How climate extremes - not means - define a species' geographic range boundary via a demographic tipping point

Abstract

Species' geographic range limits interest biologists and resource managers alike; however, scientists lack strong mechanistic understanding of the factors that set geographic range limits in the field, especially for animals. There exists a clear need for detailed case studies that link mechanisms to spatial dynamics and boundaries because such mechanisms allow us to predict whether climate change is likely to change a species' geographic range and, if so, how abundance in marginal populations compares to the core. The bagworm Thyridopteryx ephemeraeformis (Lepidoptera: Psychidae) is a major native pest of cedars, arborvitae, junipers, and other landscape trees throughout much of North America. Across dozens of bagworm populations spread over six degrees of latitude in the American Midwest, we find latitudinal declines in fecundity and egg and pupal survivorship as you proceed towards the northern range boundary. A spatial gradient of bagworm reproductive success emerges, which is associated with a progressive decline in local abundance and an increase in the risk of local population extinction near the species' geographic range boundary. We develop a mathematical model, completely constrained by empirically estimated parameters, to explore the relative roles of reproductive asynchrony and stage-specific survivorship in generating the range limit for this species. We find that overwinter egg mortality is the biggest constraint on bagworm persistence beyond their northern range limit. Overwinter egg mortality is directly related to winter temperatures that fall below the bagworm eggs' physiological limit. This threshold, in conjunction with latitudinal declines in fecundity and pupal survivorship, creates a non-linear response to climate extremes that sets the geographic boundary and provides a path for predicting northward range expansion under altered climate conditions. Our mechanistic modeling approach demonstrates how species' sensitivity to climate extremes can create population tipping points not reflected in demographic responses to climate means, a distinction that is critical to successful ecological forecasting.