Different factors limit early- and late-season windows of opportunity for monarch development.

Seasonal windows of opportunity are intervals within a year that provide improved prospects for growth, survival, or reproduction. However, few studies have sufficient temporal resolution to examine how multiple factors combine to constrain the seasonal timing and extent of developmental opportunities. Here, we document seasonal changes in milkweed (Asclepias fascicularis)-monarch (Danaus plexippus) interactions with high resolution throughout the last three breeding seasons prior to a precipitous single-year decline in the western monarch population. Our results show early- and late-season windows of opportunity for monarch recruitment that were constrained by different combinations of factors. Early-season windows of opportunity were characterized by high egg densities and low survival on a select subset of host plants, consistent with the hypothesis that early-spring migrant female monarchs select earlier-emerging plants to balance a seasonal trade-off between increasing host plant quantity and decreasing host plant quality. Late-season windows of opportunity were coincident with the initiation of host plant senescence, and caterpillar success was negatively correlated with heatwave exposure, consistent with the hypothesis that late-season windows were constrained by plant defense traits and thermal stress. Throughout this study, climatic and microclimatic variations played a foundational role in the timing and success of monarch developmental windows by affecting bottom-up, top-down, and abiotic limitations. More exposed microclimates were associated with higher developmental success during cooler conditions, and more shaded microclimates were associated with higher developmental success during warmer conditions, suggesting that habitat heterogeneity could buffer the effects of climatic variation. Together, these findings show an important dimension of seasonal change in milkweed-monarch interactions and illustrate how different biotic and abiotic factors can limit the developmental success of monarchs across the breeding season. These results also suggest the potential for seasonal sequences of favorable or unfavorable conditions across the breeding range to strongly affect monarch population dynamics.

The complexity of global change and its effects on insects.

Global change includes multiple overlapping and interacting drivers: 1) climate change, 2) land use change, 3) novel chemicals, and 4) the increased global transport of organisms. Recent studies have documented the complex and counterintuitive effects of these drivers on the behavior, life histories, distributions, and abundances of insects. This complexity arises from the indeterminacy of indirect, non-additive and combined effects. While there is wide consensus that global change is reorganizing communities, the available data are limited. As the pace of anthropogenic changes outstrips our ability to document its impacts, ongoing change may lead to increasingly unpredictable outcomes. This complexity and uncertainty argue for renewed efforts to address the fundamental drivers of global change.

Long-term exposure to higher temperature increases the thermal sensitivity of grazer metabolism and movement.

Ecological studies of global warming impacts have many constraints. Organisms are often exposed to higher temperatures for short periods of time, probably underestimating their ability to acclimate or adapt relative to slower but real rates of warming. Many studies also focus on a limited number of traits and miss the multifaceted effects that warming may have on organisms, from physiology to behaviour. Organisms exhibit different movement traits, some of which are primarily driven by metabolic processes and others by decision-making, which should influence the extent to which temperature affects them. We collected snails from streams that have been differentially heated by geothermal activity for decades to determine how long-term exposure to different temperatures affected their metabolism and movement. Additionally, we collected snails from a cold stream (5°C) and measured their metabolism and movement at higher temperatures (short-term exposure). We used respirometry to measure metabolic rates and automated in situ image-based tracking to quantify several movement traits from 5 to 21°C. Long-term exposure to higher temperatures resulted in a greater thermal sensitivity of metabolic rate compared to snails exposed for short durations, highlighting the need for caution when conducting acute temperature exposures in global warming research. Average speed, which is largely driven by metabolism, also increased more with temperature for long-term exposure compared to short-term exposure. Movement traits we interpret as more decision-based, such as time spent moving and trajectory shape, were less affected by temperature. Step length increased and step angle decreased at higher temperatures for both long- and short-term exposure, resulting in overall straighter trajectories. The power-law exponent of the step length distributions and fractal dimension of trajectories were independent of temperature, however, suggesting that snails retained the same movement strategy. The observed changes in snail movement at higher temperatures should lead to higher encounter rates and more efficient searching, providing a behavioural mechanism for stronger plant-herbivore interactions in warmer environments. Our research is among the first to show that temperature has contrasting effects on different movement traits, which may be determined by the metabolic contribution to those behaviours.