RESUMO
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.
RESUMO
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.
