Combining photoperiod and thermal responses to predict phenological mismatch for introduced insects.

A wide variety of organisms use the regular seasonal changes in photoperiod as a cue to align their life cycles with favorable conditions. Yet the phenological consequences of photoperiodism for organisms exposed to new climates are often overlooked. We present a conceptual approach and phenology model that maps voltinism (generations per year) and the degree of phenological mismatch that can arise when organisms with a short-day diapause response are introduced to new regions or are otherwise exposed to new climates. Our degree-day-based model combines continent-wide spatialized daily climate data, calculated date-specific and latitude-specific day lengths, and experimentally determined developmental responses to both photoperiod and temperature. Using the case of the knotweed psyllid Aphalara itadori, a new biological control agent being introduced from Japan to North America and Europe to control an invasive weed, we show how incorporating a short-day diapause response will result in geographic patterns of attempted voltinism that are strikingly different from the potential number of generations based on degree-days alone. The difference between the attempted and potential generations represents a quantitative measure of phenological mismatch between diapause timing and the end of the growing season. We conclude that insects moved from lower to higher latitudes (or to cooler climates) will tend to diapause too late, potentially resulting in high mortality from inclement weather, and those moved from higher to lower latitude (to warmer climates) may be prone to diapausing too early, therefore not fully exploiting the growing season and/or suffering from insufficient reserves for the longer duration in diapause. Mapped output reveals a central region with good phenology match that shifts north or south depending on the geographic source of the insect and its corresponding critical photoperiod for diapause. These results have direct relevance for efforts to establish populations of classical biocontrol agents. More generally, our approach and model could be applied to a wide variety of photoperiod- and temperature-sensitive organisms that are exposed to changes in climate, including resident and invasive agricultural pests and species of conservation concern.

The Effects of Photoperiod on Diapause Induction in Hypena opulenta (Lepidoptera: Erebidae), a Biological Control Agent Against Invasive Swallow-Worts in North America.

Many insects exhibit a short-day diapause response, whereby diapause is induced when daylength falls below a critical threshold. This response is an adaptation to ensure synchrony between periods of insect activity, and the availability of resources, but it can cause problems when organisms are moved to new locations, where early or late-induced diapause can prove a barrier to establishment. We explored the role of photoperiod in diapause induction in Hypena opulenta, a recently introduced classical biological control agent for invasive swallow-worts in North America. We conducted four experimental cage releases as well as a growth chamber experiment to determine the threshold photoperiod for diapause induction in H. opulenta. We determined that the critical photoperiod for inducing diapause in 50% of H. opulenta is 15 h 35 min, which the moth only experiences in the Ottawa release site around summer solstice. This may lead to univoltinism, premature diapause, and poor establishment at some North American release sites. Our results can inform practical aspects of the biological control program for H. opulenta, such as fine-tuning methodologies for stockpiling diapausing pupae in the laboratory and narrowing down the optimal time window for releases at a given location. Additionally, our results will be important for the development of a temperature-based phenology model to more accurately predict voltinism in H. opulenta across the invasive range of swallow-worts in North America.

The influence of matrix habitat on Aphthona flea beetle immigration to leafy spurge patches.

Variation in movement ability by insects among different non-habitat (matrix) types may have important implications for both metapopulation dynamics and weed biocontrol practices. We used a mark-recapture experiment to explore the effects of two different matrix habitats (grass vs shrub) on the ability of two species of Aphthona (Chrysomelidae: Coleoptera) flea beetle to immigrate to patches of the invasive weed, leafy spurge. Using generalized linear models, we compared effects of the matrix habitat types, species and sex on observed immigration probabilities. Our analyses demonstrated that one species (A. nigriscutis) had a much higher immigration probability when moving through a grass-dominated matrix than a shrub-dominated matrix whereas immigration probabilities for the second species (A. lacertosa) were similar in both matrix habitats but significantly lower overall than for A. nigriscutis. Furthermore, A. nigriscutis females were more likely to immigrate to spurge patches embedded in a grass matrix than in shrub, whereas the opposite occurred for males. Our results suggest that metapopulation dynamics may be strongly affected by the type(s) of matrix habitat present on a landscape. These effects also suggest that release strategies for weed biocontrol should be tailored according to the structure of the landscape into which releases are planned. In addition, even closely related species can have significantly different movement abilities which will also affect release strategies.