Abstract. The literature on cross-layer protocols, protocol improvements, and design methodologies for wireless sensor networks (WSNs) is reviewed and a taxonomy is proposed. The communication protocols devised for WSNs that focus on cross-layer design techniques are reviewed and classified, based on the network layers they aim at replacing in the classical open system interconnection (OSI) network stack. Furthermore, systematic methodologies for the design of cross-layer solution for sensor networks as resource allocation problems in the framework of non-linear optimization are discussed. Open research issues in the development of cross-layer design methodologies for sensor networks are discussed and possible research directions are indicated. Finally, possible shortcomings of cross-layer design techniques such as lack of modularity, decreased robustness, and instability are discussed, and precautionary guidelines are presented.