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Defining phylogenetic networks using ancestral profiles

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 Added by Charles Semple
 Publication date 2020
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and research's language is English




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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 restriction on the network, that of being `stack-free, is required in order for the original uniqueness claim to hold. On the other hand, if the additional stack-free restriction is lifted, we establish an alternative result; namely, there is uniqueness within the class of orchard networks up to the resolution of vertices of high in-degree.



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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 uniquely reconstructed from its `ancestral profile (the number of paths from each ancestral vertex to each element in $X$). We show that reconstruction holds (even within the class of all networks) for a class of networks we call `orchard networks, and we provide a polynomial-time algorithm for reconstructing any orchard network from its ancestral profile. Our approach relies on establishing a structural theorem for orchard networks, which also provides for a fast (polynomial-time) algorithm to test if any given network is of orchard type. Since the class of orchard networks includes tree-sibling tree-consistent networks and tree-child networks, our result generalise reconstruction results from 2008 and 2009. Orchard networks allow for an unbounded number $k$ of reticulation vertices, in contrast to tree-sibling tree-consistent networks and tree-child networks for which $k$ is at most $2|X|-4$ and $|X|-1$, respectively.
In the study of rooted phylogenetic networks, analyzing the set of rooted phylogenetic trees that are embedded in such a network is a recurring task. From an algorithmic viewpoint, this analysis almost always requires an exhaustive search of a particular multiset $S$ of rooted phylogenetic trees that are embedded in a rooted phylogenetic network $mathcal{N}$. Since the size of $S$ is exponential in the number of reticulations of $mathcal{N}$, it is consequently of interest to keep this number as small as possible but without loosing any element of $S$. In this paper, we take a first step towards this goal by introducing the notion of a non-essential arc of $mathcal{N}$, which is an arc whose deletion from $mathcal{N}$ results in a rooted phylogenetic network $mathcal{N}$ such that the sets of rooted phylogenetic trees that are embedded in $mathcal{N}$ and $mathcal{N}$ are the same. We investigate the popular class of tree-child networks and characterize which arcs are non-essential. This characterization is based on a family of directed graphs. Using this novel characterization, we show that identifying and deleting all non-essential arcs in a tree-child network takes time that is cubic in the number of leaves of the network. Moreover, we show that deciding if a given arc of an arbitrary phylogenetic network is non-essential is $Pi_2^P$-complete.
Tree-child networks, one of the prominent network classes in phylogenetics, have been introduced for the purpose of modeling reticulate evolution. Recently, the first author together with Gittenberger and Mansouri (2019) showed that the number ${rm TC}_{ell,k}$ of tree-child networks with $ell$ leaves and $k$ reticulation vertices has the first-order asymptotics [ {rm TC}_{ell,k}sim c_kleft(frac{2}{e}right)^{ell}ell^{ell+2k-1},qquad (ellrightarrowinfty). ] Moreover, they also computed $c_k$ for $k=1,2,$ and $3$. In this short note, we give a second approach to the above result which is based on a recent (algorithmic) approach for the counting of tree-child networks due to Cardona and Zhang (2020). This second approach is also capable of giving a simple, closed-form expression for $c_k$ for all $kgeq 0$.
In previous work, we gave asymptotic counting results for the number of tree-child and normal networks with $k$ reticulation vertices and explicit exponential generating functions of the counting sequences for $k=1,2,3$. The purpose of this note is two-fold. First, we make some corrections to our previous approach which overcounted the above numbers and thus gives erroneous exponential generating functions (however, the overcounting does not effect our asymptotic counting results). Secondly, we use our (corrected) exponential generating functions to derive explicit formulas for the number of tree-child and normal networks with $k=1,2,3$ reticulation vertices. This re-derives recent results of Carona and Zhang, answers their question for normal networks with $k=2$, and adds new formulas in the case $k=3$.
Rooted phylogenetic networks provide a way to describe species relationships when evolution departs from the simple model of a tree. However, networks inferred from genomic data can be highly tangled, making it difficult to discern the main reticulation signals present. In this paper, we describe a natural way to transform any rooted phylogenetic network into a simpler canonical network, which has desirable mathematical and computational properties, and is based only on the visible nodes in the original network. The method has been implemented and we demonstrate its application to some examples.
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