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Specialized natural enemies have long been used to implement the biological control of invasive insects. Although research tracking populations following biological control introductions has traditionally focused on the impact of the introduced agent, recent studies and reviews have reflected an appreciation of the complex interactions of the introduced specialist agents with native generalist natural enemies. These interactions can be neutral, antagonistic, or complementary. Here we studied the invasive defoliator winter moth (Operophtera brumata) in the Northeast USA to investigate the role of native, generalist pupal predators along with the introduced, host‐specific parasitoid Cyzenis albicans. Prior research in Canada has shown that predation of winter moth pupae from native generalists increased after C. albicans was established as a biological control agent. To explain this phenomenon, the following hypotheses were suggested: (H1) parasitoids suppress the winter moth population to a density that can be maintained by generalist predators, (H2) unparasitized pupae are preferred by predators and therefore experience higher mortality rates, or (H3) C. albicans sustains higher predator populations throughout the year more effectively than winter moth alone. We tested these hypotheses by deploying winter moth pupae over 6 years spanning 2005 to 2017 and by modeling pupal predation rates as a function of winter moth density and C. albicans establishment. We also compared predation rates of unparasitized and parasitized pupae and considered additional mortality by a native pupal parasitoid. We found support for the first hypothesis; we detected both temporal and spatial density dependence, but only in the latter years of the study when winter moth densities were low. We found no evidence for the latter two hypotheses. Our findings suggest that pupal predators have a regulatory effect on winter moth populations only after populations have been reduced, presumably by the introduction of the host‐specific parasitoid C. albicans. [ABSTRACT FROM AUTHOR]

The success or failure of an introduced biological control agent may depend on its rate of mortality from disease, predation, and hyperparasitism. Cyzenis albicans Fallén (Diptera: Tachinidae) was introduced to the northeastern U.S. as a biocontrol agent of the invasive species winter moth, Operophtera brumata L. (Lepidoptera: Geometridae). This study aimed to determine the rates of mortality from predation by generalist ground predators and hyperparasitism of C. albicans puparia, identify any hyperparasitoids, and assess the impact of predation and hyperparasitism on the potential success of C. albicans in controlling the winter moth. Mortality of C. albicans puparia was primarily due to predation, but there was also hyperparasitism. Predation and parasitism of C. albicans puparia were consistently high across the six study sites and two years of study, but somewhat lower than was reported from British Columbia, where successful establishment of C. albicans in the 1970s was followed by a decrease in winter moth densities. In this study, three genera of ichneumonid hyperparasitoids were detected and identified using a combination of morphological and molecular approaches: Phygadeuon (1 species), Pimpla (2 species), and Gelis (2 species), all of which contain species with broad host ranges and were likely acting as facultative hyperparsitoids. We conclude that while total mortality of C. albicans puparia is high, it is unlikely to have a significant effect on biological control of winter moth in this system, although it may explain why C . albicans has been slow to establish in this region. Our study emphasizes the importance of assessing the mortality of introduced biological control agents caused by native predators and hyperparasitoids. [ABSTRACT FROM AUTHOR]