Assessing sampling sufficiency of network metrics using bootstrap

Sampling the full diversity of interactions in an ecological community is a highly intensive effort. Recent studies have demonstrated that many network metrics are sensitive to both sampling effort and network size. Here, we develop a statistical framework, based on bootstrap resampling, that aims to assess sampling sufficiency for some of the most widely used metrics in network ecology, namely connectance, nestedness (NODF-nested overlap and decreasing fill) and modularity (using the QuaBiMo algorithm). Our framework can generate confidence intervals for each network metric with increasing sample size (i.e., the number of sampled interaction events, or number of sampled individuals), which can be used to evaluate sampling sufficiency. The sample is considered sufficient when the confidence limits reach stability or lie within an acceptable level of precision for the aims of the study. We illustrate our framework with data from three quantitative networks of plant and frugivorous birds, varying in size from 16 to 115 species, and 17 to 2,745 interactions. These data sets illustrate that, for the same dataset, sampling sufficiency may be reached at different sample sizes depending on the metric of interest. The bootstrap confidence limits reached stability for the two largest networks, but were wide and unstable with increasing sample size for all three metrics estimated based on the smallest network. The bootstrap method is useful to empirical ecologists to indicate the minimum number of interactions necessary to reach sampling sufficiency for a specific network metric. It is also useful to compare sampling techniques of networks in their capacity to reach sampling sufficiency. Our method is general enough to be applied to different types of metrics and networks.