arXiv subjects

Steven Kelk

Publications and source records attributed to Steven Kelk.

At least 19 recordsLinked to original sources

arXiv2026

A kernel for the maximum agreement forest problem on multiple binary phylogenetic trees

The maximum agreement forest (MAF) problem in phylogenetics takes as input a set t >= 2 of binary phylogenetic trees T on the same set of taxa X. It asks for a partition of X into the smallest number of blocks such that the subtrees induced by these blocks are disjoint and have common topology across all the trees in T. We produce a modified version of the well-known chain reduction rule in order to prove that after exhaustive application of reduction rules each tree has O( t * r * k ) leaves, where k is the natural parameter (the number of blocks) and r=min{max{k,3},t+1}}. We prove this bound for both the unrooted and rooted version of the problem, and demonstrate that the bound r, the length to which common chains are truncated, is tight. Our results constitute the first kernels for MAF in the t>2 regime.

math.CO↗

arXiv2025

Computational support for case-heavy proofs in mathematical phylogenetics

In this note we demonstrate that a number of case-heavy combinatorial proofs in the mathematical phylogenetics literature can be proven more compactly using computational support. We use these techniques to also prove several new combinatorial lemmas that would have taken considerable effort to prove by hand. We are optimistic that similar approaches can be deployed more widely in phylogenetics.

q-bio.PE↗

arXiv2025

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.

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arXiv2025

IMProofBench: Benchmarking AI on Research-Level Mathematical Proof Generation

As the mathematical capabilities of large language models (LLMs) improve, it becomes increasingly important to evaluate their performance on research-level tasks at the frontier of mathematical knowledge. However, existing benchmarks are limited, as they focus solely on final-answer questions or high-school competition problems. To address this gap, we introduce IMProofBench, a private benchmark consisting of 77 peer-reviewed problems developed by expert mathematicians. Each problem requires a detailed proof and is paired with subproblems that have final answers, supporting both an evaluation by human experts and a large-scale quantitative analysis through automated grading. Furthermore, unlike prior benchmarks, the evaluation setup simulates a realistic research environment: models operate in an agentic framework with tools like web search for literature review and mathematical software such as SageMath. Our results show that current LLMs can already solve a significant percentage of research-level questions. IMProofBench will continue to evolve as a dynamic benchmark in collaboration with the mathematical community, ensuring its relevance for evaluating the next generation of LLMs.

cs.CL↗

arXiv2025

A strengthened bound on the number of states required to characterize maximum parsimony distance

In this article we prove that the distance $d_{\mathrm{MP}}(T_1,T_2) = k$ between two unrooted binary phylogenetic trees $T_1, T_2$ on the same set of taxa can be defined by a character that is convex on one of $T_1, T_2$ and which has at most $2k$ states. This significantly improves upon the previous bound of $7k-5$ states. We also show that for every $k \geq 1$ there exist two trees $T_1, T_2$ with $d_{\mathrm{MP}}(T_1,T_2) = k$ such that at least $k+1$ states are necessary in any character that achieves this distance and which is convex on one of $T_1, T_2$. We augment these lower and upper bounds with an empirical analysis which shows that in practice significantly fewer than $k+1$ states are usually required.

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arXiv2025

Coconvex characters on collections of phylogenetic trees

In phylogenetics, a key problem is to construct evolutionary trees from collections of characters where, for a set X of species, a character is simply a function from X onto a set of states. In this context, a key concept is convexity, where a character is convex on a tree with leaf set X if the collection of subtrees spanned by the leaves of the tree that have the same state are pairwise disjoint. Although collections of convex characters on a single tree have been extensively studied over the past few decades, very little is known about coconvex characters, that is, characters that are simultaneously convex on a collection of trees. As a starting point to better understand coconvexity, in this paper we prove a number of extremal results for the following question: What is the minimal number of coconvex characters on a collection of n-leaved trees taken over all collections of size t >= 2, also if we restrict to coconvex characters which map to k states? As an application of coconvexity, we introduce a new one-parameter family of tree metrics, which range between the coarse Robinson-Foulds distance and the much finer quartet distance. We show that bounds on the quantities in the above question translate into bounds for the diameter of the tree space for the new distances. Our results open up several new interesting directions and questions which have potential applications to, for example, tree spaces and phylogenomics.

math.CO↗

arXiv2025

Bounding the SNPR distance between two tree-child networks using generalised agreement forests

Agreement forests continue to play a central role in the comparison of phylogenetic trees since their introduction more than 25 years ago. More specifically, they are used to characterise several distances that are based on tree rearrangement operations and related quantifiers of dissimilarity between phylogenetic trees. In addition, the concept of agreement forests continues to underlie most advancements in the development of algorithms that exactly compute the aforementioned measures. In this paper, we introduce agreement digraphs, a concept that generalises agreement forests for two phylogenetic trees to two phylogenetic networks. Analogous to the way in which agreement forests compute the subtree prune and regraft distance between two phylogenetic trees but inherently more complex, we then use agreement digraphs to bound the subnet prune and regraft distance between two tree-child networks from above and below and show that our bounds are tight.

math.CO↗

arXiv2024

A branch-&-price approach to the unrooted maximum agreement forest problem

We propose the first branch-&-price algorithm for the maximum agreement forest problem on unrooted binary trees: given two unrooted X-labelled binary trees we seek to partition X into a minimum number of blocks such that the induced subtrees are disjoint and have the same topologies in both trees. We provide a dynamic programming algorithm for the weighted maximum agreement subtree problem to solve the pricing problem. When combined with rigorous polynomial-time pre-processing our branch-&-price algorithm exhibits (beyond) state-of-the-art performance.

cs.DS↗

arXiv2024

Split-or-decompose: Improved FPT branching algorithms for maximum agreement forests

Phylogenetic trees are leaf-labelled trees used to model the evolution of species. In practice it is not uncommon to obtain two topologically distinct trees for the same set of species, and this motivates the use of distance measures to quantify dissimilarity. A well-known measure is the maximum agreement forest (MAF): a minimum-size partition of the leaf labels which splits both trees into the same set of disjoint, leaf-labelled subtrees (up to isomorphism after suppressing degree-2 vertices). Computing such a MAF is NP-hard and so considerable effort has been invested in finding FPT algorithms, parameterised by $k$, the number of components of a MAF. The state of the art has been unchanged since 2015, with running times of $O^*(3^k)$ for unrooted trees and $O^*(2.3431^k)$ for rooted trees. In this work we present improved algorithms for both the unrooted and rooted cases, with runtimes $O^*(2.846^k)$ and $O^*(2.3391^k)$ respectively. The key to our improvement is a novel branching strategy in which we show that any overlapping components obtained on the way to a MAF can be `split' by a branching rule with favourable branching factor, and then the problem can be decomposed into disjoint subproblems to be solved separately. We expect that this technique may be more widely applicable to other problems in algorithmic phylogenetics.

cs.DS↗

arXiv2024

A 2-approximation algorithm for the softwired parsimony problem on binary, tree-child phylogenetic networks

Finding the most parsimonious tree inside a phylogenetic network with respect to a given character is an NP-hard combinatorial optimization problem that for many network topologies is essentially inapproximable. In contrast, if the network is a rooted tree, then Fitch's well-known algorithm calculates an optimal parsimony score for that character in polynomial time. Drawing inspiration from this we here introduce a new extension of Fitch's algorithm which runs in polynomial time and ensures an approximation factor of 2 on binary, tree-child phylogenetic networks, a popular topologically-restricted subclass of phylogenetic networks in the literature. Specifically, we show that Fitch's algorithm can be seen as a primal-dual algorithm, how it can be extended to binary, tree-child networks and that the approximation guarantee of this extension is tight. These results for a classic problem in phylogenetics strengthens the link between polyhedral methods and phylogenetics and can aid in the study of other related optimization problems on phylogenetic networks.

q-bio.PE↗

arXiv2024

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.

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arXiv2024

Approximation Ratio of the Min-Degree Greedy Algorithm for Maximum Independent Set on Interval and Chordal Graphs

In this article we prove that the minimum-degree greedy algorithm, with adversarial tie-breaking, is a $(2/3)$-approximation for the Maximum Independent Set problem on interval graphs. We show that this is tight, even on unit interval graphs of maximum degree 3. We show that on chordal graphs, the greedy algorithm is a $(1/2)$-approximation and that this is again tight. These results contrast with the known (tight) approximation ratio of $\frac{3}{\Delta+2}$ of the greedy algorithm for general graphs of maximum degree $\Delta$.

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arXiv2023

Relaxed Agreement Forests

There are multiple factors which can cause the phylogenetic inference process to produce two or more conflicting hypotheses of the evolutionary history of a set X of biological entities. That is: phylogenetic trees with the same set of leaf labels X but with distinct topologies. This leads naturally to the goal of quantifying the difference between two such trees T_1 and T_2. Here we introduce the problem of computing a 'maximum relaxed agreement forest' (MRAF) and use this as a proxy for the dissimilarity of T_1 and T_2, which in this article we assume to be unrooted binary phylogenetic trees. MRAF asks for a partition of the leaf labels X into a minimum number of blocks S_1, S_2, ... S_k such that for each i, the subtrees induced in T_1 and T_2 by S_i are isomorphic up to suppression of degree-2 nodes and taking the labels X into account. Unlike the earlier introduced maximum agreement forest (MAF) model, the subtrees induced by the S_i are allowed to overlap. We prove that it is NP-hard to compute MRAF, by reducing from the problem of partitioning a permutation into a minimum number of monotonic subsequences (PIMS). Furthermore, we show that MRAF has a polynomial time O(log n)-approximation algorithm where n=|X| and permits exact algorithms with single-exponential running time. When at least one of the two input trees has a caterpillar topology, we prove that testing whether a MRAF has size at most k can be answered in polynomial time when k is fixed. We also note that on two caterpillars the approximability of MRAF is related to that of PIMS. Finally, we establish a number of bounds on MRAF, compare its behaviour to MAF both in theory and in an experimental setting and discuss a number of open problems.

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arXiv2023

Agreement forests of caterpillar trees: complexity, kernelization and branching

Given a set $X$ of species, a phylogenetic tree is an unrooted binary tree whose leaves are bijectively labelled by $X$. Such trees can be used to show the way species evolve over time. One way of understanding how topologically different two phylogenetic trees are, is to construct a minimum-size agreement forest: a partition of $X$ into the smallest number of blocks, such that the blocks induce homeomorphic, non-overlapping subtrees in both trees. This comparison yields insight into commonalities and differences in the evolution of $X$ across the two trees. Computing a smallest agreement forest is NP-hard (Hein, Jiang, Wang and Zhang, Discrete Applied Mathematics 71(1-3), 1996). In this work we study the problem on caterpillars, which are path-like phylogenetic trees. We will demonstrate that, even if we restrict the input to this highly restricted subclass, the problem remains NP-hard and is in fact APX-hard. Furthermore we show that for caterpillars two standard reductions rules well known in the literature yield a tight kernel of size at most $7k$, compared to $15k$ for general trees (Kelk and Simone, SIAM Journal on Discrete Mathematics 33(3), 2019). Finally we demonstrate that we can determine if two caterpillars have an agreement forest with at most $k$ blocks in $O^*(2.49^k)$ time, compared to $O^*(3^k)$ for general trees (Chen, Fan and Sze, Theoretical Computater Science 562, 2015), where $O^*(.)$ suppresses polynomial factors.

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arXiv2023

Snakes and Ladders: a Treewidth Story

Let $G$ be an undirected graph. We say that $G$ contains a ladder of length $k$ if the $2 \times (k+1)$ grid graph is an induced subgraph of $G$ that is only connected to the rest of $G$ via its four cornerpoints. We prove that if all the ladders contained in $G$ are reduced to length 4, the treewidth remains unchanged (and that this bound is tight). Our result indicates that, when computing the treewidth of a graph, long ladders can simply be reduced, and that minimal forbidden minors for bounded treewidth graphs cannot contain long ladders. Our result also settles an open problem from algorithmic phylogenetics: the common chain reduction rule, used to simplify the comparison of two evolutionary trees, is treewidth-preserving in the display graph of the two trees.

math.CO↗

arXiv2022

Deep kernelization for the Tree Bisection and Reconnnect (TBR) distance in phylogenetics

We describe a kernel of size 9k-8 for the NP-hard problem of computing the Tree Bisection and Reconnect (TBR) distance k between two unrooted binary phylogenetic trees. We achieve this by extending the existing portfolio of reduction rules with three novel new reduction rules. Two of the rules are based on the idea of topologically transforming the trees in a distance-preserving way in order to guarantee execution of earlier reduction rules. The third rule extends the local neighbourhood approach introduced in (Kelk and Linz, Annals of Combinatorics 24(3), 2020) to more global structures, allowing new situations to be identified when deletion of a leaf definitely reduces the TBR distance by one. The bound on the kernel size is tight up to an additive term. Our results also apply to the equivalent problem of computing a Maximum Agreement Forest (MAF) between two unrooted binary phylogenetic trees. We anticipate that our results will be more widely applicable for computing agreement-forest based dissimilarity measures.

cs.DS↗

arXiv2022

Cyclic generators and an improved linear kernel for the rooted subtree prune and regraft distance

The rooted subtree prune and regraft (rSPR) distance between two rooted binary phylogenetic trees is a well-studied measure of topological dissimilarity that is NP-hard to compute. Here we describe an improved linear kernel for the problem. In particular, we show that if the classical subtree and chain reduction rules are augmented with a modified type of chain reduction rule, the resulting trees have at most 9k-3 leaves, where k is the rSPR distance; and that this bound is tight. The previous best-known linear kernel had size O(28k). To achieve this improvement we introduce cyclic generators, which can be viewed as cyclic analogues of the generators used in the phylogenetic networks literature. As a corollary to our main result we also give an improved weighted linear kernel for the minimum hybridization problem on two rooted binary phylogenetic trees.

cs.DS↗

arXiv2021

Convex characters, algorithms and matchings

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. In \cite{KelkS17} it was shown how to efficiently count, list and sample certain restricted subfamilies of convex characters, and algorithmic applications were given. We continue this work in a number of directions. First, we show how combining the enumeration of convex characters with existing parameterised algorithms can be used to speed up exponential-time algorithms for the \emph{maximum agreement forest problem} in phylogenetics. Second, we re-visit the quantity $g_2(T)$, defined as the number of convex characters on $T$ in which each state appears on at least 2 taxa. We use this to give an algorithm with running time $O( \phi^{n} \cdot \text{poly}(n) )$, where $\phi \approx 1.6181$ is the golden ratio and $n$ is the number of taxa in the input trees, for computation of \emph{maximum parsimony distance on two state characters}. By further restricting the characters counted by $g_2(T)$ we open an interesting bridge to the literature on enumeration of matchings. By crossing this bridge we improve the running time of the aforementioned parsimony distance algorithm to $O( 1.5895^{n} \cdot \text{poly}(n) )$, and obtain a number of new results in themselves relevant to enumeration of matchings on at-most binary trees.

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