arXiv subjects

Niels Holtgrefe

Publications and source records attributed to Niels Holtgrefe.

8 recordsLinked to original sources

arXiv2026

On Tree-Network Distinguishability and Full Identifiability of Phylogenetic Networks

Phylogenetic networks generalize phylogenetic trees to evolutionary histories that include reticulate events such as recombination, horizontal gene transfer, and hybridization. Under a Markov model of nucleotide substitution, a phylogenetic network determines a distribution of leaf-patterns. Here, we study the identifiability of the network topology from this distribution under the Jukes-Cantor (JC), Kimura 2-parameter (K2P), and Kimura 3-parameter (K3P) models. Our first result is that the semi-directed network parameter of a level-1 phylogenetic network (modulo redirecting triangles) is fully identifiable under all three models, on a biologically reasonable parameter space in which substitution rates are probabilistic and mixing parameters are non-trivial (i.e., not 0 or 1). In contrast to the generic identifiability established in prior work, this holds at every point of the parameter space, not merely off of a measure-zero subset. Our second result distinguishes phylogenetic networks from phylogenetic trees, on the same parameter space, under JC. We prove that no phylogenetic network and phylogenetic tree can induce the same leaf-pattern distribution unless the network is a tree, possibly augmented with certain substructures called 2-blobs. This means the presence of reticulate evolution creates, in most cases, a detectable signature in the leaf-pattern distribution. More broadly, these results have consequences for identifiability beyond the models and network classes studied here, including for several coalescent-based models.

q-bio.PE↗

arXiv2026

Tractable Maximization of Budgeted Phylogenetic Diversity on Networks Utilizing Node Scanwidth

Background: Identifying a subset of taxa that maximizes phylogenetic diversity is a cornerstone of quantitative conservation planning. Traditionally, phylogenetic diversity is defined over a phylogenetic tree in which leaves resemble present-day taxa and the branch lengths capture the estimated evolutionary distinctiveness. While maximizing phylogenetic diversity is computationally tractable on trees with unit costs, the problem becomes computationally intractable when transitioning to phylogenetic networks or to budgeted versions in which protecting taxa incurs non-homogeneous costs. This paper addresses these two challenges together, providing definitions and a comprehensive analysis of three distinct variants of budgeted phylogenetic diversity on networks.Results: We conduct our study through the lens of a small structural parameter, node scanwidth ($nsw$), which measures the tree-likeness of a phylogenetic network. Given a tree-extension of width $nsw$, we show that two of the considered variants can be optimized in $\mathcal{O}^*(3^{nsw} \cdot B^2)$ time, where $B$ is the budget. For the computationally harder third variant, we provide an algorithm to compute phylogenetic diversity scores in $\mathcal{O}^*(4^{nsw})$ time. We further contribute the first exact algorithms to compute node scanwidth itself. On highly reticulated, simulated networks with several hundred taxa and heterogeneous costs, our implementation computes phylogenetic diversity scores and optimal node scanwidth in fractions of a second, and the budgeted optimization algorithms significantly outperform existing benchmarks previously limited to unit-cost scenarios. Conclusions: Node scanwidth proves to be a small and computable parameter that makes budgeted phylogenetic diversity tractable on realistic networks, scaling comfortably to a thousand taxa. This narrows the gap between reali...

cs.DS↗

arXiv2026

Bounds on the sequence length sufficient to reconstruct binary level-$1$ phylogenetic networks under the CFN model

Phylogenetic trees and networks are graphs used to model evolutionary relationships, with trees representing strictly branching histories and networks allowing for events in which lineages merge, called reticulation events. While the question of data sufficiency has been studied extensively in the context of trees, it remains largely unexplored for networks. In this work we take a first step in this direction by establishing bounds on the amount of genomic data required to reconstruct binary level-$1$ semi-directed phylogenetic networks, which are binary networks in which reticulation events are indicated by directed edges, all other edges are undirected, and cycles are vertex-disjoint. For this class, methods have been developed recently that are statistically consistent. Roughly speaking, such methods are guaranteed to reconstruct the correct network assuming infinitely long genomic sequences. Here we consider the question whether networks from this class can be uniquely and correctly reconstructed from finite sequences. Specifically, we present an inference algorithm that takes as input genetic sequence data, and demonstrate that the sequence length sufficient to reconstruct the correct network with high probability, under the CFN model of evolution, scales logarithmically, polynomially, or polylogarithmically with the number of taxa, depending on the parameter regime. As part of our contribution, we also present novel inference rules for quartet data in the semi-directed phylogenetic network setting.

q-bio.PE↗

arXiv2026

Exact and Heuristic Computation of the Scanwidth of Directed Acyclic Graphs

To measure the tree-likeness of a directed acyclic graph (DAG), a new width parameter that considers the directions of the arcs was recently introduced: scanwidth. We present the first algorithm that efficiently computes the exact scanwidth of general DAGs. For DAGs with one root and scanwidth $k$ it runs in $O(k \cdot n^k \cdot m)$ time. The algorithm also functions as an FPT algorithm with complexity $O(2^{4 \ell - 1} \cdot \ell \cdot n + n^2)$ for phylogenetic networks of level-$\ell$, a type of DAG used to depict evolutionary relationships among species. Our algorithm performs well in practice, being able to compute the scanwidth of synthetic networks up to 30 reticulations and 100 leaves within 500 seconds. Furthermore, we propose a heuristic that obtains an average practical approximation ratio of 1.5 on these networks. While we prove that the scanwidth is bounded from below by the treewidth of the underlying undirected graph, experiments suggest that for networks the parameters are close in practice.

cs.DS↗

arXiv2026

Limits of Kernelization and Parametrization for Phylogenetic Diversity with Dependencies

In the Maximize Phylogenetic Diversity problem, we are given a phylogenetic tree that represents the genetic proximity of species, and we are asked to select a subset of species of maximum phylogenetic diversity to be preserved through conservation efforts, subject to budgetary constraints that allow only k species to be saved. This neglects that it is futile to preserve a predatory species if we do not also preserve at least a subset of the prey it feeds on. Thus, in the Optimizing PD with Dependencies ($ε$-PDD) problem, we are additionally given a food web that represents the predator-prey relationships between species. The goal is to save a set of k species of maximum phylogenetic diversity such that for every saved species, at least one of its prey is also saved. This problem is NP-hard even when the phylogenetic tree is a star. The $α$-PDD problem alters PDD by requiring that at least some fraction $α$ of the prey of every saved species are also saved. In this paper, we study the parameterized complexity of $α$-PDD. We prove that the problem is W[1]-hard and in XP when parameterized by the solution size k, the diversity threshold D, or their complements. When parameterized by the vertex cover number of the food web, $α$-PDD is fixed-parameter tractable (FPT). A key measure of the computational difficulty of a problem that is FPT is the size of the smallest kernel that can be obtained. We prove that, when parameterized by the distance to clique, 1-PDD admits a linear kernel. Our main contribution is to prove that $α$-PDD does not admit a polynomial kernel when parameterized by the vertex cover number plus the diversity threshold D, even if the phylogenetic tree is a star. This implies the non-existence of a polynomial kernel for $α$-PDD also when parameterized by a range of structural parameters of the food web, such as its dist[...]

cs.CC↗

arXiv2025

Characterizing semi-directed phylogenetic networks and their multi-rootable variants

In evolutionary biology, phylogenetic networks are graphs that provide a flexible framework for representing complex evolutionary histories that involve reticulate evolutionary events. Recently phylogenetic studies have started to focus on a special class of such networks called semi-directed networks. These graphs are defined as mixed graphs that can be obtained by de-orienting some of the arcs in some rooted phylogenetic network, that is, a directed acyclic graph whose leaves correspond to a collection of species and that has a single source or root vertex. However, this definition of semi-directed networks is implicit in nature since it is not clear when a mixed-graph enjoys this property or not. In this paper, we introduce novel, explicit mathematical characterizations of semi-directed networks, and also multi-semi-directed networks, that is, mixed graphs that can be obtained from directed phylogenetic networks that may have more than one root. In addition, through extending foundational tools from the theory of rooted networks into the semi-directed setting - such as cherry picking sequences, omnians, and path partitions - we characterize when a (multi-)semi-directed network can be obtained by de-orienting some rooted network that is contained in one of the well-known classes of tree-child, orchard, tree-based or forest-based networks. These results address structural aspects of (multi-)semi-directed networks and pave the way to improved theoretical and computational analyses of such networks, for example, within the development of algebraic evolutionary models that are based on such networks.

q-bio.PE↗

arXiv2025

Distinguishing Phylogenetic Level-2 Networks with Quartets and Inter-Taxon Quartet Distances

The inference of phylogenetic networks, which model complex evolutionary processes including hybridization and gene flow, remains a central challenge in evolutionary biology. Until now, statistically consistent inference methods have been limited to phylogenetic level-1 networks, which allow no interdependence between reticulate events. In this work, we establish the theoretical foundations for a statistically consistent inference method for a much broader class: semi-directed level-2 networks that are outer-labeled planar and galled. We precisely characterize the features of these networks that are distinguishable from the topologies of their displayed quartet trees. Moreover, we prove that an inter-taxon distance derived from these quartets is circular decomposable, enabling future robust inference of these networks from quartet data, such as concordance factors obtained from gene tree distributions under the Network Multispecies Coalescent model. Our results also have novel identifiability implications across different data types and evolutionary models, applying to any setting in which displayed quartets can be distinguished.

q-bio.PE↗

arXiv2025

Reconstructing semi-directed level-1 networks using few quarnets

Semi-directed networks are partially directed graphs that model evolution where the directed edges represent reticulate evolutionary events. We present an algorithm that reconstructs binary $n$-leaf semi-directed level-1 networks in $O( n^2)$ time from its quarnets (4-leaf subnetworks). Our method assumes we have direct access to all quarnets, yet uses only an asymptotically optimal number of $O(n \log n)$ quarnets. When the network is assumed to contain no triangles, our method instead relies only on four-cycle quarnets and the splits of the other quarnets. A variant of our algorithm works with quartets rather than quarnets and we show that it reconstructs most of a semi-directed level-1 network from an asymptotically optimal $O(n \log n)$ of the quartets it displays. Additionally, we provide an $O(n^3)$ time algorithm that reconstructs the tree-of-blobs of any binary $n$-leaf semi-directed network with unbounded level from $O(n^3)$ splits of its quarnets.

q-bio.PE↗