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The use of flying platforms such as unmanned aerial vehicles (UAVs), popularly known as drones, is rapidly growing. In particular, with their inherent attributes such as mobility, flexibility, and adaptive altitude, UAVs admit several key potential applications in wireless systems. On the one hand, UAVs can be used as aerial base stations to enhance coverage, capacity, reliability, and energy efficiency of wireless networks. On the other hand, UAVs can operate as flying mobile terminals within a cellular network. Such cellular-connected UAVs can enable several applications ranging from real-time video streaming to item delivery. In this paper, a comprehensive tutorial on the potential benefits and applications of UAVs in wireless communications is presented. Moreover, the important challenges and the fundamental tradeoffs in UAV-enabled wireless networks are thoroughly investigated. In particular, the key UAV challenges such as 3D deployment, performance analysis, channel modeling, and energy efficiency are explored along with representative results. Then, open problems and potential research directions pertaining to UAV communications are introduced. Finally, various analytical frameworks and mathematical tools, such as optimization theory, machine learning, stochastic geometry, transport theory, and game theory are described. The use of such tools for addressing unique UAV problems is also presented. In a nutshell, this tutorial provides key guidelines on how to analyze, optimize, and design UAV-based wireless communication systems.