Summary1. Despite recent interest in linkages between above-and below-ground communities and their consequences for ecosystem processes, much remains unknown about their responses to long-term ecosystem change. We synthesize multiple lines of evidence from a long-term 'natural experiment' to illustrate how ecosystem retrogression (the decline in ecosystem process rates due to long-term absence of major disturbance) drives vegetation change, and thus above-ground and below-ground carbon (C) sequestration, and communities of consumer biota. 2. Our study system involves 30 islands in Swedish boreal forest that form a 5000-year, fire-driven retrogressive chronosequence. Here, retrogression leads to lower plant productivity and slower decomposition and a community shift from plants with traits associated with resource acquisition to those linked with resource conservation. 3. We present consistent evidence that above-ground ecosystem C sequestration declines, while below-ground and total C storage increases linearly for at least 5000 years following fire absence. This increase is driven primarily by changes in vegetation characteristics, impairment of decomposer organisms and absence of humus combustion. 4. Data from contrasting trophic groups show that during retrogression, biomass or abundance of plants and decomposer biota decreases, while that of above-ground invertebrates and birds increases, due to different organisms accessing resources via distinct energy channels. Meanwhile, diversity measures of vascular plants and above-ground (but not below-ground) consumers respond positively to retrogression. 5. We show that taxonomic richness of plants and above-ground consumers are positively correlated with total ecosystem C storage, suggesting that conserving old-growth forests simultaneously maximizes biodiversity and C sequestration. However, we find little observational or experimental evidence that plant diversity is a major driver of ecosystem C storage on the islands relative to other biotic and abiotic factors. 6. Synthesis. Our study reveals that across contrasting islands differing in exposure to a key extrinsic driver (historical disturbance regime and resulting retrogression), there are coordinated responses of soil fertility, vegetation, consumer communities and ecosystem C sequestration, which all feed back to one another. It also highlights the value of well-replicated natural experiments for tackling questions about above-ground-below-ground linkages over temporal and spatial scales that are otherwise unachievable.

Fire is the primary form of disturbance in temperate and boreal forest ecosystems. However, our knowledge of the biochemical mechanisms by which fire stimulates forest N cycling is incomplete. Charcoal is a major byproduct of forest fires and is ubiquitous in soils of most forest ecosystems, yet the biological function of charcoal in soils of forest ecosystems has been greatly overlooked. We conducted a suite of laboratory experiments on soils from ponderosa pine (Pinus ponderosa Laws) forests to determine the influence of charcoal on soil N dynamics and in particular, nitrification. The addition of NH 4 1 to forest soils had absolutely no effect on nitrification demonstrating that this process is not substrate limited. The amendment of these soils with NH 4 1 and field collected charcoal (1% w/w) significantly increased the nitrification potential, net nitrification, gross nitrification, and decreased the solution concentrations of plant secondary compounds (phenolics). Charcoal had no effect on nitrification in soils (from a grassland site) that had naturally high rates of nitrifier activity. The increase in gross nitrification in forest soils and lack of effect on grassland soils suggests that charcoal may alleviate factors that otherwise inhibit the activity of the nitrifying microbial community in forest soils. These results reveal the biological importance of charcoal and advance our mechanistic understanding of how fire drives nutrient cycling in temperate and boreal ecosystems.

Productivity in boreal ecosystems is primarily limited by available soil nitrogen (N), and there is substantial interest in understanding whether deposition of anthropogenically derived reactive nitrogen (N r ) results in greater N availability to woody vegetation, which could result in greater carbon (C) sequestration. One factor that may limit the acquisition of N r by woody plants is the presence of bryophytes, which are a significant C and N pool, and a location where associative cyanobacterial N-fixation occurs. Using a replicated stand-scale N-addition experiment (five levels: 0, 3, 6, 12, and 50 kg N ha À1 yr À1 ; n 5 6) in the boreal zone of northern Sweden, we tested the hypothesis that sequestration of N r into bryophyte tissues, and downregulation of N-fixation would attenuate N r inputs, and thereby limit anthropogenic N r acquisition by woody plants. Our data showed that N-fixation per unit moss mass and per unit area sharply decreased with increasing N addition. Additionally, the tissue N concentrations of Pleuorzium schreberi increased and its biomass decreased with increasing N addition. This response to increasing N addition caused the P. schreberi N pool to be stable at all but the highest N addition rate, where it significantly decreased. The combined effects of changed N-fixation and P. schreberi biomass N accounted for 56.7% of cumulative N r additions at the lowest N r addition rate, but only a minor fraction for all other treatments. This 'bryophyte effect' can in part explain why soil inorganic N availability and acquisition by woody plants (indicated by their d 15 N signatures) remained unchanged up to N addition rates of 12 kg ha À1 yr À1 or greater. Finally, we demonstrate that approximately 71.8% of the boreal forest experiences N r deposition rates at or below 3 kg ha À1 yr À1 , suggesting that bryophytes likely limit woody plant acquisition of ambient anthropogenic N r inputs throughout a majority of the boreal forest.

Human societies depend on an Earth System that operates within a constrained range of nutrient 68 availability, yet the recent trajectory of terrestrial nitrogen (N) availability is uncertain. 69 Examining patterns of foliar N concentrations ([N]) and isotope ratios (δ 15 N) from more than 42,000 samples acquired over years, here we show that foliar [N] declined by 8% and foliar δ 15 N declined by 0.8 -1.9 ‰. Examining patterns across different climate spaces, foliar δ 15 N declined across the entire range of MAT and MAP tested. These results suggest declines in N supply relative to plant demand at the global scale. In all, there are now multiple lines of evidence of declining N availability in many unfertilized terrestrial ecosystems, including declines in δ 15 N of tree rings and leaves from herbarium samples over the past 75-150 years. 76These patterns are consistent with the proposed consequences of elevated atmospheric CO 2 and longer growing seasons. These declines will limit future terrestrial C uptake and increase nutritional stress for herbivores. 235 much. Preventing these declines in N availability further emphasizes the need to reduce 236 anthropogenic CO 2 emissions.Data and code availability. The datasets generated during and/or analysed during the current study are available in the Dryad repository [link to be generated upon acceptance]. All code used for statistical analyses and figure generation are available on Dryad (XXX).

Ecosystems in the far north, including arctic and boreal biomes, are a globally significant pool of carbon (C). Global change is proposed to influence both C uptake and release in these ecosystems, thereby potentially affecting whether they act as C sources or sinks. Bryophytes (i.e., mosses) serve a variety of key functions in these systems, including their association with nitrogen (N2 )-fixing cyanobacteria, as thermal insulators of the soil, and producers of recalcitrant litter, which have implications for both net primary productivity (NPP) and heterotrophic respiration. While ground-cover bryophytes typically make up a small proportion of the total biomass in northern systems, their combined physical structure and N2 -fixing capabilities facilitate a disproportionally large impact on key processes that control ecosystem C and N cycles. As such, the response of bryophyte-cyanobacteria associations to global change may influence whether and how ecosystem C balances are influenced by global change. Here, we review what is known about their occurrence and N2 -fixing activity, and how bryophyte systems will respond to several key global change factors. We explore the implications these responses may have in determining how global change influences C balances in high northern latitudes.