Organismal responses to artificial light at night in terrestrial ecosystems vary across lunar cycles

Globally, nocturnal lightscapes are now determined by both moonlight and light pollution, or artificial light at night (ALAN). Organisms respond both to changes in moonlight across lunar cycles and to alterations in light conditions due to artificial light. The interaction of natural and artificial light is a critical aspect to incorporate into our understanding of how crepuscular and nocturnal ecology is altered in anthropogenically-modified landscapes. In this manuscript we review the rapidly expanding body of research on ecological impacts of ALAN to (1) assess patterns of lunar data inclusion and (2) summarize documented interactions of moonlight and ALAN. Three-fourths (72%) of 379 papers reviewed did not incorporate moonlight into their statistical analyses and experimental design. Only 12% directly investigated interaction effects of moonlight and ALAN. However, 70% of these studies reported an interactive effect. Considering this stark contrast, as a precursor to our literature review, we present an overview of moonlight and the lunar cycle for biologists. The overarching trend emerging from the literature is that biological impacts of ALAN decrease with increasing moonlight, although the opposite is true in some cases. After summarizing the literature, we present general hypotheses regarding the interaction of the lunar cycle and ALAN. These hypotheses consider the different forms of ALAN encountered by organisms (i.e. skyglow and light sources) and account for the influence of cloud cover. Finally, we suggest best practices for incorporating moonlight into future research on biological impacts of ALAN.