Abstract The arthropods and microbes found in trees attacked by the southern pine beetle (Dendroctonus frontalis Zimmermann) have long been referred to as “southern pine beetle associates.” This community includes predators, parasitoids, competitors, mutualists, symbionts, and taxa whose relationship to southern pine beetle may be unclear or not well-described. We present the first systematic review of the southern pine beetle-associated community. We performed a systematic search using the Web of Science Core Collection to identify sources addressing some aspect of southern pine beetle symbiosis or facilitation. Our search returned 231 results; of these, 174 papers fit our criteria for analysis. We also incorporated 198 additional sources from field-specific journals and government publications. From each paper, we extracted taxon names for all organisms identified as southern pine beetle-associates, any description of their ecological role, and the year, location, and study methodology. Our review reveals patterns in the literature and tells the history of this line of research. We present 760 taxa that have been reported as southern pine beetle associates. Not surprisingly given its role as a key natural enemy, the predatory clerid, Thanasimus dubius (Fabricius), was the most reported southern pine beetle associate. The number of publications increased markedly in the 1970s, coinciding with major federal funding for southern pine beetle research. Using the resulting database of southern pine beetle associates and their reported roles, we explored the central question, “what is an associate?” We categorize each southern pine beetle-associate pairing as direct or indirect: these range from observed predation and parasitism (direct association) to co-occurrence in southern pine beetle-attacked trees or in funnel traps (indirect association).
Kanaskie et al. (Tue,) studied this question.