Embracing scale-dependence to achieve a deeper understanding of biodiversity and its change across communities

Abstract Because biodiversity is multidimensional and scale-dependent, it is challenging to estimate its change. However, it is unclear (1) how much scale-dependence matters for empirical studies, and (2) if it does matter, how exactly we should quantify biodiversity change. To address the first question, we analyzed studies with comparisons among multiple assemblages, and found that rarefaction curves frequently crossed, implying reversals in the ranking of species richness across spatial scales. Moreover, the most frequently measured aspect of diversity—species richness—was poorly correlated with other measures of diversity. Second, we collated studies that included spatial scale in their estimates of biodiversity change in response to ecological drivers and found frequent and strong scale-dependence, including nearly 10% of studies which showed that biodiversity changes switched directions across scales. Having established the complexity of empirical biodiversity comparisons, we describe a synthesis of methods based on rarefaction curves that allow more explicit analyses of spatial and sampling effects on biodiversity comparisons. We use a case study of nutrient additions in experimental ponds to illustrate how this multi-dimensional and multi-scale perspective informs the responses of biodiversity to ecological drivers. Statement of Authorship JC and BM conceived the study and the overall approach, and all authors participated in multiple working group meetings to develop and refine the approach. BM collected the data for the meta-analysis that led to Fig. 2,3; JC collected the data for the metaanalysis that led to Figure 4 and S1; SB and FM did the analyses for Figures 2-4; DM, FM and XX wrote the code for the analysis used for the recipe and case study in Figure 6. JC, BM and NG wrote first drafts of most sections, and all authors contributed substantially to revisions. Figure 1. A. Individual-based rarefaction curves of three hypothetical communities (labelled A,B, C) where ranked differences between communities are consistent across scales. B. Individual-based rarefaction curves of three hypothetical communities (labelled A,B, C) where rankings between communities switch because of differences in the total numbers of species, and their relative abundances. Dotted vertical lines illustrate sampling scales where rankings switch. These curves were generated using the sim_sad function from the mobsim R package (May et al. 2018). Figure 2. Bivariate relationships between N, S PIE and S for 346 communities across the 37 datasets taken from McGill (2011b)(see Appendix 1). (A) S as a function of N; (B) S as a function of S PIE . (N vs S PIE not shown). Black lines depict the relationships across studies (and correspond to R 2 fixed); colored points and lines show the relationships within studies. All axes are log-scale. Insets are histograms of the study-level slopes, with the solid line representing the slope across all studies. Gray bars indicate the study-level slope did not differ from zero, blue indicates a significant positive slope, and red indicates a significant negative slope. Figure 3. Representative rarefaction curves, the proportion of curves that crossed, and counts of how often curves crossed. (A) Rarefaction curves for different local communities within two datasets: marine invertebrates (nematodes) along a gradient from a waste plant outlet (Lambshead 1986), and trees in a Ugandan rainforest (Eggeling 1947); axes are log-transformed. (B) Counts of how many times pairs of rarefaction curves (from the same community) crossed; y-axis is on a log-scale. Data accessibility statement All data for meta-analyses and case study will be deposited in a publically available repository with DOI upon acceptance (available in link for submission).