Phylogenetic diversity (PD) depends on sampling intensity, which complicates the comparison of PD between samples of different depth. One approach to dealing with differing sample depth for a given diversity statistic is to rarefy, which means to take a random subset of a given size of the original sample. Exact analytical formulae for the mean and variance of species richness under rarefaction have existed for some time but no such solution exists for PD. We have derived exact formulae for the mean and variance of PD under rarefaction. We show that these formulae are correct by comparing exact solution mean and variance to that calculated by repeated random (Monte Carlo) subsampling of a dataset of stem counts of woody shrubs of Toohey Forest, Queensland, Australia. We also demonstrate the application of the method using two examples: identifying hotspots of mammalian diversity in Australasian ecoregions, and characterising the human vaginal microbiome. There is a very high degree of correspondence between the analytical and random subsampling methods for calculating mean and variance of PD under rarefaction, although the Monte Carlo method requires a large number of random draws to converge on the exact solution for the variance. Rarefaction of mammalian PD of ecoregions in Australasia to a common standard of 25 species reveals very different rank orderings of ecoregions, indicating quite different hotspots of diversity than those obtained for unrarefied PD. The application of these methods to the vaginal microbiome shows that a classical score used to quantify bacterial vaginosis is correlated with the shape of the rarefaction curve. The analytical formulae for the mean and variance of PD under rarefaction are both exact and more efficient than repeated subsampling. Rarefaction of PD allows for many applications where comparisons of samples of different depth is required.
The Fair Proportion of a species in a phylogenetic tree is a very simple measure that has been used to assess its value relative to the overall phylogenetic diversity represented by the tree. It has recently been proved by Fuchs and Jin to be equal to the Shapley Value of the coallitional game that sends each subset of species to its rooted Phylogenetic Diversity in the tree. We prove in this paper that this result extends to the natural translations of the Fair Proportion and the rooted Phylogenetic Diversity to rooted phylogenetic networks. We also generalize to rooted phylogenetic networks …
In biodiversity conservation it is often necessary to prioritize the species to conserve. Existing approaches to prioritization, e.g. the Fair Proportion Index and the Shapley Value, are based on phylogenetic trees and rank species according to their contribution to overall phylogenetic diversity. However, in many cases evolution is not treelike and thus, phylogenetic networks have come to the fore as a generalization of phylogenetic trees, allowing for the representation of non-treelike evolutionary events, such as horizontal gene transfer or hybridization. Here, we extend the concepts of phy…
Phylogenetic Diversity (PD) is a prominent quantitative measure of the biodiversity of a collection of present-day species (taxa). This measure is based on the evolutionary distance among the species in the collection. Loosely speaking, if $\mathcal{T}$ is a rooted phylogenetic tree whose leaf set $X$ represents a set of species and whose edges have real-valued lengths (weights), then the PD score of a subset $S$ of $X$ is the sum of the weights of the edges of the minimal subtree of $\mathcal{T}$ connecting the species in $S$. In this paper, we define several natural variants of the PD score …
It is generally accepted that "diversity" is associated with success in evolutionary algorithms. However, diversity is a broad concept that can be measured and defined in a multitude of ways. To date, most evolutionary computation research has measured diversity using the richness and/or evenness of a particular genotypic or phenotypic property. While these metrics are informative, we hypothesize that other diversity metrics are more strongly predictive of success. Phylogenetic diversity metrics are a class of metrics popularly used in biology, which take into account the evolutionary history …
Phylogenetic diversity indices provide a formal way to apportion 'evolutionary heritage' across species. Two natural diversity indices are Fair Proportion (FP) and Equal Splits (ES). FP is also called 'evolutionary distinctiveness' and, for rooted trees, is identical to the Shapley Value (SV), which arises from cooperative game theory. In this paper, we investigate the extent to which FP and ES can differ, characterise tree shapes on which the indices are identical, and study the equivalence of FP and SV and its implications in more detail. We also define and investigate analogues of these ind…
Phylogenetic diversity indices such as the Fair Proportion (FP) index are frequently discussed as prioritization criteria in biodiversity conservation. They rank species according to their contribution to overall diversity by taking into account the unique and shared evolutionary history of each species as indicated by its placement in an underlying phylogenetic tree. Traditionally, phylogenetic trees were inferred from single genes and the resulting gene trees were assumed to be a valid estimate for the species tree, i.e., the "true" evolutionary history of the species under consideration. Ho…
In conservation biology, phylogenetic diversity (PD) provides a way to quantify the impact of the current rapid extinction of species on the evolutionary `Tree of Life'. This approach recognises that extinction not only removes species but also the branches of the tree on which unique features shared by the extinct species arose. In this paper, we investigate three questions that are relevant to PD. The first asks how many sets of species of given size $k$ preserve the maximum possible amount of PD in a given tree. The number of such maximum PD sets can be very large, even for moderate-sized p…
Planning for the protection of species often involves difficult choices about which species to prioritize, given constrained resources. One way of prioritizing species is to consider their "evolutionary distinctiveness", i.e. their relative evolutionary isolation on a phylogenetic tree. Several evolutionary isolation metrics or phylogenetic diversity indices have been introduced in the literature, among them the so-called Fair Proportion index (also known as the "evolutionary distinctiveness" score). This index apportions the total diversity of a tree among all leaves, thereby providing a simp…
Properties of biodiversity indices that incorporate future extinction risk
University of Canterbury（坎特伯雷大学）；New Jersey Institute of Technology（新泽理工学院）；Northwestern University（西北大学）；The University of Chicago（芝加哥大学）；Simon Fraser University（西蒙菲莎大学）
We describe the use of the Frechet mean and variance in the Billera-Holmes-Vogtmann (BHV) treespace to summarize and explore the diversity of a set of phylogenetic trees. We show that the Frechet mean is comparable to other summary methods, and, despite its stickiness property, is more likely to be binary than the majority-rules consensus tree. We show that the Frechet variance is faster and more precise than commonly used variance measures. The Frechet mean and variance are more theoretically justified, and more robust, than previous estimates of this type, and can be estimated reasonably eff…
The Shapley Value and the Fair Proportion Index of phylogenetic trees have been frequently discussed as prioritization tools in conservation biology. Both indices rank species according to their contribution to total phylogenetic diversity, allowing for a simple conservation criterion. While both indices have their specific advantages and drawbacks, it has recently been shown that both values are closely related. However, as different authors use different definitions of the Shapley Value, the specific degree of relatedness depends on the specific version of the Shapley Value - it ranges from …
