When two trees go to war

Rooted phylogenetic networks are often constructed by combining trees, clusters, triplets or characters into a single network that in some well-defined sense simultaneously represents them all. We review these four models and investigate how they are related. In general, the model chosen influences the minimum number of reticulation events required. However, when one obtains the input data from two binary trees, we show that the minimum number of reticulations is independent of the model. The number of reticulations necessary to represent the trees, triplets, clusters (in the softwired sense) and characters (with unrestricted multiple crossover recombination) are all equal. Furthermore, we show that these results also hold when not the number of reticulations but the level of the constructed network is minimised. We use these unification results to settle several complexity questions that have been open in the field for some time. We also give explicit examples to show that already for data obtained from three binary trees the models begin to diverge

A Practical Algorithm for Reconstructing Level-1 Phylogenetic Networks

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 exact and a greedy algorithm both of which are of independent theoretical interest. Most importantly, LEV1ATHAN runs in polynomial time and always constructs a level-1 network. If the data is consistent with a phylogenetic tree, then the algorithm constructs such a tree. Moreover, if the input triplet set is dense and, in addition, is fully consistent with some level-1 network, it will find such a network. The potential of LEV1ATHAN is explored by means of an extensive simulation study and a biological data set. One of our conclusions is that LEV1ATHAN is able to construct networks consistent with a high percentage of input triplets, even when these input triplets are affected by a low to moderate level of noise

Reconstructing a phylogenetic level-1 network from quartets

We describe a method that will reconstruct an unrooted binary phylogenetic level-1 network on n taxa from the set of all quartets containing a certain fixed taxon, in O(n^3) time. We also present a more general method which can handle more diverse quartet data, but which takes O(n^6) time. Both methods proceed by solving a certain system of linear equations over GF(2). For a general dense quartet set (containing at least one quartet on every four taxa) our O(n^6) algorithm constructs a phylogenetic level-1 network consistent with the quartet set if such a network exists and returns an (O(n^2) sized) certificate of inconsistency otherwise. This answers a question raised by Gambette, Berry and Paul regarding the complexity of reconstructing a level-1 network from a dense quartet set

Phylogenetic Networks Do not Need to Be Complex: Using Fewer Reticulations to Represent Conflicting Clusters

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 reticulate events such as hybridizations, recombinations and horizontal gene transfers. Here we present the new CASS algorithm that can combine any set of clusters into a phylogenetic network. We show that the networks constructed by CASS are usually simpler than networks constructed by other available methods. Moreover, we show that CASS is guaranteed to produce a network with at most two reticulations per biconnected component, whenever such a network exists. We have implemented CASS and integrated it in the freely available Dendroscope software

Hierarchies from lowest stable ancestors in nonbinary phylogenetic networks

The reconstruction of the evolutionary history of a set of species is an important problem in classification and phylogenetics. Phylogenetic networks are a generalization of evolutionary trees that are used to represent histories for species that have undergone reticulate evolution, an important evolutionary force for many organisms (e.g. plants or viruses). In this paper, we present a novel approach to understanding the structure of networks that are not necessarily binary. More specifically, we define the concept of a closed set and show that the collection of closed sets of a network forms a hierarchy, and that this hierarchy can be deduced from either the subtrees or subnetworks on all 3-subsets. This allows us to also show that closed sets generalize the concept of the SN-sets of a binary network, sets which have proven very useful in elucidating the structure of binary networks. We also characterize the minimal closed sets (under set inclusion) for a special class of networks (2-terminal networks). Taken together, we anticipate that our results should be useful for the development of new phylogenetic network reconstruction algorithms

A Survey of Combinatorial Methods for Phylogenetic Networks

The evolutionary history of a set of species is usually described by a rooted phylogenetic tree. Although it is generally undisputed that bifurcating speciation events and descent with modifications are major forces of evolution, there is a growing belief that reticulate events also have a role to play. Phylogenetic networks provide an alternative to phylogenetic trees and may be more suitable for data sets where evolution involves significant amounts of reticulate events, such as hybridization, horizontal gene transfer, or recombination. In this article, we give an introduction to the topic of phylogenetic networks, very briefly describing the fundamental concepts and summarizing some of the most important combinatorial methods that are available for their computation