Uniqueness, intractability and exact algorithms: reflections on level-k phylogenetic networks

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 prove that (1) for all k of at least one it is NP-hard to construct a level-k network consistent with all input triplets, and (2) for all k it is NP-hard to construct a level-k network consistent with a maximum number of input triplets, even when the input is dense. As a response to this intractability we give an exact algorithm for constructing level-1 networks consistent with a maximum number of input triplets

Constructing level-2 phylogenetic networks from triplets

Jansson and Sung showed that, given a dense set of input triplets T (representing hypotheses about the local evolutionary relationships of triplets of species), it is possible to determine in polynomial time whether there exists a level-1 network consistent with T, and if so to construct such a network. They also showed that, unlike in the case of trees (i.e. level-0 networks), the problem becomes NP-hard when the input is non-dense. Here we further extend this work by showing that, when the set of input triplets is dense, the problem is even polynomial-time solvable for the construction of level-2 networks. This shows that, assuming density, it is tractable to construct plausible evolutionary histories from input triplets even when such histories are heavily non-tree like. This further strengthens the case for the use of triplet-based methods in the construction of phylogenetic networks. We also show that, in the non-dense case, the level-2 problem remains NP-hard

A Note on Encodings of Phylogenetic Networks of Bounded Level

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 one of the above collections encode (i.e. uniquely describe) the network that induces it? In this note, we present a complete answer to this question for the special case of a level-1 (phylogenetic) network by characterizing those level-1 networks for which an encoding in terms of one (or equivalently all) of the above collections exists. Given that this type of network forms the first layer of the rich hierarchy of level-k networks, k a non-negative integer, it is natural to wonder whether our arguments could be extended to members of that hierarchy for higher values for k. By giving examples, we show that this is not the case

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 phylogenetic level-1 networks from nondense binet and trinet sets

Binets and trinets are phylogenetic networks with two and three leaves, respectively. Here we consider the problem of deciding if there exists a binary level-1 phylogenetic network displaying a given set T of binary binets or trinets over a taxon set X, and constructing such a network whenever it exists. We show that this is NP-hard for trinets but polynomial-time solvable for binets. Moreover, we show that the problem is still polynomial-time solvable for inputs consisting of binets and trinets as long as the cycles in the trinets have size three. Finally, we present an O(3^{|X|} poly(|X|)) time algorithm for general sets of binets and trinets. The latter two algorithms generalise to instances containing level-1 networks with arbitrarily many leaves, and thus provide some of the first supernetwork algorithms for computing networks from a set of rooted 1 phylogenetic networks

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

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

Level-k Phylogenetic Network can be Constructed from a Dense Triplet Set in Polynomial Time

Given a dense triplet set $\mathcal{T}$, there arise two interesting questions: Does there exists any phylogenetic network consistent with $\mathcal{T}$? And if so, can we find an effective algorithm to construct one? For cases of networks of levels $k=0$ or 1 or 2, these questions were answered with effective polynomial algorithms. For higher levels $k$, partial answers were recently obtained with an $O(|\mathcal{T}|^{k+1})$ time algorithm for simple networks. In this paper we give a complete answer to the general case. The main idea is to use a special property of SN-sets in a level-k network. As a consequence, we can also find the level-k network with the minimum number of reticulations in polynomial time