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The different sciences furnish us with a wide variety of explanations: some work at macroscopic scales, some work at microscopic scales, and some operate across different levels. How do these different explanatory levels relate to one another, and what is an explanatory level in the first place? Over the last 50 years, more and more philosophers--both reductionists and anti-reductionists--no longer subscribe to the idea that the best explanation resides at the fundamental physical level. New challenges arise from the success of scientific explanations employing multi-level models which mix levels of explanation, from distinctive differences between levels structures in biology, cognitive science, and social science, from the apparently radical reimagining of the explanatory role of spacetime in our current best theories of fundamental physics, and from the enduring mystery of how higher-level explanations are possible in the first place. These questions naturally connect to classic philosophical ways of thinking about the relationships between levels: reduction, emergence, and fundamentality. This volume presents a snapshot of cutting-edge research on explanatory levels, from their conceptual foundations to the details of how they are used in scientific practice.