Defoliation-induced changes in foliage quality may trigger broad-scale insect outbreaks.

Top-down effects, like predation, are drivers of insect outbreaks, but bottom-up effects, like host nutritional quality, also influence outbreaks and could in turn be altered by insect-caused defoliation. We evaluated the prediction that herbivory leads to a positive feedback on outbreak severity as nutrient concentration in plant tissues increases through improved soil nutrient availability from frass and litter deposition. Over seven years of a spruce budworm outbreak, we quantified litter nutrient fluxes, soil nitrogen availability, and host tree foliar nutrient status along a forest susceptibility gradient. As the outbreak progressed, both soil nutrient fluxes and availability increased which, in turn, improved foliage quality in surviving host trees. This is consistent with boosted insect fitness and increased population density and defoliation as outbreaks grow. Our results suggest that a positive bottom-up feedback to forest ecosystems from defoliation may result in conditions favorable to self-amplifying population dynamics in insect herbivores that can contribute to driving broad-scale outbreaks.

Improved performance of the eastern spruce budworm on black spruce as warming temperatures disrupt phenological defences.

Phenological shifts, induced by global warming, can lead to mismatch between closely interacting species. The eastern spruce budworm, Choristoneura fumiferana, an important outbreaking insect defoliator in North America, mainly feeds on balsam fir, Abies balsamea, which has historically been well synchronized with the insect. But as climate change pushes the northern range limit of the budworm further north into the boreal forest, the highly valuable black spruce, Picea mariana, historically protected against the budworm by its late budburst phenology, is suffering increased defoliation during the current outbreak. We tested the hypothesis that rising temperatures can lead, not to a mismatch, but to an improved match between the budworm and black spruce through differential phenological advancement. For 3 years, eastern spruce budworm larvae were reared from instar 2 to pupae, on both black spruce and balsam fir, in a temperature free-air controlled enhancement experiment (T-FACE) consisting in 24 field plots, half of which were heated at +2°C from March to October. Our results show that every year, larval development was faster on heated trees and pupation was earlier than on unheated trees. Bud development was also accelerated in heated trees of both species. However, there was no difference in mass between pupae that developed at +2°C and controls at the end of the season. Finally, we found no difference either in development rate or pupal mass between larvae reared on black spruce and those reared on balsam fir. This suggests that under higher temperature regimes, eastern spruce budworm will be as successful on black spruce as on balsam fir, as black spruce budburst becomes better synchronized with the insect's emergence from diapause. This could lead to critical changes in outbreak dynamics and severity with important ecological state shifts at the landscape level.

Phenological synchrony between eastern spruce budworm and its host trees increases with warmer temperatures in the boreal forest.

Climate change is predicted to alter relationships between trophic levels by changing the phenology of interacting species. We tested whether synchrony between two critical phenological events, budburst of host species and larval emergence from diapause of eastern spruce budworm, increased at warmer temperatures in the boreal forest in northeastern Canada. Budburst was up to 4.6 ± 0.7 days earlier in balsam fir and up to 2.8 ± 0.8 days earlier in black spruce per degree increase in temperature, in naturally occurring microclimates. Larval emergence from diapause did not exhibit a similar response. Instead, larvae emerged once average ambient temperatures reached 10°C, regardless of differences in microclimate. Phenological synchrony increased with warmer microclimates, tightening the relationship between spruce budworm and its host species. Synchrony increased by up to 4.5 ± 0.7 days for balsam fir and up to 2.8 ± 0.8 days for black spruce per degree increase in temperature. Under a warmer climate, defoliation could potentially begin earlier in the season, in which case, damage on the primary host, balsam fir may increase. Black spruce, which escapes severe herbivory because of a 2-week delay in budburst, would become more suitable as a resource for the spruce budworm. The northern boreal forest could become more vulnerable to outbreaks in the future.

Model-specification uncertainty in future forest pest outbreak.

Climate change will modify forest pest outbreak characteristics, although there are disagreements regarding the specifics of these changes. A large part of this variability may be attributed to model specifications. As a case study, we developed a consensus model predicting spruce budworm (SBW, Choristoneura fumiferana [Clem.]) outbreak duration using two different predictor data sets and six different correlative methods. The model was used to project outbreak duration and the uncertainty associated with using different data sets and correlative methods (=model-specification uncertainty) for 2011-2040, 2041-2070 and 2071-2100, according to three forcing scenarios (RCP 2.6, RCP 4.5 and RCP 8.5). The consensus model showed very high explanatory power and low bias. The model projected a more important northward shift and decrease in outbreak duration under the RCP 8.5 scenario. However, variation in single-model projections increases with time, making future projections highly uncertain. Notably, the magnitude of the shifts in northward expansion, overall outbreak duration and the patterns of outbreaks duration at the southern edge were highly variable according to the predictor data set and correlative method used. We also demonstrated that variation in forcing scenarios contributed only slightly to the uncertainty of model projections compared with the two sources of model-specification uncertainty. Our approach helped to quantify model-specification uncertainty in future forest pest outbreak characteristics. It may contribute to sounder decision-making by acknowledging the limits of the projections and help to identify areas where model-specification uncertainty is high. As such, we further stress that this uncertainty should be strongly considered when making forest management plans, notably by adopting adaptive management strategies so as to reduce future risks.