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Shifting range limits are predicted for many species as the climate warms. However, the rapid pace of climate change will challenge the natural dispersal capacity of long-lived, sessile organisms such as forest trees. Adaptive responses of populations will, therefore, depend on levels of genetic variation and plasticity for climate-responsive traits, which likely vary across the range due to expansion history and current patterns of selection. Here, we study levels of genetic and plastic variation for phenology and growth traits in populations of red spruce ( Picea rubens ), from the range core to the highly fragmented trailing edge. We measured more than 5000 offspring sampled from three genetically distinct regions (core, margin and edge) grown in three common gardens replicated along a latitudinal gradient. Genetic variation in phenology and growth showed low to moderate heritability and differentiation among regions, suggesting some potential to respond to selection. Phenology traits were highly plastic, but this plasticity was generally neutral or maladaptive in the effect on growth, revealing a potential liability under warmer climates. These results suggest future climate adaptation will depend on the regional availability of genetic variation in red spruce and provide a resource for the design and management of assisted gene flow. This article is part of the theme issue ‘Species’ ranges in the face of changing environments (Part II)’. 

Understanding the factors influencing the current distribution of genetic diversity across a species range is one of the main questions of evolutionary biology, especially given the increasing threat to biodiversity posed by climate change. Historical demographic processes such as population expansion or bottlenecks and decline are known to exert a predominant influence on past and current levels of genetic diversity, and revealing this demo‐genetic history can have immediate conservation implications. We used a whole‐exome capture sequencing approach to analyze polymorphism across the gene space of red spruce (Picea rubens Sarg.), an endemic and emblematic tree species of eastern North America high‐elevation forests that are facing the combined threat of global warming and increasing human activities. We sampled a total of 340 individuals, including populations from the current core of the range in northeastern USA and southeastern Canada and from the southern portions of its range along the Appalachian Mountains, where populations occur as highly fragmented mountaintop “sky islands.” Exome capture baits were designed from the closely relative white spruce (P. glauca Voss) transcriptome, and sequencing successfully captured most regions on or near our target genes, resulting in the generation of a new and expansive genomic resource for studying standing genetic variation in red spruce applicable to its conservation. Our results, based on over 2 million exome‐derived variants, indicate that red spruce is structured into three distinct ancestry groups that occupy different geographic regions of its highly fragmented range. Moreover, these groups show small Ne , with a temporal history of sustained population decline that has been ongoing for thousands (or even hundreds of thousands) of years. These results demonstrate the broad potential of genomic studies for revealing details of the demographic history that can inform management and conservation efforts of nonmodel species with active restoration programs, such as red spruce. 

Abstract. Fire frequency exerts a fundamental control on productivity and nutrientcycling in savanna ecosystems. Individual fires often increaseshort-term nitrogen (N) availability to plants, but repeated burningcauses ecosystem N losses and can ultimately decrease soil organicmatter and N availability. However, these effects remain poorlyunderstood due to limited long-term biogeochemical data. Here, weevaluate how fire frequency and changing vegetation compositioninfluenced wood stable N isotopes (δ15N) across space andtime at one of the longest running prescribed burn experimentsin the world (established in 1964). We developed multiple δ15N recordsacross a burn frequency gradient from precisely dated Quercus macrocarpa tree rings in an oak savanna at Cedar Creek EcosystemScience Reserve, Minnesota, USA. Sixteen trees were sampled across fourtreatment stands that varied with respect to the temporal onset of burning and burnfrequency but were consistent in overstory species representation, soilcharacteristics, and topography. Burn frequency ranged from an unburnedcontrol stand to a high-fire-frequency stand that had burned in 4 ofevery 5 years during the past 55 years. Because N stocks and net Nmineralization rates are currently lowest in frequently burned stands,we hypothesized that wood δ15N trajectories would declinethrough time in all burned stands, but at a rate proportional to the firefrequency. We found that wood δ15N records within each standwere remarkably coherent in their mean state and trend through time. Agradual decline in wood δ15N occurred in the mid-20thcentury in the no-, low-, and medium-fire stands, whereas there was notrend in the high-fire stand. The decline in the three stands did notsystematically coincide with the onset of prescribed burning. Thus, wefound limited evidence for variation in wood δ15N that couldbe attributed directly to long-term fire frequency in this prescribedburn experiment in temperate oak savanna. Our wood δ15Nresults may instead reflect decadal-scale changes in vegetationcomposition and abundance due to early- to mid-20th-century firesuppression. 

Fire activity is changing dramatically across the globe, with uncertain effects on ecosystem processes, especially below‐ground. Fire‐driven losses of soil carbon (C) are often assumed to occur primarily in the upper soil layers because the repeated combustion of above‐ground biomass limits organic matter inputs into surface soil. However, C losses from deeper soil may occur if frequent burning reduces root biomass inputs of C into deep soil layers or stimulates losses of C via leaching and priming.

To assess the effects of fire on soil C, we sampled 12 plots in a 51‐year‐long fire frequency manipulation experiment in a temperate oak savanna, where variation in prescribed burning frequency has created a gradient in vegetation structure from closed‐canopy forest in unburned plots to open‐canopy savanna in frequently burned plots.

Soil C stocks were nonlinearly related to fire frequency, with soil C peaking in savanna plots burned at an intermediate fire frequency and declining in the most frequently burned plots. Losses from deep soil pools were significant, with the absolute difference between intermediately burned plots versus most frequently burned plots more than doubling when the full 1 m sample was considered rather than the top 0–20 cm alone (losses of 98.5 Mg C/ha [−76%] and 42.3 Mg C/ha [−68%] in the full 1 m and 0–20 cm layers respectively). Compared to unburned forested plots, the most frequently burned plots had 65.8 Mg C/ha (−58%) less C in the full 1 m sample. Root biomass below the top 20 cm also declined by 39% with more frequent burning. Concurrent fire‐driven losses of nitrogen and gains in calcium and phosphorus suggest that burning may increase nitrogen limitation and play a key role in the calcium and phosphorus cycles in temperate savannas.

Synthesis. Our results illustrate that fire‐driven losses in soil C and root biomass in deep soil layers may be critical factors regulating the net effect of shifting fire regimes on ecosystem C in forest‐savanna transitions. Projected changes in soil C with shifting fire frequencies in savannas may be 50% too low if they only consider changes in the topsoil.