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How the ED Method Converts PD-Dissimilarities to Estimates of Gains and Losses

“ED” refers to a specific family of “environmental diversity” calculations (Faith and Walker 1996a, b, c; Faith 2003; Faith et al. 2003, 2004). ED typically uses an environmental gradients space, derived using species compositional dissimilarities and ordination methods (Faith and Walker 1996a, b, c). ED has been implemented as a surrogates strategy in biodiversity conservation-planning software that evaluates nominated sets of localities or finds best sites to add to an existing set. For example, ED provided the first integration of 'costs' into regional biodiversity planning based on comparing gains or 'ED-complementarity' values to marginal costs to facilitate trade-offs, balancing biodiversity conservation and other needs of society (Faith et al. 1996).

In order to understand the applicability of ED to PD-dissimilarities, we have to consider ED's assumptions and then examine a simple example analysis. I referred above to unimodal response (Fig. 1b) and the shared-habitat/shared-features companion model to PD's shared-ancestry/shared-features model. ED explicitly builds on this general unimodal response of species (or other elements) to environmental gradients (for background, see Austin 1985; Faith et al. 1987). ED's environmental space typically is derived using compositional dissimilarities (including those estimated GDM) and ordination methods (for review, see Faith et al. 2004). The dissimilarities, the ordination methods and GDM all are relatively robust approaches under a general model of unimodal responses to environmental gradients (Fig. 1b; Faith et al. 1987; Faith and Walker 1996a; Ferrier et al. 2009).

The unimodal response model not only guides the inference of an environmental space using ordination methods (Faith et al. 1987), but also defines how ED methods should effectively sample that environmental space in order to capture biodiversity. ED is based on the idea that many different species (or other elements of biodiversity) respond to similar environmental gradients, and exhibit a general unimodal response at different positions along those gradients (Fig.1b). It follows that effective representation of these gradients (say, by a proposed set of protected areas) should deliver good representation of the various species or phylogenetic branches.

The assumption of a general unimodal response model directly leads to the use of p-median (and related) criteria for ED's estimation of the number of species represented by a given set of localities in the environmental space or ordination. A p-median criterion is based on a sum of the distances in an environmental space. Each distance in this summation is that between a hypothetical point ('demand point') in the space and its nearest site (among all sites in some selected subset). The selected sites, for example, might be nominated protected-area localities. ED is defined based on this calculation. The 'continuous' version of ED refers to the case where the demand points are hypothetical points distributed uniformly throughout the continuous environmental space. Faith and Walker (1994, 1996a) demonstrated that, under a simple unimodal response model, species representation will be maximised by a selected set of sites if and only if it satisfies this continuous p-median criterion. Note that the ED score, because it counts un-represented species based on a sum of distances, is numerically small when the number of represented species is large (see example calculations below and in Faith and Walker 1996a). The ED surrogates approach therefore provides a rationale for interpreting high environmental diversity for a set of localities as implying high biodiversity for the set (see Beier and Albuquerque 2015).

I referred above to the p-median and related criteria. ED is not defined by any a priori choice of the p-median criterion. Instead, the various ED calculations emerge from the assumption of an underlying unimodal response model. In the simple case, unimodal response implies that features are effectively counted up when we apply calculations linked to the p-median; in other cases, the model implies calculations that are modifications of the simple p-median. Simple ED variants include weighting of demand points when species richness varies over the space (Faith and Walker 1996a; Faith et al. 2004), and creating an extended environmental space ('extended polytope'; Faith and Walker 1994, 1996a, b; Faith et al. 2004; see also Hortal et al. 2009). These options modify the parameters used in calculating the p-median. In a later section, I will consider an ED variant that departs from p-median in order to capture expected diversity or persistence.

When extended to features and branches from a phylogeny, the unimodal response model supports an expectation that ED is compatible with Bray-Curtis type PD-dissimilarities. Does this unimodal model (as idealised in Fig. 1b) apply when the elements are branches or features? Certainly, this relationship can be expected, given that PD-dissimilarity operates as if it is a standard Bray Curtis dissimilarity, but applied to features, not species. The robust ordination of such dissimilarities should produce general unimodal responses, as in the species-level case (Faith et al. 1987).

PD and PD-dissimilarities are commonly applied to molecular phylogenetic trees and microbial community data; here, PD analyses overcome the typical absence of defined microbial species. However, there has not been any clear model linking branches to gradients in such studies. Faith et al. (2009) presented an example documenting unimodal response of branches based on a gradient space for microbial communities, sampled in house dust (Fig. 2; Rintala et al. 2008). In Fig. 2, arrows at the right side indicate major gradients revealed by the ordination of the PD-dissimilarities. The solid dots in the space indicate different communities or sample localities. A sample locality represents the branch corresponding to a given family if the locality has one or more descendants of that branch in the phylogeny (for details see Rintala et al. 2008; Faith et al. 2009).

For the ordination space of Rintala et al., Faith et al. (2009) showed that all but 3 of the 56 phylogenetic branches (corresponding to identified families) have a clear unimodal response in the gradients space. Here, a response was recorded as unimodal only if a simple shape could enclose all sample sites representing the given branch (and not include any other sites). This unimodal response for phylogenetic features or branches is a critical property: it provides theoretical justification for GDM on PD-dissimilarities and it accords with the assumptions of the ED (environmental diversity) method.

Extending this example, I now will illustrate the application of the ED method to the PD-based environmental space of Rintala et al. (Fig. 3). In Fig. 3a, the space (from Fig. 2) is filled with ED “demand points”. In Fig. 3b, the ED value is calculated as the sum of the distances from each demand point to its nearest sample/site. In Fig. 3c, sample site x is assumed lost and ED is re-calculated. In Fig. 3d, alternatively, sample site y is lost and ED is re-calculated. We can see from the plots that the loss of sample x clearly results in a greater sum of distances. The loss of sample/ site x would imply much greater loss of phylogenetic diversity compared to loss of sample/site y, as indicated by the amount of change in the sums of distances (Fig. 3c,d). This result corresponds to the intuition that sample x, in filling a larger gap in the space relative to sample y, is likely to uniquely represent more features.

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