Sampling bias is a challenge for quantifying specialization and network structure: lessons from a quantitative niche model

Network approaches have become a popular tool for understanding ecological complexity in a changing world. Many network descriptors relate directly or indirectly to specialization, which is a central concept in ecology and measured in different ways. Unfortunately, quantification of specialization and network structure using field data can suffer from sampling effects. Previous studies evaluating such sampling effects either used field data where the true network structure is unknown, or they simulated sampling based on completely generalized interactions. Here, we used a quantitative niche model to generate bipartite networks representing a wide range of specialization and evaluated potential sampling biases for a large set of specialization and network metrics for different network sizes. We show that with sample sizes realistic for species-rich networks, all metrics are biased towards overestimating specialization (and underestimating generalization and niche overlap). Importantly, this sampling bias depends on the true degree of specialization and is strongest for generalized networks. We show that methods used for empirical data may misrepresent sampling bias: null models simulating generalized interactions may overestimate bias, whereas richness estimators may strongly overestimate sampling completeness. Some network metrics are barely related between small and large sub-samples of the same network and thus may often not be meaningful. Small samples also overestimate interspecific variation of specialization within generalized networks. While new approaches to deal with these challenges have to be developed, we also identify metrics that are relatively unbiased and fairly consistent across sampling intensities and we identify a provisional rule of thumb for the number of observations required for accurate estimates. Our quantitative niche model can help understand variation in network structure capturing both sampling effects and biological meaning. This is needed to connect network science to fundamental ecological theory and to give robust quantitative answers for applied ecological problems.