Lethal combination for seedlings: extreme heat drives mortality of drought-exposed high-elevation pine seedlings.

BACKGROUND AND AIMS: Hotter drought- and biotically-driven tree mortality are expected to increase with climate change in much of the western United States, and species persistence will depend upon ongoing establishment under novel conditions or migration to track ecological niche requirements. High-elevation tree species may be particularly vulnerable to increasing water stress as snowpack declines, increasing the potential for adult mortality and simultaneous regeneration failures. Seedling survival will be determined by ecophysiological limitations in response to changing water availability and temperature. METHODS: We exposed seedlings from populations of Pinus longaeva, Pinus flexilis, and Pinus albicaulis to severe drought and concurrent temperature stress in common gardens testing timing of drought onset under two different temperature regimes. We monitored seedling functional traits, physiological function, and survival. KEY RESULTS: The combined stressors of water limitation and extreme heat led to conservative water use strategies and declines in physiological function, with these joint stressors ultimately exceeding species' tolerances and leading to complete episodic mortality across all species. Growing conditions were the primary determinant of seedling trait expression, with seedlings exhibiting more drought-resistant traits such as lower specific leaf area in the hottest, driest treatment conditions. Water stress-induced stomatal closure was also widely apparent. Under adequate soil moisture, seedlings endured prolonged exposure to high air and surface temperatures, suggesting broad margins for survival. CONCLUSIONS: The critical interaction between soil moisture and temperature suggests that rising temperatures will exacerbate growing season moisture stress. Our results highlight the importance of local conditions over population- and species-level influences in shaping strategies for stress tolerance and resistance to desiccation at this early life stage. By quantifying some of the physiological consequences of drought and heat that lead to seedling mortality, we can better understand the future effects of global change on the composition and distribution of high-elevation conifer forests.

Reforestation of high elevation pines: Direct seeding success depends on seed source and sowing environment.

Forest persistence in regions impacted by increasing water and temperature stress will depend upon species' ability to either rapidly adjust to novel conditions or migrate to track ecological niches. Predicted, rapid climate change is likely to outpace the adaptive and migratory capacity of long-lived isolated tree species, and reforestation may be critical to species' persistence. Facilitating persistence both within and beyond a species' range requires identification of seed lots best adapted to the current and future conditions predicted with rapid climate change. We evaluate variation in emergent seedling performance that leads to differential survival among species and populations for three high elevation five-needle pines. We paired a fully reciprocal field common garden experiment with a greenhouse common garden study to (1) quantify variation in seedling emergence and functional traits, (2) ask how functional traits affect performance under different establishment conditions, and (3) evaluate whether trait and performance variation demonstrates local adaptation and plasticity. Among study species-limber, Great Basin bristlecone, and whitebark pines-we found divergence in emergence and functional traits, though soil moisture was the strongest driver of seedling emergence and abundance across all species. Generalist limber pine had a clear emergence advantage as well as traits associated with drought adaptation, while edaphic specialist bristlecone pine was characterized by low emergence yet high early survival once established. Despite evidence for edaphic specialization, soil characteristics alone did not explain bristlecone success. Across species, trait-environment relationships provided some evidence for local adaptation in drought-adapted traits, but we found no evidence of local adaptation in emergence or survival at this early life stage. For managers looking to promote persistence, sourcing seed from drier environments is likely to impart greater drought resistance into reforestation efforts through strategies such as greater root investment, increasing the probability of early seedling survival. This research demonstrates, through a rigorous reciprocal transplant experimental design, that it may be possible to select climate- and soil-appropriate seed sources for reforestation. However, planting success will ultimately rely on a suitable establishment environment, requiring careful consideration of interannual climate variability for management interventions in these climate and disturbance-impacted tree species.

Persistent yet vulnerable: resurvey of an Abies ecotone reveals few differences but vulnerability to climate change.

Climate change is shifting forest tree species distributions across elevational and latitudinal gradients, and these changes are often pronounced at ecotones where species meet their climatic bounds and are replaced by other species. Using an extensive ecotone composed of lower-montane white fir (Abies concolor var. lowiana) and upper-montane red fir (Abies magnifica var. magnifica) in the central Sierra Nevada range of California, USA, we (1) examined how the demographics of the ecotone have responded to recent climate using a field observational study and a historical dataset, (2) quantified climate drivers across species life stages using contemporary demographic data, and (3) tested the potential impacts of future climate on species-specific seedling survival and growth in a fully factorial growth chamber experiment that varied temperature, growing season length, and water availability. A re-examination of the ecotone midpoint after 35 yr suggested a reduction in A. concolor sapling and tree densities and a rise in A. magnifica proportional dominance between surveys. Seedling abundances across the ecotone indicated that A. magnifica tends to dominate the regeneration layer and currently forms an important component of the seedling community at elevations below those where A. magnifica saplings or trees begin to co-dominate stands. Observational and experimental assessments suggest that temperature and precipitation serve as important drivers, differentiating A. concolor vs. A. magnifica distributions, and are primary stressors at the seedling stage. Seedlings of both species were adversely affected by experimental climate treatments, although A. concolor exhibited greater survival and a more conservative growth strategy under extreme climatic stress than A. magnifica. Projections indicate that historical climate conditions will rise by an amount greater than the ecotone's current elevational extent by the end of the 21st century. Differential drivers of species abundances suggest that the projected climate will expand conditions that promote A. concolor abundance and impede A. magnifica abundance across the ecotone; however, disturbance activity and microclimatic conditions will also influence regeneration and overstory tree dynamics. Our study demonstrates the importance of quantifying species-specific responses to climate and indicates that widespread regeneration failure may be one possible consequence in which species exhibit strong sensitivity to projected climate conditions.

From canopy to seed: Loss of snow drives directional changes in forest composition.

Climate change is altering the conditions for tree recruitment, growth, and survival, and impacting forest community composition. Across southeast Alaska, USA, and British Columbia, Canada, Callitropsis nootkatensis (Alaska yellow-cedar) is experiencing extensive climate change-induced canopy mortality due to fine-root death during soil freezing events following warmer winters and the loss of insulating snowpack. Here, we examine the effects of ongoing, climate-driven canopy mortality on forest community composition and identify potential shifts in stand trajectories due to the loss of a single canopy species. We sampled canopy and regenerating forest communities across the extent of C. nootkatensis decline in southeast Alaska to quantify the effects of climate, community, and stand-level drivers on C. nootkatensis canopy mortality and regeneration as well as postdecline regenerating community composition. Across the plot network, C. nootkatensis exhibited significantly higher mortality than co-occurring conifers across all size classes and locations. Regenerating community composition was highly variable but closely related to the severity of C. nootkatensis mortality. Callitropsis nootkatensis canopy mortality was correlated with winter temperatures and precipitation as well as local soil drainage, with regenerating community composition and C. nootkatensis regeneration abundances best explained by available seed source. In areas of high C. nootkatensis mortality, C. nootkatensis regeneration was low and replaced by Tsuga. Our study suggests that climate-induced forest mortality is driving alternate successional pathways in forests where C. nootkatensis was once a major component. These pathways are likely to lead to long-term shifts in forest community composition and stand dynamics. Our analysis fills a critical knowledge gap on forest ecosystem response and rearrangement following the climate-driven decline of a single species, providing new insight into stand dynamics in a changing climate. As tree species across the globe are increasingly stressed by climate change-induced alteration of suitable habitat, identifying the autecological factors contributing to successful regeneration, or lack thereof, will provide key insight into forest resilience and persistence on the landscape.