Responses of bryosphere fauna to drought across a boreal forest chronosequence.

Projected changes in precipitation regimes can greatly impact soil biota, which in turn alters key ecosystem functions. In moss-dominated ecosystems, the bryosphere (i.e., the ground moss layer including live and senesced moss) plays a key role in carbon and nutrient cycling, and it hosts high abundances of microfauna (i.e., nematodes and tardigrades) and mesofauna (i.e., mites and springtails). However, we know very little about how bryosphere fauna responds to precipitation, and whether this response changes across environmental gradients. Here, we used a mesocosm experiment to study the effect of volume and frequency of precipitation on the abundance and community composition of functional groups of bryosphere fauna. Hylocomium splendens bryospheres were sampled from a long-term post-fire boreal forest chronosequence in northern Sweden which varies greatly in environmental conditions. We found that reduced precipitation promoted the abundance of total microfauna and of total mesofauna, but impaired predaceous/omnivorous nematodes, and springtails. Generally, bryosphere fauna responded more strongly to precipitation volume than to precipitation frequency. For some faunal functional groups, the effects of precipitation frequency were stronger at reduced precipitation volumes. Context-dependency effects were found for microfauna only: microfauna was more sensitive to precipitation in late-successional forests (i.e., those with lower productivity and soil nutrient availability) than in earlier-successional forests. Our results also suggest that drought-induced changes in trophic interactions and food resources in the bryosphere may increase faunal abundance. Consequently, drier bryospheres that may result from climate change could promote carbon and nutrient turnover from fauna activity, especially in older, less productive forests.

Effects of plant functional group removal on CO2 fluxes and belowground C stocks across contrasting ecosystems.

Changes in plant communities can have large effects on ecosystem carbon (C) dynamics and long-term C stocks. However, how these effects are mediated by environmental context or vary among ecosystems is not well understood. To study this, we used a long-term plant removal experiment set up across 30 forested lake islands in northern Sweden that collectively represent a strong gradient of soil fertility and ecosystem productivity. We measured forest floor CO2 exchange and aboveground and belowground C stocks for a 22-yr experiment involving factorial removal of the two dominant functional groups of the boreal forest understory, namely ericaceous dwarf shrubs and feather mosses, on each of the 30 islands. We found that long-term shrub and moss removal increased forest floor net CO2 loss and decreased belowground C stocks consistently across the islands irrespective of their productivity or soil fertility. However, we did see context-dependent responses of respiration to shrub removals because removals only increased respiration on islands of intermediate productivity. Both CO2 exchange and C stocks responded more strongly to shrub removal than to moss removal. Shrub removal reduced gross primary productivity of the forest floor consistently across the island gradient, but it had no effect on respiration, which suggests that loss of belowground C caused by the removals was driven by reduced litter inputs. Across the island gradient, shrub removal consistently depleted C stocks in the soil organic horizon by 0.8 kg C/m2 . Our results show that the effect of plant functional group diversity on C dynamics can be relatively consistent across contrasting ecosystems that vary greatly in productivity and soil fertility. These findings underline the key role of understory vegetation in forest C cycling, and suggest that global change leading to changes in the relative abundance of both shrubs and mosses could impact on the capacity of boreal forests to store C.

Increased fire severity alters initial vegetation regeneration across Calluna-dominated ecosystems.

Calluna vulgaris-dominated habitats are valued for ecosystem services such as carbon storage and for their conservation importance. Climate and environmental change are altering their fire regimes. In particular, more frequent summer droughts will result in higher severity wildfires. This could alter the plant community composition of Calluna habitats and thereby influence ecosystem function. To study the effect of fire severity on community composition we used rain-out shelters to simulate drought prior to experimental burns at two Calluna-dominated sites, a raised bog and a heathland. We analysed species abundance in plots surveyed ca. 16 months after fire in relation to burn severity (indicated by fire-induced soil heating). We found that fire severity was an important control on community composition at both sites. Higher fire severity increased the abundance of ericoids, graminoids and acrocarpous mosses, and decreased the abundance of pleurocarpous mosses compared to lower severity fires. At the raised bog, the keystone species Sphagnum capillifolium and Eriophorum vaginatum showed no difference in regeneration with fire severity. Species and plant functional type beta-diversity increased following fire, and was similar in higher compared to lower severity burns. Our results further our understanding of the response of Calluna-dominated habitats to projected changes in fire regimes, and can assist land managers using prescribed fires in selecting burning conditions to achieve management objectives.

Burning increases post-fire carbon emissions in a heathland and a raised bog, but experimental manipulation of fire severity has no effect.

Large amounts of carbon are stored in northern peatlands. There is concern that greater wildfire severity following projected increases in summer drought will lead to higher post-fire carbon losses. We measured soil carbon dynamics in a Calluna heathland and a raised peat bog after experimentally manipulating fire severity. A gradient of fire severity was achieved by simulating drought in 2 × 2 m plots. Ecosystem respiration (ER), net ecosystem exchange (NEE), methane (CH4) flux and concentration of dissolved organic carbon ([DOC], measured at the raised bog only) were measured for up to two years after burning. The response of these carbon fluxes to increased fire severity in drought plots was similar to plots burnt under ambient conditions associated with traditional managed burning. Averaged across all burnt plots, burning altered mean NEE from a net carbon sink at the heathland (-0.33 µmol CO2 m-2 s-1 in unburnt plots) to a carbon source (0.50 µmol m-2 s-1 in burnt plots) and at the raised bog (-0.38 and 0.16 µmol m-2 s-1, respectively). Burning also increased CH4 flux at the raised bog (from 1.16 to 25.3 nmol m-2 s-1 in the summer, when it accounted for 79% of the CO2-equivalent emission). Burning had no significant effect on soil water [DOC].

Fire severity is more sensitive to low fuel moisture content on Calluna heathlands than on peat bogs.

Moorland habitats dominated by the dwarf shrub Calluna vulgaris provide important ecosystem services. Drought is projected to intensify throughout their range, potentially leading to increased fire severity as moisture is a key control on severity. We studied the effect of low fuel moisture content (FMC) on fire severity by using 2×2m rain-out shelters prior to completing 19 experimental fires in two sites in Scotland (UK): a dry heath with thin organic soils and a raised bog with deep, saturated peat, both dominated by Calluna vulgaris. Reduced FMC of the moss and litter (M/L) layer at both sites, and the soil moisture of the dry heath, increased fire-induced consumption of the M/L layer and soil heating at both sites. Increase in fire severity was greater at the dry heath than at the raised bog, e.g. average maximum temperatures at the soil surface increased from 31°C to 189°C at the dry heath, but only from 10°C to 15°C at the raised bog. Substantial M/L layer consumption was observed when its FMC was below 150%. This led to larger seasonal and daily soil temperature fluctuation, particularly at the dry heath during warm months. The results suggest that low FMC following predicted changes in climate are likely to increase wildfire severity and that the impact on vegetation composition and carbon stores may be greater at heathlands than at peatlands. Managed burning aiming to minimise fire severity (e.g. ignition of the M/L layer and exposure to lethal temperatures of ericoid seeds) should be carried out when the FMC of the M/L layer is above 150% and the FMC of the soil is above 200-300%.

Leaving moss and litter layers undisturbed reduces the short-term environmental consequences of heathland managed burns.

Variation in the structure of ground fuels, i.e. the moss and litter (M/L) layer, may be an important control on fire severity in heather moorlands and thus influence vegetation regeneration and soil carbon dynamics. We completed experimental fires in a Calluna vulgaris-dominated heathland to study the role of the M/L layer in determining (i) fire-induced temperature pulses into the soil and (ii) post-fire soil thermal dynamics. Manually removing the M/L layer before burning increased fire-induced soil heating, both at the soil surface and 2 cm below. Burnt plots where the M/L layer was removed simulated the fuel structure after high severity fires where ground fuels are consumed but the soil does not ignite. Where the M/L layer was manually removed, either before or after the fire, post-fire soil thermal dynamics showed larger diurnal and seasonal variation, as well as similar patterns to those observed after wildfires, compared to burnt plots where the M/L layer was not manipulated. We used soil temperatures to explore potential changes in post-fire soil respiration. Simulated high fire severity (where the M/L layer was manually removed) increased estimates of soil respiration in warm months. With projected fire regimes shifting towards higher severity fires, our results can help land managers develop strategies to balance ecosystem services in Calluna-dominated habitats.