Borthwick et al. Landscape Complementation and Gene Flow continuous HSM with GSM might perform better because of their combined ability to effectively represent and quantify the continuous degree of landscape complementation (i.e., availability of complementary habitats in vicinity) found at and in-between sites, on which these beetles depend. Our findings may inform future studies that seek to model habitat connectivity in complex heterogeneous landscapes as natural habitats continue to become more fragmented in the Anthropocene.