How does habitat fragmentation affect the biodiversity and ecosystem functioning relationship?

Context: The relationship between biodiversity and ecosystem functioning (BEF) has been a central topic in ecology for more than 20 years. While experimental and theoretical studies have produced much knowledge of how biodiversity affects ecosystem functioning, it remains poorly understood how habitat fragmentation affects the BEF relationship. Objectives: To develop a framework that connects habitat fragmentation to the BEF relationship from a landscape perspective. Methods: We reviewed the literature on habitat fragmentation, BEF, and related fields, and developed a framework to analyze how habitat fragmentation affects the BEF relationship through altering biodiversity, environmental conditions, and both, based on the pattern-process-scale perspective in landscape ecology. Results: Our synthesis of the literature suggests that habitat fragmentation can alter BEF relationship through several processes. First, habitat fragmentation causes the non-random loss of species that make major contributions to ecosystem functioning (decreasing sampling effect), and reduces mutualistic interactions (decreasing complementarity effects) regardless of the changes in species richness. Second, environmental conditions within patches and ecological flows among patches vary significantly with the degree of fragmentation, which potentially contributes to and modulates the BEF relationship. Conclusions: Habitat fragmentation can affect the BEF relationship directly by altering community composition, as well as indirectly by changing environmental conditions within and among habitat patches on both local and landscape levels. The BEF relationship obtained from small plots and over short time periods may not fully represent that in real landscapes that are fragmented, dynamic, and continuously influenced by myriad human activities on different scales in time and space.