers have been added across different characters without proper justification. This practice, however, has caused the impacts of characters to be inflated or deflated without a valid reason. To resolve this methodological deficiency, here I present a corrected parsimony criterion for reconstructing phylogenies. In essence, character rather than step is the most fundamental unit. Ac…
In this paper we investigate mathematical questions concerning the reliability (reconstruction accuracy) of Fitch's maximum parsimony algorithm for reconstructing the ancestral state given a phylogenetic tree and a character. In particular, we consider the question whether the maximum parsimony method applied to a subset of taxa can reconstruct the ancestral state of the root more accurately than when applied to all taxa, and we give an example showing that this indeed is possible. A surprising feature of our example is that ignoring a taxon closer to the root improves the reliability of the m…
Enumerating monophyletic characters in mathematical phylogenetics
Institute of Mathematics and Computer Science, Greifswald University（格赖夫斯瓦尔德大学数学与计算机科学研究所）；School of Mathematics and Statistics, University of New South Wales（新南威尔士大学数学与统计学院）；Department of Mathematical Sciences, New Jersey Institute of Technology（新泽西理工学院数学科学系）；National Institute for Theory and Mathematics in Biology, Northwestern University and The University of Chicago（西北大学和芝加哥大学生物理论与数学国家研究所）
Applying a method to reconstruct a phylogenetic tree from random data provides a way to detect whether that method has an inherent bias towards certain tree `shapes'. For maximum parsimony, applied to a sequence of random 2-state data, each possible binary phylogenetic tree has exactly the same distribution for its parsimony score. Despite this pleasing and slightly surprising symmetry, some binary phylogenetic trees are more likely than others to be a most parsimonious (MP) tree for a sequence of $k$ such characters, as we show. For $k=2$, and unrooted binary trees on six taxa, any tree with …
Inferring the ancestral state at the root of a phylogenetic tree from states observed at the leaves is a problem arising in evolutionary biology. The simplest technique -- majority rule -- estimates the root state by the most frequently occurring state at the leaves. Alternative methods -- such as maximum parsimony - explicitly take the tree structure into account. Since either method can outperform the other on particular trees, it is useful to consider the accuracy of the methods on trees generated under some evolutionary null model, such as a Yule pure-birth model. In this short note, we an…
Within the field of phylogenetics there is great interest in distance measures to quantify the dissimilarity of two trees. Recently, a new distance measure has been proposed: the Maximum Parsimony (MP) distance. This is based on the difference of the parsimony scores of a single character on both trees under consideration, and the goal is to find the character which maximizes this difference. Here we show that computation of MP distance on two \emph{binary} phylogenetic trees is NP-hard. This is a highly nontrivial extension of an earlier NP-hardness proof for two multifurcating phylogenetic t…
Inheritance Entropy: A Model-Independent Method to Probe the Hereditary Structure of Cell Lineage Trees
Sapienza Università di Roma（罗马第一大学）；Italian Institute of Technology（意大利理工学院）；Consiglio Nazionale delle Ricerche（国家研究委员会）；Istituto Nazionale di Fisica Nucleare（国家核物理研究所）；Fondazione IRCCS San Gerardo dei Tintori（圣杰拉尔多代蒂尼奥里IRCCS基金会）
The problem Parsimony Haplotyping (PH) asks for the smallest set of haplotypes which can explain a given set of genotypes, and the problem Minimum Perfect Phylogeny Haplotyping (MPPH) asks for the smallest such set which also allows the haplotypes to be embedded in a perfect phylogeny, an evolutionary tree with biologically-motivated restrictions. For PH, we extend recent work by further mapping the interface between ``easy'' and ``hard'' instances, within the framework of (k,l)-bounded instances where the number of 2's per column and row of the input matrix is restricted. By exploring, in the…
This paper studies a Markov chain for phylogenetic reconstruction which uses a popular transition between tree topologies known as subtree pruning-and-regrafting (SPR). We analyze the Markov chain in the simpler setting that the generating tree consists of very short edge lengths, short enough so that each sample from the generating tree (or character in phylogenetic terminology) is likely to have only one mutation, and that there enough samples so that the data looks like the generating distribution. We prove in this setting that the Markov chain is rapidly mixing, i.e., it quickly converges …
Phylogenetic networks are often constructed by merging multiple conflicting phylogenetic signals into a directed acyclic graph. It is interesting to explore whether a network constructed in this way induces biologically-relevant phylogenetic signals that were not present in the input. Here we show that, given a multiple alignment A for a set of taxa X and a rooted phylogenetic network N whose leaves are labelled by X, it is NP-hard to locate the most parsimonious phylogenetic tree displayed by N (with respect to A) even when the level of N - the maximum number of reticulation nodes within a bi…
Maximum likelihood is one of the most widely used techniques to infer evolutionary histories. Although it is thought to be intractable, a proof of its hardness has been lacking. Here, we give a short proof that computing the maximum likelihood tree is NP-hard by exploiting a connection between likelihood and parsimony observed by Tuffley and Steel.
The so-called binary perfect phylogeny with persistent characters has recently been thoroughly studied in computational biology as it is less restrictive than the well known binary perfect phylogeny. Here, we focus on the notion of (binary) persistent characters, i.e. characters that can be realized on a phylogenetic tree by at most one $0 \rightarrow 1$ transition followed by at most one $1 \rightarrow 0$ transition in the tree, and analyze these characters under different aspects. First, we illustrate the connection between persistent characters and Maximum Parsimony, where we characterize p…
Reconciliation methods aim at recovering macro evolutionary events and at localizing them in the species history, by observing discrepancies between gene family trees and species trees. In this article we introduce an Integer Linear Programming (ILP) approach for the NP-hard problem of computing a most parsimonious time-consistent reconciliation of a gene tree with a species tree when dating information on speciations is not available. The ILP formulation, which builds upon the DTL model, returns a most parsimonious reconciliation ranging over all possible datings of the nodes of the species t…
This paper introduces constNJ, the first algorithm for phylogenetic reconstruction of sets of trees with constrained pairwise rooted subtree-prune regraft (rSPR) distance. We are motivated by the problem of constructing sets of trees which must fit into a recombination, hybridization, or similar network. Rather than first finding a set of trees which are optimal according to a phylogenetic criterion (e.g. likelihood or parsimony) and then attempting to fit them into a network, constNJ estimates the trees while enforcing specified rSPR distance constraints. The primary input for constNJ is a co…
In evolutionary biology, genetic sequences carry with them a trace of the underlying tree that describes their evolution from a common ancestral sequence. The question of how many sequence sites are required to recover this evolutionary relationship accurately depends on the model of sequence evolution, the substitution rate, divergence times and the method used to infer phylogenetic history. A particularly challenging problem for phylogenetic methods arises when a rapid divergence event occurred in the distant past. We analyse an idealised form of this problem in which the terminal edges of a…
A Supertree synthesizes the topologies of a set of phylogenetic trees carrying overlapping taxa set. In process, conflicts in the tree topologies are aimed to be resolved with the consensus clades. Such a problem is proved to be NP-hard. Various heuristics on local search, maximum parsimony, graph cut, etc. lead to different supertree approaches, of which the most popular methods are based on analyzing fixed size subtree topologies (such as triplets or quartets). Time and space complexities of these methods, however, depend on the subtree size considered. Our earlier proposed supertree method …
Evolution is a process that is influenced by various environmental factors, e.g. the interactions between different species, genes, and biogeographical properties. Hence, it is interesting to study the combined evolutionary history of multiple species, their genes, and the environment they live in. A common approach to address this research problem is to describe each individual evolution as a phylogenetic tree and construct a tree reconciliation which is parsimonious with respect to a given event model. Unfortunately, most of the previous approaches are designed only either for host-parasite …
Estimating the phylogeny of the genus Homo is entering a new phase of vastly improved data and methodology. There is increasing evidence of 6 to 10 competing species/lineages at any point in the last half million years, making the elucidation of the relationships of individual specimens particularly important. Recent estimates of the phylogeny of key specimens include Waddell (2013, 2014, 2015, 2016), and Mounier et al. (2016). These are made with quite different data (3D skull shapes and discrete morphological characters, respectively) and methods of analysis (unweighted least squares fitting…
Phylogenetic trees are used to model evolution: leaves are labelled to represent contemporary species ("taxa") and interior vertices represent extinct ancestors. Informally, convex characters are measurements on the contemporary species in which the subset of species (both contemporary and extinct) that share a given state, form a connected subtree. Given an unrooted, binary phylogenetic tree T on a set of n >= 2 taxa, a closed (but fairly opaque) expression for the number of convex characters on T has been known since 1992, and this is independent of the exact topology of T. In this note we p…
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 …
Terraces are potentially large sets of trees with precisely the same likelihood or parsimony score, which can be induced by missing sequences in partitioned multi-locus phylogenetic data matrices. The set of trees on a terrace can be characterized by enumeration algorithms or consensus methods that exploit the pattern of partial taxon coverage in the data, independent of the sequence data themselves. Terraces add ambiguity and complexity to phylogenetic inference particularly in settings where inference is already challenging: data sets with many taxa and relatively few loci. In this paper we …
