Factors affecting severity of wildfires in Scottish heathlands and blanket bogs.

Temperate heathlands and blanket bogs are globally rare and face growing wildfire threats. Ecosystem impacts differ between low and high severity fires, where severity reflects immediate fuel consumption. This study assessed factors influencing fire severity in Scottish heathlands and blanket bogs, including the efficacy of the Canadian Fire Weather Index System (CFWIS). Using remote sensing, we measured the differenced Normalised Burn Ratio at 92 wildfire sites from 2015 to 2021. We used Generalised Additive Mixed Models to investigate the impact of topography, habitat wetness, CFWIS components and 30-day weather on severity. Dry heath exhibited higher severity than wet heath and blanket bog, and slope, elevation and south facing aspect were positively correlated to severity. Weather effects were less clear due to data scale differences, yet still indicated weather's significant role in severity. Rainfall had an increasingly negative effect from approximately 15 days before the fire, whilst temperature had an increasingly positive effect. Vapour Pressure Deficit (VPD) was the weather variable with highest explanatory value, and predicted severity better than any CFWIS component. The best-explained fire severity model (R2 = 0.25) incorporated topography, habitat wetness wind and VPD on the day of the fire. The Drought Code (DC), predicting organic matter flammability at ≥10 cm soil depth, was the CFWIS component with the highest predictive effect across habitats. Our findings suggest that wildfires in wet heath and blanket bogs are typically characterised by low severity, but that warmer, drier weather may increase the risk of severe, smouldering fires which threaten peatland carbon stores.

Legacy effects of nitrogen and phosphorus additions on vegetation and carbon stocks of upland heaths.

Soil carbon (C) pools and plant community composition are regulated by nitrogen (N) and phosphorus (P) availability. Atmospheric N deposition impacts ecosystem C storage, but the direction of response varies between systems. Phosphorus limitation may constrain C storage response to N, hence P application to increase plant productivity and thus C sequestration has been suggested. We revisited a 23-yr-old field experiment where N and P had been applied to upland heath, a widespread habitat supporting large soil C stocks. At 10 yr after the last nutrient application we quantified long-term changes in vegetation composition and in soil and vegetation C and P stocks. Nitrogen addition, particularly when combined with P, strongly influenced vegetation composition, favouring grasses over Calluna vulgaris, and led to a reduction in vegetation C stocks. However, soil C stocks did not respond to nutrient treatments. We found 40% of the added P had accumulated in the soil. This study showed persistent effects of N and N + P on vegetation composition, whereas effects of P alone were small and showed recovery. We found no indication that P application could mitigate the effects of N on vegetation or increase C sequestration in this system.

Combination of herbivore removal and nitrogen deposition increases upland carbon storage.

Ecosystem carbon (C) accrual and storage can be enhanced by removing large herbivores as well as by the fertilizing effect of atmospheric nitrogen (N) deposition. These drivers are unlikely to operate independently, yet their combined effect on aboveground and belowground C storage remains largely unexplored. We sampled inside and outside 19 upland grazing exclosures, established for up to 80 years, across an N deposition gradient (5-24 kg N ha(-1) yr(-1) ) and found that herbivore removal increased aboveground plant C stocks, particularly in moss, shrubs and litter. Soil C storage increased with atmospheric N deposition, and this was moderated by the presence or absence of herbivores. In exclosures receiving above 11 kg N ha(-1) year(-1) , herbivore removal resulted in increased soil C stocks. This effect was typically greater for exclosures dominated by dwarf shrubs (Calluna vulgaris) than by grasses (Molinia caerulea). The same pattern was observed for ecosystem C storage. We used our data to predict C storage for a scenario of removing all large herbivores from UK heathlands. Predictions were made considering herbivore removal only (ignoring N deposition) and the combined effects of herbivore removal and current N deposition rates. Predictions including N deposition resulted in a smaller increase in UK heathland C storage than predictions using herbivore removal only. This finding was driven by the fact that the majority of UK heathlands receive low N deposition rates at which herbivore removal has little effect on C storage. Our findings demonstrate the crucial link between herbivory by large mammals and atmospheric N deposition, and this interaction needs to be considered in models of biogeochemical cycling.

Slow recovery of High Arctic heath communities from nitrogen enrichment.

Arctic ecosystems are strongly nutrient limited and exhibit dramatic responses to nitrogen (N) enrichment, the reversibility of which is unknown. This study uniquely assesses the potential for tundra heath to recover from N deposition and the influence of phosphorus (P) availability on recovery. We revisited an experiment in Svalbard, established in 1991, in which N was applied at rates representing atmospheric N deposition in Europe (10 and 50 kg N ha(-1)  yr(-1) ; 'low' and 'high', respectively) for 3-8 yr. We investigated whether significant effects on vegetation composition and ecosystem nutrient status persisted up to 18 yr post-treatment. Although the tundra heath is no longer N saturated, N treatment effects persist and are strongly P-dependent. Vegetation was more resilient to N where no P was added, although shrub cover is still reduced in low-N plots. Where P was also added (5 kg P ha(-1)  yr(-1) ), there are still effects of low N on community composition and nutrient dynamics. High N, with and without P, has many lasting impacts. Importantly, N + P has caused dramatically increased moss abundance, which influences nutrient dynamics. Our key finding is that Arctic ecosystems are slow to recover from even small N inputs, particularly where P is not limiting.

Calluna vulgaris-dominated upland heathland sequesters more CO2 annually than grass-dominated upland heathland.

It has been shown in many habitats worldwide, that a shift in vegetation composition between woody shrub and graminoid dominance can influence carbon (C) cycling. Due to land management practices and environmental change, UK upland heath vegetation has historically undergone shifts in dominance from the woody dwarf shrub Calluna vulgaris (Calluna) to species poor graminoid swards. The consequences of this for C sequestration are unknown. We compared annual net ecosystem exchange (NEE) of carbon dioxide (CO2) between building phase Calluna- and grass-dominated communities within three upland heaths in Scotland, measuring c. monthly over a year. Light and temperature response curves were generated, and the parameters derived were applied to continuous light and temperature data to extrapolate CO2 fluxes over the full year and generate estimates of annual CO2 sequestration for each vegetation type. Grass-dominated communities had higher ecosystem respiration rates than Calluna-dominated communities, attributed to graminoids having greater metabolic demands and producing more labile litter which decomposes readily. Both communities had similar gross primary productivity over the year; the net result being higher NEE within the Calluna-dominated than the grass-dominated community (-2.36 ± 0.23 and -1.78 ± 0.18 µmol CO2m(-2)s(-1) respectively). Modelled CO2 fluxes over a year showed both communities to be CO2 sinks. The Calluna-dominated community sequesters -3.45 ± 0.96 t C ha(-1)yr(-1), double that sequestered by the grass-dominated community at 1.61 ± 0.57 t C ha(-1)yr(-1). Potential rate of C sequestration by upland heath is comparable to that of woodland and the increase in total sequestration that could be gained from habitat restoration may equate to c. 60% of the annual UK C sink attributed to forest land management. National C sequestration by heathlands is also more than double that by peatlands. Management of graminoid-dominated upland heath should promote Calluna re-establishment, thus providing a C benefit in addition to benefits to biodiversity, grazing and sporting interests.

Root traits predict decomposition across a landscape-scale grazing experiment.

Root litter is the dominant soil carbon and nutrient input in many ecosystems, yet few studies have considered how root decomposition is regulated at the landscape scale and how this is mediated by land-use management practices. Large herbivores can potentially influence below-ground decomposition through changes in soil microclimate (temperature and moisture) and changes in plant species composition (root traits). To investigate such herbivore-induced changes, we quantified annual root decomposition of upland grassland species in situ across a landscape-scale livestock grazing experiment, in a common-garden experiment and in laboratory microcosms evaluating the influence of key root traits on decomposition. Livestock grazing increased soil temperatures, but this did not affect root decomposition. Grazing had no effect on soil moisture, but wetter soils retarded root decomposition. Species-specific decomposition rates were similar across all grazing treatments, and species differences were maintained in the common-garden experiment, suggesting an overriding importance of litter type. Supporting this, in microcosms, roots with lower specific root area (m(2) g(-1)) or those with higher phosphorus concentrations decomposed faster. Our results suggest that large herbivores alter below-ground carbon and nitrogen dynamics more through their effects on plant species composition and associated root traits than through effects on the soil microclimate.

Isolation of microsatellite primers for Melampyrum sylvaticum (Orobanchaceae), an endangered plant in the United Kingdom.

PREMISE OF THE STUDY: Microsatellite markers were developed for the hemiparasitic plant Melampyrum sylvaticum to investigate the breeding system, genetic diversity, and structure of populations in the United Kingdom, Sweden, and Norway. METHODS AND RESULTS: Microsatellites were isolated from genomic DNA using an enrichment protocol. Twenty-nine loci were characterized in two individuals from each of 15 geographically disparate populations ("global"). Seven polymorphic loci were further characterized in one population ("local"). The number of alleles per locus ranged from two to 12 in the global sample and one to seven in the local sample. The expected heterozygosity ranged from 0-0.75, the observed heterozygosity from 0-0.1, and the inbreeding coefficient from 0.84-1 in the local sample. CONCLUSIONS: The results show the utility of these novel polymorphic microsatellite markers for further conservation genetic analyses. The strong deficit of heterozygosity across all loci in the local sample suggests the species may be inbreeding.

Balancing positive and negative plant interactions: how mosses structure vascular plant communities.

Our understanding of positive and negative plant interactions is primarily based on vascular plants, as is the prediction that facilitative effects dominate in harsh environments. It remains unclear whether this understanding is also applicable to moss-vascular plant interactions, which are likely to be influential in low-temperature environments with extensive moss ground cover such as boreal forest and arctic tundra. In a field experiment in high-arctic tundra, we investigated positive and negative impacts of the moss layer on vascular plants. Ramets of the shrub Salix polaris, herb Bistorta vivipara, grass Alopecurus borealis and rush Luzula confusa were transplanted into plots manipulated to contain bare soil, shallow moss (3 cm) and deep moss (6 cm) and harvested after three growing seasons. The moss layer had both positive and negative impacts upon vascular plant growth, the relative extent of which varied among vascular plant species. Deep moss cover reduced soil temperature and nitrogen availability, and this was reflected in reduced graminoid productivity. Shrub and herb biomass were greatest in shallow moss, where soil moisture also appeared to be highest. The relative importance of the mechanisms by which moss may influence vascular plants, through effects on soil temperature, moisture and nitrogen availability, was investigated in a phytotron growth experiment. Soil temperature, and not nutrient availability, determined Alopecurus growth, whereas Salix only responded to increased temperature if soil nitrogen was also increased. We propose a conceptual model showing the relative importance of positive and negative influences of the moss mat on vascular plants along a gradient of moss depth and illustrate species-specific outcomes. Our findings suggest that, through their strong influence on the soil environment, mat-forming mosses structure the composition of vascular plant communities. Thus, for plant interaction theory to be widely applicable to extreme environments such as the Arctic, growth forms other than vascular plants should be considered.

Assessing the recovery potential of alpine moss-sedge heath: reciprocal transplants along a nitrogen deposition gradient.

The potential of alpine moss-sedge heath to recover from elevated nitrogen (N) deposition was assessed by transplanting Racomitrium lanuginosum shoots and vegetation turfs between 10 elevated N deposition sites (8.2-32.9 kg ha(-1) yr(-1)) and a low N deposition site, Ben Wyvis (7.2 kg ha(-1) yr(-1)). After two years, tissue N of Racomitrium shoots transplanted from higher N sites to Ben Wyvis only partially equilibrated to reduced N deposition whereas reciprocal transplants almost matched the tissue N of indigenous moss. Unexpectedly, moss shoot growth was stimulated at higher N deposition sites. However, moss depth and biomass increased in turfs transplanted to Ben Wyvis, apparently due to slower shoot turnover (suggested to result partly from decreased tissue C:N slowing decomposition), whilst abundance of vascular species declined. Racomitrium heath has the potential to recover from the impacts of N deposition; however, this is constrained by the persistence of enhanced moss tissue N contents.

Recovery of ecosystem carbon fluxes and storage from herbivory.

The carbon (C) sink strength of arctic tundra is under pressure from increasing populations of arctic breeding geese. In this study we examined how CO2 and CH4 fluxes, plant biomass and soil C responded to the removal of vertebrate herbivores in a high arctic wet moss meadow that has been intensively used by barnacle geese (Branta leucopsis) for ca. 20 years. We used 4 and 9 years old grazing exclosures to investigate the potential for recovery of ecosystem function during the growing season (July 2007). The results show greater above- and below-ground vascular plant biomass within the grazing exclosures with graminoid biomass being most responsive to the removal of herbivory whilst moss biomass remained unchanged. The changes in biomass switched the system from net emission to net uptake of CO2 (0.47 and -0.77 µmol m-2 s-1 in grazed and exclosure plots, respectively) during the growing season and doubled the C storage in live biomass. In contrast, the treatment had no impact on the CH4 fluxes, the total litter C pool or the soil C concentration. The rapid recovery of the above ground biomass and CO2 fluxes demonstrates the plasticity of this high arctic ecosystem in terms of response to changing herbivore pressure.

Herbivore impacts to the moss layer determine tundra ecosystem response to grazing and warming.

Herbivory and climate are key environmental drivers, shaping ecosystems at high latitudes. Here, we focus on how these two drivers act in concert, influencing the high arctic tundra. We aim to investigate mechanisms through which herbivory by geese influences vegetation and soil processes in tundra ecosystems under ambient and warmed conditions. To achieve this, two grazing treatments, clipping plus faecal additions and moss removal, were implemented in conjunction with passive warming. Our key finding was that, in many cases, the tundra ecosystem response was determined by treatment impacts on the moss layer. Moss removal reduced the remaining moss layer depth by 30% and increased peak grass biomass by 27%. These impacts were probably due to observed higher soil temperatures and decomposition rates associated with moss removal. The positive impact of moss removal on grass biomass was even greater with warming, further supporting this conclusion. In contrast, moss removal reduced dwarf shrub biomass possibly resulting from increased exposure to desiccating winds. An intact moss layer buffered the soil to increased air temperature and as a result there was no response of vascular plant productivity to warming over the course of this study. In fact, moss removal impacts on soil temperature were nearly double those of warming, suggesting that the moss layer is a key component in controlling soil conditions. The moss layer also absorbed nutrients from faeces, promoting moss growth. We conclude that both herbivory and warming influence this high arctic ecosystem but that herbivory is the stronger driver of the two. Disturbance to the moss layer resulted in a shift towards a more grass-dominated system with less abundant mosses and shrubs, a trend that was further enhanced by warming. Thus herbivore impacts to the moss layer are key to understanding arctic ecosystem response to grazing and warming.

Habitat type determines herbivory controls over CO2 fluxes in a warmer Arctic.

High-latitude ecosystems store large amounts of carbon (C); however, the C storage of these ecosystems is under threat from both climate warming and increased levels of herbivory. In this study we examined the combined role of herbivores and climate warming as drivers of CO2 fluxes in two typical high-latitude habitats (mesic heath and wet meadow). We hypothesized that both herbivory and climate warming would reduce the C sink strength of Arctic tundra through their combined effects on plant biomass and gross ecosystem photosynthesis and on decomposition rates and the abiotic environment. To test this hypothesis we employed experimental warming (via International Tundra Experiment [ITEX] chambers) and grazing (via captive Barnacle Geese) in a three-year factorial field experiment. Ecosystem CO2 fluxes (net ecosystem exchange of CO2, ecosystem respiration, and gross ecosystem photosynthesis) were measured in all treatments at varying intensity over the three growing seasons to capture the impact of the treatments on a range of temporal scales (diurnal, seasonal, and interannual). Grazing and warming treatments had markedly different effects on CO2 fluxes in the two tundra habitats. Grazing caused a strong reduction in CO2 assimilation in the wet meadow, while warming reduced CO2 efflux from the mesic heath. Treatment effects on net ecosystem exchange largely derived from the modification of gross ecosystem photosynthesis rather than ecosystem respiration. In this study we have demonstrated that on the habitat scale, grazing by geese is a strong driver of net ecosystem exchange of CO2, with the potential to reduce the CO2 sink strength of Arctic ecosystems. Our results highlight that the large reduction in plant biomass due to goose grazing in the Arctic noted in several studies can alter the C balance of wet tundra ecosystems. We conclude that herbivory will modulate direct climate warming responses of Arctic tundra with implications for the ecosystem C balance; however, the magnitude and direction of the response will be habitat-specific.

Short-term exposure to elevated atmospheric CO2 benefits the growth of a facultative annual root hemiparasite, Rhinanthus minor (L.), more than that of its host, Poa pratensis (L.).

The effects of elevated CO2 (650 ppm) on interactions between a chlorophyllous parasitic angiosperm, Rhinanthus minor (L.) and a host, Poa pratensis (L.) were investigated. R. minor benefited from elevated CO2, with both photosynthesis and biomass increasing, and transpiration and tissue N concentration remaining unaffected. However, this did not alleviate the negative effect of the parasite on the host; R. minor reduced host photosynthesis, transpiration, leaf area and biomass, irrespective of CO2 concentration. Elevated CO2 resulted in increased host photosynthesis, but there was no concomitant increase in biomass and foliar N decreased. It appears that the parasite may reduce host growth more by competition for nitrogen than for carbon. Contrary to expectation, R. minor did not reduce the productivity of the host-parasite association, and it actually contributed to the stimulation of productivity of the association by elevated CO2.

Differential responses of UK upland plants to nitrogen deposition.

Native upland species, Nardus stricta, Eriophorum vaginatum, Erica cinerea and Vaccinium vitis-idaea were given 3 or 60 kg N ha-1 yr-1 , over 2 yr, applied as a mist (NH4 NO3 ). The high N treatment increased above-ground biomass in all four species, but only significantly in E. cinerea, E. vaginatum and N. stricta. Biomass increases in E. vaginatum and N. stricta resulted from enhanced tiller production rather than shoot elongation. Root growth increased in N. stricta, so that root∶shoot ratio in this species was unchanged by N. Root growth in E. vaginatum, E. cinerea and V. vitis-idaea did not respond to N and their root∶shoot ratios decreased. Tissue N concentrations increased in both shoots and roots of all species in response to N. The accumulated foliar N did not increase the proportion of N allocated to Rubisco and the photosynthetic capacities of N. stricta, E. vaginatum and V. vitis-idaea were unchanged. Thus growth responses to N were due to altered allocation rather than increased rate of photosynthesis per unit leaf area. The high N treatment increased flower production significantly in E. cinerea but not in the other species. Although in this experiment dwarf shrubs were more responsive than graminoids to N, in the field at current N inputs the enhanced tillering of the graminoids may be more competitively advantageous, especially where gaps develop in the canopy. Thus increasing N deposition may lead to increased grassiness of upland heath, and in particular, a spread of N. stricta.