Functional attributes of conifers expanding into temperate semi-arid grasslands modulate carbon and nitrogen fluxes in response to prescribed fire

Abstract

Fire exclusion is a key factor driving conifer expansion into temperate semi-arid grasslands. However, it remains unclear how reintroducing fire affects the aboveground storage of carbon (C) and nitrogen (N) in the expanding tree species and belowground in soils. To assess the impacts of fire reintroduction C and N pools and fluxes in areas of conifer expansion we targeted a region of the Northern Great Plains that has experienced extensive woody plant expansion (WPE) of two species: ponderosa pine (Pinus ponderosa) and juniper (Juniperus spp). We quantified tree mortality of both species to estimate the amount of dead biomass C and N produced by a recent prescribed fire, in addition to changes in soil C, pyrogenic C (PyC), and N concentrations across a woody-cover gradient using a before/after/control experimental design. Post-fire soil chemical analysis revealed a 2 year increase in mineral soil C, PyC and N, suggesting the return of fire led to the transfer of partially combusted plant organic matter back to the soil. Further, we found that functional trait differences between the two species influenced the distribution of living conifer biomass-N prior to fire. Despite junipers having 41% less total aboveground biomass than ponderosa, they contained two times more aboveground N. Prescribed fire resulted in 88% mortality of all mature juniper stems and increased fire severity correlated with greater pre-fire juniper cover. Ponderosa mortality varied by size class, with > 40 cm stem diameter class having only 28% mortality. High mortality and greater aboveground N storage in juniper biomass, compared to ponderosa, led to 77% of the total conifer biomass N lost. Consequently, the functional attributes of expanding trees differentially contribute to fluxes of C and N after the return of fire, with junipers acting as conduits for N movement due to their relatively higher N content in less fire-resistant tissues and ponderosa serving as important and more stable storage pools for C. Together, these findings highlight the importance of considering species-specific traits when planning WPE management strrategies at landscape-scales, particularly when goals include C storage or soil nutrient status.

Graphical abstract

The hypothesized effects of how carbon (C) and nitrogen (N) fluxes respond to the return of fire in grasslands experiencing conifer expansion. The question marks represent the input of fire-altered biomass from conifer trees into the soil and the transformation of living to dead tree biomass which are the focus of this study.