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Wireless sensor networks (WSNs) use small nodes with constrained capabilities to sense, collect, and disseminate information in many types of applications. As sensor networks become wide-spread, security issues become a central concern, especially in mission-critical tasks. In this paper, we identify the threats and vulnerabilities to WSNs and summarize the defense methods based on the networking protocol layer analysis first. Then we give a holistic overview of security issues. These issues are divided into seven categories: cryptography, key management, attack detections and preventions, secure routing, secure location security, secure data fusion, and other security issues. Along the way we analyze the advantages and disadvantages of current secure schemes in each category. In addition, we also summarize the techniques and methods used in these categories, and point out the open research issues and directions in each area.