Behavior of class-level landscape metrics across gradients of class aggregation and area

Habitat loss and fragmentation processes strongly affect biodiversity conservation in landscapes undergoing anthropogenic land use changes. Many attempts have been made to use landscape structure metrics to quantify the independent and joint effects of these processes. Unfortunately, ecological interpretation of those metrics has been plagued by lack of thorough understanding of their theoretical behavior. We explored behavior of 50 metrics in neutral landscapes across a 21-step gradient in aggregation and a 19-step gradient in area using a full factorial design with 100 replicates of each of the 399 combinations of the two factors to assess how well metrics reflected changes in landscape structure. Metric values from real landscapes were used to determine the extent of neutral landscape space that is represented in real landscapes. We grouped metrics into three major behavioral classes: strongly related to focal class area (n = 15), strongly related to aggregation (n = 7), and jointly responding to area and aggregation (n = 28). Metrics strongly related to class area exhibited a variety of distinct behaviors, and many of these metrics have unique interpretations that make each of them particularly useful in certain applications. Metrics strongly related to aggregation, independent of class area, are particularly useful in assessing effects of fragmentation. Moreover, metrics in this group exhibited a range of specific behaviors, highlighting subtle but different aspects of landscape aggregation even though we controlled only one aspect of aggregation. The non-linear behavior exhibited by many metrics renders interpretation difficult and use of linear analytical techniques inappropriate under many circumstances. Ultimately, comprehensive characterization of landscapes undergoing habitat loss and fragmentation will require using several metrics distributed across behavioral groups.