Improved description of terrestrial habitat types by including microbial communities as indicators.

Soils host diverse communities of microorganisms essential for ecosystem functions and soil health. Despite their importance, microorganisms are not covered by legislation protecting biodiversity or habitats, such as the Habitats Directive. Advances in molecular methods have caused breakthroughs in microbial community analysis, and recent studies have shown that parts of the communities are habitat-specific. If distinct microbial communities are present in the habitat types defined in the Habitats Directive, the Directive may be improved by including these communities. Thus, monitoring and reporting of biodiversity and conservation status of habitat types could be based not only on plant communities but also on microbial communities. In the present study, bacterial and plant communities were examined in six habitat types defined in the Habitats Directive by conducting botanical surveys and collecting soil samples for amplicon sequencing across 19 sites in Denmark. Furthermore, selected physico-chemical properties expected to differ between habitat types and explain variations in community composition of bacteria and vegetation were analysed (pH, electrical conductivity (EC), soil texture, soil water repellency, soil organic carbon content (OC), inorganic nitrogen, and in-situ water content (SWC)). Despite some variations within the same habitat type and overlaps between habitat types, habitat-specific communities were observed for both bacterial and plant communities, but no correlation was observed between the alpha diversity of vegetation and bacteria. PERMANOVA analysis was used to evaluate the variables best able to explain variation in the community composition of vegetation and bacteria. Habitat type alone could explain 46% and 47% of the variation in bacterial and plant communities, respectively. Excluding habitat type as a variable, the best model (pH, SWC, OC, fine silt, and Shannon's diversity index for vegetation) could explain 37% of the variation for bacteria. For vegetation, the best model (pH, EC, ammonium content and Shannon's diversity index for bacteria) could explain 25% of the variation. Based on these results, bacterial communities could be included in the Habitats Directive to improve the monitoring, as microorganisms are more sensitive to changes in the environment compared to vegetation, which the current monitoring is based on.

Resilience of temperate peatland vegetation communities to wildfire depends upon burn severity and pre-fire species composition.

Peatland ecosystems are of global conservation and environmental importance storing globally significant amounts of ancient carbon, regulating regional temperatures and hydrological regimes, and supporting unique biodiversity. Livestock grazing, land-use change, drainage, nutrient and acid deposition, and wildfire threaten the composition and function of many peatlands including those in the uplands of the United Kingdom. Presently, little is known about either the short- or long-term effects of wildfires within these systems in the UK. Our study aimed to evaluate how plant communities respond to wildfires across a range of vegetation communities, soil types, and burn severities. We evaluated wildfire burn severity using the ground-based Composite Burn Index adapted for treeless peatlands. Using paired burned-unburned plots, we quantified differences in the abundance of plant families and functional groups, vegetation diversity, and community composition. Multivariate differences in composition between burned and unburned areas were used as an index of community resilience to fire. Plots in heathland communities with shallow organic soils burned at the highest severities and had the greatest reductions in plant diversity and richness. There were significant declines in plot-scale species richness and diversity with increasing burn severity. Graminoids were resilient to fire whilst Ericaceae tended to increase with higher severity. Bryophyte composition was substantially altered-pleurocarpous species declined and acrocarpous species increased with greater burn severity. Community resilience was related to ground layer burn severity with higher burn severity driving greater changes in communities. Wildfire effects on temperate peatlands are a function of fire weather and site environmental and ecological characteristics. Management policy should ensure that the risk of severe wildfires is mitigated to protect ecosystem function and biodiversity. This will require system-specific fire management prescriptions across the gradient of peatland soil and vegetation types.

Long-term temporal patterns in ecosystem carbon flux components and overall balance in a heathland ecosystem.

Terrestrial ecosystems have strong feedback to atmospheric CO2 concentration and climate change. However, the long-term whole life cycle dynamics of ecosystem carbon (C) fluxes and overall balance in some ecosystem types, such as heathland ecosystems, have not been thoroughly explored. We studied the changes in ecosystem CO2 flux components and overall C balance over a full ecosystem lifecycle in stands of Calluna vulgaris (L.) Hull by using a chronosequence of 0, 12, 19 and 28 years after vegetation cutting. Overall, the ecosystem C balance was highly nonlinear over time and exhibited a sinusoidal-like curvature of C sink/source change over the three-decade timescale. After cutting, plant-related C flux components of gross photosynthesis (PG), aboveground autotrophic respiration (Raa) and belowground autotrophic respiration (Rba) were higher at the young age (12 years) than at middle (19 years) and old (28 years) ages. The young ecosystem was a C sink (12 years: -0.374 kg C m-2 year-1) while it became a C source with aging (19 years: 0.218 kg C m-2 year-1) and when dying (28 years: 0.089 kg C m-2 year-1). The post-cutting C compensation point was observed after four years, while the cumulative C loss in the period after cutting had been compensated by an equal amount of C uptake after seven years. Annual ecosystem C payback from the ecosystem to the atmosphere started after 16 years. This information may be used directly for optimizing vegetation management practices for maximal ecosystem C uptake capacity. Our study highlights that whole life cycle observational data of changes in C fluxes and balance in ecosystems are important and the ecosystem model needs to take the successional stage and vegetation age into account when projecting component C fluxes, ecosystem C balance, and overall feedback to climate change.

Contrasting plant-soil-microbial feedbacks stabilize vegetation types and uncouple topsoil C and N stocks across a subarctic-alpine landscape.

Global vegetation regimes vary in belowground carbon (C) and nitrogen (N) dynamics. However, disentangling large-scale climatic controls from the effects of intrinsic plant-soil-microbial feedbacks on belowground processes is challenging. In local gradients with similar pedo-climatic conditions, effects of plant-microbial feedbacks may be isolated from large-scale drivers. Across a subarctic-alpine mosaic of historic grazing fields and surrounding heath and birch forest, we evaluated whether vegetation-specific plant-microbial feedbacks involved contrasting N cycling characteristics and C and N stocks in the organic topsoil. We sequenced soil fungi, quantified functional genes within the inorganic N cycle, and measured 15 N natural abundance. In grassland soils, large N stocks and low C : N ratios associated with fungal saprotrophs, archaeal ammonia oxidizers, and bacteria capable of respiratory ammonification, indicating maintained inorganic N cycling a century after abandoned reindeer grazing. Toward forest and heath, increasing abundance of mycorrhizal fungi co-occurred with transition to organic N cycling. However, ectomycorrhizal fungal decomposers correlated with small soil N and C stocks in forest, while root-associated ascomycetes associated with small N but large C stocks in heath, uncoupling C and N storage across vegetation types. We propose that contrasting, positive plant-microbial feedbacks stabilize vegetation trajectories, resulting in diverging soil C : N ratios at the landscape scale.

Quantitative assessment of the dispersal of soil-dwelling oribatid mites via rodents in restored heathlands.

Heathland restoration using topsoil removal requires the re-colonization of above- and belowground communities. Oribatid mites play a key role in the comminution of organic matter and are frequently early colonizers during succession despite their limited mobility. Whereas the assembly of their communities may take decades, passive dispersal likely dominates colonization processes, but especially dispersal via other animals (phoresy) remains poorly studied. Compared to other potential hosts, movement habits and ecology of small rodents may provide dispersal advantages to oribatid communities. We studied dispersal of oribatid mites via small rodents in restored heathland sites of different age. We measured movement patterns of small rodents and extracted mites from their pelts and nests to estimate annual contributions of these rodents to the dispersal of oribatids. We also discussed phoretic estimates reported on other host groups as a reference. Probability estimates of oribatids in pelts and nests showed lower occurrence frequencies compared to other reported phoretic hosts. However, local rodent communities may aid the dispersal of up to 41,000 oribatid mites per year. We highlight the high diversity of oribatid species mounting rodents, unlike strong species-specific filters reported in other passive pathways. We found that over half (58%) of the oribatid species reproduced asexually and over a third (32%) had a soil-dwelling lifestyle. We also observed that rodents often travel short distances below 40 m, but occasionally reach distances of up to 100 m, especially in earlier successional stages. Synthesis and applications. Our results suggest that rodents may contribute to assembly processes of soil-dwelling oribatid communities given the slow turnover rate of this group in heathlands. This is accomplished through short-distance dispersal, and especially in sites at early stages of succession. To our knowledge, we are the first to quantitatively assess the potential dispersal of oribatid mites via rodents.

Displacement Effects of Conservation Grazing on Red Deer (Cervus elaphus) Spatial Behaviour.

Conservation grazing uses semi-feral or domesticated herbivores to limit encroachment in open areas and to promote biodiversity. However, we are still unaware of its effects on wild herbivores. This study investigates the influence of herded sheep and goats on red deer (Cervus elaphus) spatial behavior by testing three a-priori hypotheses: (i) red deer are expected to avoid areas used by livestock, as well as adjacent areas, when livestock are present, albeit (ii) red deer increase the use of these areas when sheep and goats are temporarily absent and (iii) there is a time-lagged disruption in red deer spatial behavior when conservation grazing practice ends. Using GPS-telemetry data on red deer from a German heathland area, we modelled their use of areas grazed by sheep and goats, using mixed-effect logistic regression. Additionally, we developed seasonal resource selection functions (use-availability design) to depict habitat selection by red deer before, during, and after conservation grazing. Red deer used areas less during conservation grazing throughout all times of the day and there was no compensatory use during nighttime. This effect mostly persisted within 21 days after conservation grazing. Effects on habitat selection of red deer were detectable up to 3000 meters away from the conservation grazing sites, with no signs of either habituation or adaption. For the first time, we demonstrate that conservation grazing can affect the spatio-temporal behavior of wild herbivores. Our findings are relevant for optimizing landscape and wildlife management when conservation grazing is used in areas where wild herbivores are present.

Habitat Protection Approaches Facilitate Conservation of Overlooked Fungal Diversity - A Case Study From the Norwegian Coastal Heathland System.

European coastal heathlands are distinct ecosystems shaped by land use tradition and they have experienced an 80% area reduction from their historical maximum. These mosaics of mires and wind exposed patches have ericaceous shrub dominated vegetation, and soils within coastal heathlands are characterized by low pH and high levels of recalcitrant debris. Using a culture-based approach with molecular identification of isolates, we characterized root-associated fungal communities of six ericaceous species in eight heathland localities along Norway's western coast. Site-level alpha diversity ranged from 21-38 OTUs, while the total estimated gamma diversity for culturable heathland root fungi was 190-231 OTUs. Most species recovered are previously reported at low abundance in Norway, suggesting the biodiversity in this community is underreported, rather than novel for science. The fungi recovered were primarily Ascomycota, specifically endophytic Phialocephala, and Pezicula, and no host specificity was observed in the communities. The fungal communities exhibited high turnover and low nestedness, both between ericaceous hosts and across heathland sites. We observed no spatial patterns in fungal betadiversity, and this heterogeneity may be a product of the unique historic land use practices at each locality creating a distinct mycofloral "fingerprint". Robust diversity estimates will be key for managing fungal biodiversity in coastal heathlands. Our results indicate that sampling schemes that maximize the number of host plants sampled per site, rather than the number of cultures per plant yield improved alpha diversity estimates. Similarly, gamma diversity estimates are improved by maximizing the total number of localities sampled, rather than increasing the number of plants sampled per locality. We argue that while the current protected status of coastal heathland habitats and restoration efforts have knock-on effects for the conservation of fungal biodiversity, fungi have a vital functional role in the ecosystem and holistic conservation plans that consider fungal biodiversity would be beneficial.

Mapping the ecological resilience of Atlantic postglacial heathlands.

Anthropogenic heathlands are semi-natural ecosystems with a unique cultural and biodiversity value, considered worthy of preservation across most of the world. Their rate of loss, however, is alarming. Currently, we know little about the heathlands' actual span of resilience affordances and their association with abiotic and anthropogenic factors, including how much additional intervention they need to persist. Consequently, we are missing out on vital knowledge for conservation, management and the historical persistence of heathlands.This paper develops a method to assess the ecological resilience affordances of Atlantic postglacial heaths in the absence of human management. We use 12 existing cases of heathland succession to establish a four-step resilience grade for each site, which we regress onto a series of explaining factors and use it in predicting heath resilience across postglacial Atlantic Northern Europe.We find that temperature, humidity, elevation and sandiness have a positive correlation with high heathland resilience. Our predictive mapping shows an uneven distribution of ecological heath resilience across Atlantic Northern Europe within an area of 1,000 × 1,200 km of 5 × 5 km resolution.Historic heathland distributions far exceed areas that afford high heath resilience, suggesting that heath distribution and persistence depend on both abiotic and anthropogenic factors. Policy implications: The map predicting the ecological resilience of Atlantic postglacial heaths can be used by managers working towards heath preservation and restoration to prioritize conservation efforts and to plan management practices across Atlantic Northern Europe. Together with the predictive model, it provides an important initial screening tool to assess heathland resilience in the absence of management as well as the impact of atmospheric nitrogen. The results are equally relevant for scholars who are interested in humans' role in increasing and decreasing ecosystem resilience. Our predictive method can be applied in other regions across the world by adding regionally specific variables.

Prevalence and phenology of fine root endophyte colonization across populations of Lycopodiella inundata.

Mycorrhizal fungi are critical components of terrestrial habitats and agroecosystems. Recently, Mucoromycotina fine root endophyte fungi (MucFRE) were found to engage in nutritional mutualism with Lycopodiella inundata, which belongs to one of the earliest vascular plant lineages known to associate with MucFRE. The extent to which this mutualism plays a role in resilient plant populations can only be understood by examining its occurrence rate and phenological patterns. To test for prevalence and seasonality in colonization, we examined 1305 individual L. inundata roots from 275 plants collected during spring and autumn 2019 across 11 semi-natural heathlands in Britain and the Netherlands. We quantified presence/absence of fine root endophyte (FRE) hyphae and vesicles and explored possible relationships between temperature and precipitation in the months immediately before sampling. Fine root endophyte hyphae were dominant in all of the examined heathlands, and every colonized root had FRE in both cortical cells and root hairs. However, we found significant differences in colonization between the two seasons at every site. Overall, 14% of L. inundata roots were colonized in spring (2.4% with vesicles) compared with 86% in autumn (7.6% with vesicles). Colonization levels between populations were also significantly different, correlating with temperature and precipitation, suggesting some local environments may be more conducive to root and related hyphal growth. These marked seasonal differences in host-plant colonization suggest that results about FRE from single time point collections should be carefully interpreted. Our findings are relevant to habitat restoration, species conservation plans, agricultural bio-inoculation treatments, and microbial diversity studies.

Biomass, Morphology, and Dynamics of the Fine Root System Across a 3,000-M Elevation Gradient on Mt. Kilimanjaro.

Fine roots (≤2 mm) consume a large proportion of photosynthates and thus play a key role in the global carbon cycle, but our knowledge about fine root biomass, production, and turnover across environmental gradients is insufficient, especially in tropical ecosystems. Root system studies along elevation transects can produce valuable insights into root trait-environment relationships and may help to explore the evidence for a root economics spectrum (RES) that should represent a trait syndrome with a trade-off between resource acquisitive and conservative root traits. We studied fine root biomass, necromass, production, and mean fine root lifespan (the inverse of fine root turnover) of woody plants in six natural tropical ecosystems (savanna, four tropical mountain forest types, tropical alpine heathland) on the southern slope of Mt. Kilimanjaro (Tanzania) between 900 and 4,500 m a.s.l. Fine root biomass and necromass showed a unimodal pattern along the slope with a peak in the moist upper montane forest (~2,800 m), while fine root production varied little between savanna and upper montane forest to decrease toward the alpine zone. Root:shoot ratio (fine root biomass and production related to aboveground biomass) in the tropical montane forest increased exponentially with elevation, while it decreased with precipitation and soil nitrogen availability (decreasing soil C:N ratio). Mean fine root lifespan was lowest in the ecosystems with pronounced resource limitation (savanna at low elevation, alpine heathland at high elevation) and higher in the moist and cool forest belt (~1,800-3,700 m). The variation in root traits across the elevation gradient fits better with the concept of a multi-dimensional RES, as root tissue density and specific root length showed variable relations to each other, which does not agree with a simple trade-off between acquisitive and conservative root traits. In conclusion, despite large variation in fine root biomass, production, and morphology among the different plant species and ecosystems, a general belowground shift in carbohydrate partitioning is evident from 900 to 4,500 m a.s.l., suggesting that plant growth is increasingly limited by nutrient (probably N) shortage toward higher elevations.

Initial soil community drives heathland fungal community trajectory over multiple years through altered plant-soil interactions.

Dispersal limitation, biotic interactions, and environmental filters interact to drive plant and fungal community assembly, but their combined effects are rarely investigated. This study examines how different heathland plant and fungal colonization scenarios realized via three biotic treatments - addition of mature heathland-derived sod, addition of hay, and no additions - affect soil fungal community development over 6 yr along a manipulated pH gradient in a large-scale experiment starting from an agricultural, topsoil removed state. Our results show that both biotic and abiotic (pH) treatments had a persistent influence on the development of fungal communities, but that sod additions diminished the effect of abiotic treatments through time. Analysis of correlation networks between soil fungi and plants suggests that the reduced effect of pH in the sod treatment, where both soil and plant propagules were added, might be due to plant-fungal interactions since the sod additions caused stronger, more specific, and more consistent connections compared with the no addition treatment. Based on these results, we suggest that the initial availability of heathland fungal and plant taxa, which reinforce each other, can significantly steer further fungal community development to an alternative configuration, overriding the otherwise prominent effect of abiotic (pH) conditions.

New stiletto flies in the genus Anabarhynchus Macquart (Diptera: Therevidae) from Kangaroo Island, South Australia.

Anabarhynchus Macquart 1848 is a species-rich genus of stiletto flies (Diptera: Therevidae) belonging to the subfamily Therevinae, with over 113 species described from Australia. These flies are diverse and abundant in Australia's eucalypt woodland and mallee habitats. Here we describe, diagnose and illustrate a further three new Anabarhynchus species in the genus as follows: A. aurantilateralis sp. nov. and A. halmaturinus sp. nov., and A. venabrunneis sp. nov., from Kangaroo Island. These new species bring the total number of described Australian species in the genus to 116, with 13 of these known to occur on Kangaroo Island.

Pathways of change: Predicting the effects of fire on flammability.

Impacts of wildfire on humans are increasing as urban populations continue to expand into fire prone landscapes. Effective fire risk management can only be achieved if we understand and quantify how ecosystems change in response to fire and how these changes affect flammability. However, there have been limited studies to this effect with the dominant paradigm being the assumption that recently burnt vegetation is less flammable than older vegetation. To better quantify changes in flammability, we first need to quantify trajectories of changes in response to fire within individual vegetation communities. Second, we need to examine the extent to which these changes alter flammability. Here, we quantify the flammability pathways with increasing time since fire for five vegetation communities in south-eastern Australia. A total of 116 sites were measured across a range of heathland, woodland and forest ecosystems. Flammability was measured using an ecological point based mechanistic fire behaviour model that estimates three measures of flammability relevant to both fire management and research. Predicted changes in flammability varied between vegetation types with heathland and wet forests generally increasing in flammability with time since fire and tall mixed, foothills and forby forests decreasing or showing limited changes with time since fire. Variations in flammability pathways suggest fire management activities that alter fuel structure, such as prescribed burning, may only reduce flammability in a limited set of ecosystems. Incorporating these results into a landscape analysis will improve the quantification of fire risk.

Links Between Heathland Fungal Biomass Mineralization, Melanization, and Hydrophobicity.

Comprehending the decomposition process is crucial for our understanding of the mechanisms of carbon (C) sequestration in soils. The decomposition of plant biomass has been extensively studied. It revealed that extrinsic biomass properties that restrict its access to decomposers influence decomposition more than intrinsic ones that are only related to its chemical structure. Fungal biomass has been much less investigated, even though it contributes to a large extent to soil organic matter, and is characterized by specific biochemical properties. In this study, we investigated the extent to which decomposition of heathland fungal biomass was affected by its hydrophobicity (extrinsic property) and melanin content (intrinsic property). We hypothesized that, as for plant biomass, hydrophobicity would have a greater impact on decomposition than melanin content. Mineralization was determined as the mineralization of soil organic carbon (SOC) into CO2 by headspace GC/MS after inoculation by a heathland soil microbial community. Results show that decomposition was not affected by hydrophobicity, but was negatively correlated with melanin content. We argue that it may indicate that either melanin content is both an intrinsic and extrinsic property, or that some soil decomposers evolved the ability to use surfactants to access to hydrophobic biomass. In the latter case, biomass hydrophobicity should not be considered as a crucial extrinsic factor. We also explored the ecology of decomposition, melanin content, and hydrophobicity, among heathland soil fungal guilds. Ascomycete black yeasts had the highest melanin content, and hyaline Basidiomycete yeasts the lowest. Hydrophobicity was an all-or-nothing trait, with most isolates being hydrophobic.

Combining remote sensing and on-site monitoring methods to investigate footpath erosion within a popular recreational heathland environment.

Footpaths are a prominent consequence of natural area tourism and reflect damage caused to valuable, sensitive habitats by people pressure. Degradation impacts on vegetation, wildlife, on and off-site soil movement and loss, creation of additional informal off-path footpaths (desire lines), and visual destruction of landscapes. Impacts need to be measured and monitored on a large temporal and spatial scale to aid in land management to maintain access and preserve natural environments. This study combined remote sensing (Light Detection and Ranging [LiDAR] and aerial photography) with on-site measurement of footpaths within a sensitive heathland habitat (Land's End, Cornwall, UK). Soil loss, slope angle change, vegetation damage and a hydrology model were combined to comprehensively study the site. Results showed 0.09 m mean soil loss over five years, footpath widening, increasing grass cover into heathland, and water channelling on the footpaths exacerbating erosion. The environments surrounding the footpaths were affected with visitors walking off path, requiring further management and monitoring. Multiple remote sensing techniques were highly successful in comprehensively assessing the area, particularly the hydrology model, demonstrating the potential of providing a valuable objective and quantitative monitoring and management tool.

New species from Phytophthora Clade 6a: evidence for recent radiation.

During routine vegetation health surveys in the southwest of Western Australia (SWWA), several Phytophthora isolates with affinity to Clade 6a have been recovered. In this study, all known taxa from Clade 6a, P. inundata, P. humicola, P. gemini, P. 'walnut' and P. 'personii', and the new isolates were compared based on morphology and DNA sequence data from three nuclear genes and two mitochondrial genes resulting in the description of five new species, P. balyanboodja, P. condilina, P. cooljarloo, P. kwongonina and P. pseudorosacearum. With the exception of P. gemini and P. humicola, all species from Clade 6a have been recovered from natural ecosystems in SWWA. These species are morphologically similar, with predominantly ovoid sporangia and nested and extended internal proliferation. If oospores are present, they tend to be aplerotic with paragynous antheridia mostly attached adjacent to the oogonial stalk. They can all grow at 35 °C and have a fast growth rate on most agar media. These species have all been recovered from the rhizosphere soil and dead and dying plants within dry kwongon heathlands, often from water gaining sites and frequently from very isolated areas. The radiation, origin and potential ecological role of these species are discussed.

Changing contributions of stochastic and deterministic processes in community assembly over a successional gradient.

Successional dynamics in plant community assembly may result from both deterministic and stochastic ecological processes. The relative importance of different ecological processes is expected to vary over the successional sequence, between different plant functional groups, and with the disturbance levels and land-use management regimes of the successional systems. We evaluate the relative importance of stochastic and deterministic processes in bryophyte and vascular plant community assembly after fire in grazed and ungrazed anthropogenic coastal heathlands in Northern Europe. A replicated series of post-fire successions (n = 12) were initiated under grazed and ungrazed conditions, and vegetation data were recorded in permanent plots over 13 years. We used redundancy analysis (RDA) to test for deterministic successional patterns in species composition repeated across the replicate successional series and analyses of co-occurrence to evaluate to what extent species respond synchronously along the successional gradient. Change in species co-occurrences over succession indicates stochastic successional dynamics at the species level (i.e., species equivalence), whereas constancy in co-occurrence indicates deterministic dynamics (successional niche differentiation). The RDA shows high and deterministic vascular plant community compositional change, especially early in succession. Co-occurrence analyses indicate stochastic species-level dynamics the first two years, which then give way to more deterministic replacements. Grazed and ungrazed successions are similar, but the early stage stochasticity is higher in ungrazed areas. Bryophyte communities in ungrazed successions resemble vascular plant communities. In contrast, bryophytes in grazed successions showed consistently high stochasticity and low determinism in both community composition and species co-occurrence. In conclusion, stochastic and individualistic species responses early in succession give way to more niche-driven dynamics in later successional stages. Grazing reduces predictability in both successional trends and species-level dynamics, especially in plant functional groups that are not well adapted to disturbance.

Time- and age-related effects of experimentally simulated nitrogen deposition on the functioning of montane heathland ecosystems.

Ecosystems adapted to low nitrogen (N) conditions such as Calluna-heathlands are especially sensitive to enhanced atmospheric N deposition that affects many aspects of ecosystem functioning like nutrient cycling, soil properties and plant-microbial-enzyme relationships. We investigated the effects of five levels of experimentally-simulated N deposition rates (i.e., N fertilization treatments: 0, 10, 20 and 50kgNha-1yr-1 for 3years, and 56kgNha-1yr-1 for 10years) on: plant, litter, microbial biomass and soil nutrient contents, soil extracellular enzymatic activities, and plant root ericoid mycorrhizal colonization. The study was conducted in marginal montane Calluna-heathlands at different developmental stages resulting from management (young/building-phase and mature-phase). Our findings revealed that many soil properties did not show a statistically significant response to the experimental addition of N, including: total N, organic carbon (C), C:N ratio, extractable N-NO3-, available phosphorus (P), urease and ß-glucosidase enzyme activities, and microbial biomass C and N. Our results also evidenced a considerable positive impact of chronic (10-year) high-N loading on soil extractable N-NH4+, acid phosphatase enzyme activity, Calluna root mycorrhizal colonization by ericoid fungi, Calluna shoot N and P contents, and litter N content and N:P ratio. The age of heathland vegetation influenced the effects of N addition on ericoid mycorrhizal colonization, resulting in higher colonized roots in young heathlands at the control, low and medium N-input rates; and in mature ones at the high and chronically high N rates. Also, young heathlands exhibited greater soil extractable N-NO3-, available P, microbial biomass N, Calluna shoot N and P contents, and litter N content, compared to mature ones. Our results highlighted that accounting for the N-input load and duration, as well as the developmental stage of the vegetation, is important for assessing the effects of added N, particularly at the heathlands' southern distribution limit.

Diet selection and performance of horses grazing on different heathland types.

The number of horses in northern Spanish mountains has increased in recent decades, but little is known about their grazing behaviour, performance and potential for foal meat production. This research aimed to study the diet selection, liveweight (LW) changes and parasitic status of dry and lactating mares, and foals' LW gains, grazing on heathlands with different botanical composition. The experimental design consisted of three vegetation types: dominated by heather (Ericaceae) species (H), dominated by gorse (Ulex gallii; G) and co-dominated by gorse and heath-grasses (G-G), with four replicates per treatment (12 paddocks of 1.2 ha). The study lasted three grazing seasons (2010-12). Each year, 24 crossbred mature mares (310±52 kg LW) were used, managing one lactating mare with her foal plus one non-lactating mare per paddock from May to late summer or early autumn. In the case of H paddocks, animals had to be removed before (late August to early September) because of apparent loss of body condition. Animals were periodically weighed. Mares' diet composition was estimated using alkane markers, analysing the discrepancies in alkane concentrations between dietary plant components and faeces. Faecal samples were also analysed for gastrointestinal nematodes ova. Chemical composition of the main plant components (i.e. heather, gorse and grasses) revealed a low nutritive value, averaging 79, 115 and 113 g CP/kg dry matter (DM), respectively, that could restrict livestock performance. Mares initially selected gorse and grasses (0.47 and 0.40, respectively, in 2010), increasing heather consumption over time (from 0.13 in 2010 to 0.29 in 2012) as gorse availability decreased. The performance of both mares and foals was lower in H compared with G and G-G paddocks (-216 v. 347 g/day for mares, P<0.01; 278 v. 576 g/day for foals, P<0.05), whereas LW changes were more favourable in dry mares than in lactating ones (241 v. 78 g/day; P<0.05). Small strongyle (Cyathostominae) egg counts in mares' faeces increased across the grazing season with no differences between treatments. These results indicate that grazing by horses on gorse- and grass-gorse-dominated shrublands could be sustainable at least during part of the year (4 to 6 months). However, heather-dominated heathlands are not able to meet the nutritional needs of horses even for a short time (2 to 4 months). Nevertheless, the low nutritive quality of these vegetation communities, especially in autumn, requires animal access to other pastures with a higher nutritive value, or supplementary feeding, to enhance foals' growth and maintain sustainable grazing systems with productive herds.

Bryophytes and Organic layers Control Uptake of Airborne Nitrogen in Low-N Environments.

The effects of atmospheric nitrogen (N) deposition on ecosystem functioning largely depend on the retention of N in different ecosystem compartments, but accumulation and partitioning processes have rarely been quantified in long-term field experiments. In the present study we analysed for the first time decadal-scale flows and allocation patterns of N in a heathland ecosystem that has been subject to airborne N inputs over decades. Using a long-term 15N tracer experiment, we quantified N retention and flows to and between ecosystem compartments (above-ground/below-ground vascular biomass, moss layer, soil horizons, leachate). After 9 years, about 60% of the added 15N-tracer remained in the N cycle of the ecosystem. The moss layer proved to be a crucial link between incoming N and its allocation to different ecosystem compartments (in terms of a short-term capture, but long-term release function). However, about 50% of the 15N captured and released by the moss layer was not compensated for by a corresponding increase in recovery rates in any other compartment, probably due to denitrification losses from the moss layer in the case of water saturation after rain events. The O-horizon proved to be the most important long-term sink for added 15N, as reflected by an increase in recovery rates from 18 to 40% within 8 years. Less than 2.1% of 15N were recovered in the podzol-B-horizon, suggesting that only negligible amounts of N were withdrawn from the N cycle of the ecosystem. Moreover, 15N recovery was low in the dwarf shrub above-ground biomass (<3.9% after 9 years) and in the leachate (about 0.03% within 1 year), indicating still conservative N cycles of the ecosystem, even after decades of N inputs beyond critical load thresholds. The continuous accumulation of reactive forms of airborne N suggests that critical load-estimates need to account for cumulative effects of N additions into ecosystems.