Popular methods for exploring the space of rooted phylogenetic trees use rearrangement moves such as rNNI (rooted Nearest Neighbour Interchange) and rSPR (rooted Subtree Prune and Regraft). Recently, these moves were generalized to rooted phylogenetic networks, which are a more suitable representation of reticulate evolutionary histories, and it was shown that any two rooted phylogenetic networks of the same complexity are connected by a sequence of either rSPR or rNNI moves. Here, we show that this is possible using only tail moves, which are a restricted version of rSPR moves on networks that are more closely related to rSPR moves on trees. The connectedness still holds even when we restrict to distance-1 tail moves (a localized version of tail-moves). Moreover, we give bounds on the number of (distance-1) tail moves necessary to turn one network into another, which in turn yield new bounds for rSPR, rNNI and SPR (i.e. the equivalent of rSPR on unrooted networks). The upper bounds are constructive, meaning that we can actually find a sequence with at most this length for any pair of networks. Finally, we show that finding a shortest sequence of tail or rSPR moves is NP-hard.
Rooted triples, rooted binary phylogenetic trees on three leaves, are sufficient to encode rooted binary phylogenetic trees. That is, if $\mathcal T$ and $\mathcal T'$ are rooted binary phylogenetic $X$-trees that infers the same set of rooted triples, then $\mathcal T$ and $\mathcal T'$ are isomorphic. However, in general, this sufficiency does not extend to rooted binary phylogenetic networks. In this paper, we show that trinets, phylogenetic network analogues of rooted triples, are sufficient to encode rooted binary orchard networks. Rooted binary orchard networks naturally generalise roote…
While every rooted binary phylogenetic tree is determined by its set of displayed rooted triples, such a result does not hold for an arbitrary rooted binary phylogenetic network. In particular, there exist two non-isomorphic rooted binary temporal normal networks that display the same set of rooted triples. Moreover, without any structural constraint on the rooted phylogenetic networks under consideration, similarly negative results have also been established for binets and trinets which are rooted subnetworks on two and three leaves, respectively. Hence, in general, piecing together a rooted …
Metrics on rooted phylogenetic trees are integral to a number of areas of phylogenetic analysis. Cluster-similarity metrics have recently been introduced in order to limit skew in the distribution of distances, and to ensure that trees in the neighbourhood of each other have similar hierarchies. In the present paper we introduce a new cluster-similarity metric on rooted phylogenetic tree space that has an associated local operation, allowing for easy calculation of neighbourhoods, a trait that is desirable for MCMC calculations. The metric is defined by the distance on the Hasse diagram induce…
A rooted phylogenetic network is a directed acyclic graph with a single root, whose sinks correspond to a set of species. As such networks are useful for representing the evolution of species that have undergone reticulate evolution, there has been great interest in developing the theory behind and algorithms for constructing them. However, unlike evolutionary trees, these networks can be highly non-planar, which can make them difficult to visualise and interpret. Here we investigate properties of planar rooted phylogenetic networks and algorithms for deciding whether or not rooted networks ha…
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. …
Attempting to recognize a tree inside a phylogenetic network is a fundamental undertaking in evolutionary analysis. In the last few years, therefore, tree-based phylogenetic networks, which are defined by a spanning tree called a subdivision tree, have attracted attention of theoretical biologists. However, the application of such networks is still not easy, due to many problems whose time complexities are not clearly understood. In this paper, we provide a general framework for solving those various old or new problems from a coherent perspective, rather than analyzing the complexity of each …
Tree shape statistics are important for investigating evolutionary mechanisms mediating phylogenetic trees. As a step towards bridging shape statistics between rooted and unrooted trees, we present a comparison study on two subtree statistics known as numbers of cherries and pitchforks for the proportional to distinguishable arrangements (PDA) and the Yule-Harding-Kingman (YHK) models. Based on recursive formulas on the joint distribution of the number of cherries and that of pitchforks, it is shown that cherry distributions are log-concave for both rooted and unrooted trees under these two mo…
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,…
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 …
Head moves are a type of rearrangement moves for phylogenetic networks. They have mostly been studied as part of more encompassing types of moves, such as rSPR moves. Here, we study head moves as a type of moves on themselves. We show that the tiers ($k>0$) of phylogenetic network space are connected by local head moves. Then we show tail moves and head moves are closely related: sequences of tail moves can be converted to sequences of head moves and vice versa, changing the length by at most a constant factor. Because the tiers of network space are connected by rSPR moves, this gives a second…
Background. The supertree problem, i.e., the task of finding a common refinement of a set of rooted trees is an important topic in mathematical phylogenetics. The special case of a common leaf set $L$ is known to be solvable in linear time. Existing approaches refine one input tree using information of the others and then test whether the results are isomorphic. Results. A linear-time algorithm, LinCR, for constructing the common refinement $T$ of $k$ input trees with a common leaf set is proposed that explicitly computes the parent function of $T$ in a bottom-up approach. Conclusion. LinCR is…
Phylogenetic networks generalize phylogenetic trees, and have been introduced in order to describe evolution in the case of transfer of genetic material between coexisting species. There are many classes of phylogenetic networks, which can all be modeled as families of graphs with labeled leaves. In this paper, we focus on rooted and unrooted level-k networks and provide enumeration formulas (exact and asymptotic) for rooted and unrooted level-1 and level-2 phylogenetic networks with a given number of leaves. We also prove that the distribution of some parameters of these networks (such as the…
Tree-based networks are a class of phylogenetic networks that attempt to formally capture what is meant by "tree-like" evolution. A given non-tree-based phylogenetic network, however, might appear to be very close to being tree-based, or very far. In this paper, we formalise the notion of proximity to tree-based for unrooted phylogenetic networks, with a range of proximity measures. These measures also provide characterisations of tree-based networks. One measure in particular, related to the nearest neighbour interchange operation, allows us to define the notion of "tree-based rank". This pro…
Phylogenetic networks are rooted directed acyclic graphs that represent evolutionary relationships between species whose past includes reticulation events such as hybridisation and horizontal gene transfer. To search the space of phylogenetic networks, the popular tree rearrangement operation rooted subtree prune and regraft (rSPR) was recently generalised to phylogenetic networks. This new operation - called subnet prune and regraft (SNPR) - induces a metric on the space of all phylogenetic networks as well as on several widely-used network classes. In this paper, we investigate several probl…
Graph invariants are a useful tool in graph theory. Not only do they encode useful information about the graphs to which they are associated, but complete invariants can be used to distinguish between non-isomorphic graphs. Polynomial invariants for graphs such as the well-known Tutte polynomial have been studied for several years, and recently there has been interest to also define such invariants for phylogenetic networks, a special type of graph that arises in the area of evolutionary biology. Recently Liu gave a complete invariant for (phylogenetic) trees. However, the polynomial invariant…
We introduce a notion of finite sampling consistency for phylogenetic trees and show that the set of finitely sampling consistent and exchangeable distributions on n leaf phylogenetic trees is a polytope. We use this polytope to show that the set of all exchangeable and infinite sampling consistent distributions on 4 leaf phylogenetic trees is exactly Aldous' beta-splitting model and give a description of some of the vertices for the polytope of distributions on 5 leaves. We also introduce a new semialgebraic set of exchangeable and sampling consistent models we call the multinomial model and …
A Class of Unrooted Phylogenetic Networks Inspired by the Properties of Rooted Tree-Child 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）；Department of Computer Science, Middlesex University（中伦敦大学计算机科学系）；School of Computer Science, University of Auckland（奥克兰大学计算机科学学院）；Faculty of Computer Science, Dalhousie University（达尔豪斯大学计算机科学学院）
In phylogenetics, tree-based networks are used to model and visualize the evolutionary history of species where reticulate events such as horizontal gene transfer have occurred. Formally, a tree-based network $N$ consists of a phylogenetic tree $T$ (a rooted, binary, leaf-labeled tree) and so-called reticulation edges that span between edges of $T$. The network $N$ is typically visualized by drawing $T$ downward and planar and reticulation edges with one of several different styles. One aesthetic criteria is to minimize the number of crossings between tree edges and reticulation edges. This op…
Phylogenetic networks are a generalisation of phylogenetic trees that allow for more complex evolutionary histories that include hybridisation-like processes. It is of considerable interest whether a network can be considered `tree-like' or not, which lead to the introduction of \textit{tree-based} networks in the rooted, binary context. Tree-based networks are those networks which can be constructed by adding additional edges into a given phylogenetic tree, called the \textit{base tree}. Previous extensions have considered extending to the binary, unrooted case and the nonbinary, rooted case.…
Identifiability of the discrete tree parameter is a key property for phylogenetic models since it is necessary for statistically consistent estimation of the tree from sequence data. Algebraic methods have proven to be very effective at showing that tree and network parameters of phylogenetic models are identifiable, especially when the underlying models are group-based. However, since group-based models are time-reversible, only the unrooted tree topology is identifiable and the location of the root is not. In this note we show that the rooted tree parameter of the Cavender-Farris-Neyman Mode…
Rooted acyclic graphs appear naturally when the phylogenetic relationship of a set $X$ of taxa involves not only speciations but also recombination, horizontal transfer, or hybridization, that cannot be captured by trees. A variety of classes of such networks have been discussed in the literature, including phylogenetic, level-1, tree-child, tree-based, galled tree, regular, or normal networks as models of different types of evolutionary processes. Clusters arise in models of phylogeny as the sets $\mathtt{C}(v)$ of descendant taxa of a vertex $v$. The clustering system $\mathscr{C}_N$ compris…
Rooted phylogenetic networks provide a more complete representation of the ancestral relationship between species than phylogenetic trees when reticulate evolutionary processes are at play. One way to reconstruct a phylogenetic network is to consider its `ancestral profile' (the number of paths from each ancestral vertex to each leaf). In general, this information does not uniquely determine the underlying phylogenetic network. A recent paper considered a new class of phylogenetic networks called `orchard networks' where this uniqueness was claimed to hold. Here we show that an additional rest…
Phylogenetic diversity indices provide a formal way to apportion 'evolutionary heritage' across species. Two natural diversity indices are Fair Proportion (FP) and Equal Splits (ES). FP is also called 'evolutionary distinctiveness' and, for rooted trees, is identical to the Shapley Value (SV), which arises from cooperative game theory. In this paper, we investigate the extent to which FP and ES can differ, characterise tree shapes on which the indices are identical, and study the equivalence of FP and SV and its implications in more detail. We also define and investigate analogues of these ind…
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 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…
Recent investigations in computational biology focus on a family of 2-colored digraphs, called 2-colored best match graphs, which naturally arise from rooted phylogenetic trees. Actually the defining properties of such graphs are unexpectedly unusual in graph theory, and they were established only recently after the discovery of their links to evolutionary relatedness via phylogenetic trees. In this paper several results are obtained on 2-colored best match graphs which well fit in the mainstream of graph theory.
Note on the Maximum Number of Trees Displayed by a Tree-Child Network
Delft Institute of Applied Mathematics, Delft University of Technology（代尔夫特理工大学应用数学研究所）；School of Mathematics and Statistics, University of Canterbury（坎特伯雷大学数学与统计学院）
Throughout the last decade, we have seen much progress towards characterising and computing the minimum hybridisation number for a set P of rooted phylogenetic trees. Roughly speaking, this minimum quantifies the number of hybridisation events needed to explain a set of phylogenetic trees by simultaneously embedding them into a phylogenetic network. From a mathematical viewpoint, the notion of agreement forests is the underpinning concept for almost all results that are related to calculating the minimum hybridisation number for when |P|=2. However, despite various attempts, characterising thi…
Phylogenetic networks are a generalization of phylogenetic trees that are used to represent non-tree-like evolutionary histories that arise in organisms such as plants and bacteria, or uncertainty in evolutionary histories. An \emph{unrooted} phylogenetic network on a nonempty, finite set $X$ of taxa, or \emph{network}, is a connected graph in which every vertex has degree 1 or 3 and whose leaf-set is $X$. It is called a \emph{phylogenetic tree} if the underlying graph is a tree. In this paper we consider properties of \emph{tree-based networks}, that is, networks that can be constructed by ad…
Rooted phylogenetic networks provide an explicit representation of the evolutionary history of a set $X$ of sampled species. In contrast to phylogenetic trees which show only speciation events, networks can also accommodate reticulate processes (for example, hybrid evolution, endosymbiosis, and lateral gene transfer). A major goal in systematic biology is to infer evolutionary relationships, and while phylogenetic trees can be uniquely determined from various simple combinatorial data on $X$, for networks the reconstruction question is much more subtle. Here we ask when can a network be unique…
Phylogenetic networks generalise phylogenetic trees and allow for the accurate representation of the evolutionary history of a set of present-day species whose past includes reticulate events such as hybridisation and lateral gene transfer. One way to obtain such a network is by starting with a (rooted) phylogenetic tree $T$, called a base tree, and adding arcs between arcs of $T$. The class of phylogenetic networks that can be obtained in this way is called tree-based networks and includes the prominent classes of tree-child and reticulation-visible networks. Initially defined for binary phyl…
Galled trees are studied as a recombination model in theoretic population genetics. This class of phylogenetic networks has been generalized to tree-child networks, normal networks and tree-based networks by relaxing a structural condition. Although these networks are simple, their topological structures have yet to be fully understood. It is well-known that all phylogenetic trees on $n$ taxa can be generated by the insertion of the $n$-th taxa to each edge of all the phylogenetic trees on $n-1$ taxa. We prove that all tree-child networks with $k$ reticulate nodes on $n$ taxa can be uniquely g…
A multilabeled tree (or MUL-tree) is a rooted tree in which every leaf is labelled by an element from some set, but in which more than one leaf may be labelled by the same element of that set. In phylogenetics, such trees are used in biogeographical studies, to study the evolution of gene families, and also within approaches to construct phylogenetic networks. A multilabelled tree in which no leaf-labels are repeated is called a phylogenetic tree, and one in which every label is the same is also known as a tree-shape. In this paper, we consider the complexity of computing metrics on MUL-trees …
