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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.

The response of soil microbial communities to a changing climate will impact global biogeochemical cycles, potentially leading to positive and negative feedbacks. However, our understanding of how soil microbial communities respond to climate change and the implications of these changes for future soil function is limited. Here, we assess the response of soil bacterial and fungal communities to long-term experimental climate change in a heathland organo-mineral soil. We analysed microbial communities using Illumina sequencing of the 16S rRNA gene and ITS2 region at two depths, from plots undergoing 4 and 18 years of in situ summer drought or warming. We also assessed the colonisation of Calluna vulgaris roots by ericoid and dark septate endophytic (DSE) fungi using microscopy after 16 years of climate treatment. We found significant changes in both the bacterial and fungal communities in response to drought and warming, likely mediated by changes in soil pH and electrical conductivity. Changes in the microbial communities were more pronounced after a longer period of climate manipulation. Additionally, the subsoil communities of the long-term warmed plots became similar to the topsoil. Ericoid mycorrhizal colonisation decreased with depth while DSEs increased; however, these trends with depth were removed by warming. We largely ascribe the observed changes in microbial communities to shifts in plant cover and subsequent feedback on soil physicochemical properties, especially pH. Our results demonstrate the importance of considering changes in soil microbial responses to climate change across different soil depths and after extended periods of time.

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.

Conservation management is increasingly being required to support both the provision of ecosystem services and maintenance of biodiversity. However, trade‐offs can occur between biodiversity and ecosystems services. We examine whether such trade‐offs can be resolved through landscape‐scale approaches to management. We analysed the biodiversity value and provision of selected ecosystem services (carbon storage, recreation, aesthetic and timber value) on patches of lowland heathland in the southern English county of Dorset. We used transition matrices of vegetation dynamics across 112 heathland patches to forecast biodiversity and ecosystem service provision on patches of different sizes over a 27‐year timeline. Management scenarios simulated the removal of scrub and woodland and compared (i) no management (NM); (ii) all heaths managed equally (AM); and management focused on (iii) small heaths (SM) and (iv) large heaths (LM). Results highlighted a number of trade‐offs. Whereas biodiversity values were significantly lower in woodland than in dry and humid heath, timber, carbon storage and aesthetic values were highest in woodland. While recreation value was positively related to dry heath area, it was negatively related to woodland area. Multicriteria analysis ranked NM highest for aesthetic value, carbon storage and timber value. In contrast, SM ranked highest for recreation and LM highest for biodiversity value. In no scenario did the current site‐based approach to management (AM) rank highest. Synthesis and applications. Biodiversity–ecosystem service trade‐offs are reported in lowland heathland, an ecosystem type of high conservation value. Trade‐offs can be addressed through a landscape‐scale approach to management, by varying interventions according to heathland patch size. Specifically, if management for biodiversity conservation is focused on larger patches, the aesthetic, carbon storage and timber value of smaller patches would increase, as a result of woody succession. In this way, individual heathland patches of either relatively high biodiversity value or high value for provision of ecosystem services could both potentially be delivered at the landscape scale.

•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 six years 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 to no addition treatment. •Based on these results, we suggest that the initial availability of heathland fungal and plant taxa, that reinforce each other, can significantly steer further fungal community development to an alternative configuration, overriding otherwise prominent effect of abiotic (pH) conditions.

Nitrate is a limiting resource in heathland acid soils. Nitrate levels increase in heathland soils after Pteridium aquilinum invasions, and this species is assumed to biologically control nitrogen cycle processes, thus increasing nitrate availability. We compared how P. aquilinum (bracken) and Erica cinerea (bell heather) modify processes driving nitrate availability along a soil pH gradient in a Natura 2000 reserve facing bracken invasion. Soil nitrate and ammonium concentrations, substrate‐induced respiration (SIR), denitrification and nitrification enzyme activities (DEA and NEA, respectively), root procyanidin concentrations, and denitrification inhibition by procyanidins were measured on five sites under P. aquilinum and E. cinerea stands. NEA and nitrate levels were higher, and ammonium levels and SIR lower, for P. aquilinum in the most acid soils. Procyanidins from both species induced the same level of denitrification inhibition, soil nitrate being correlated with root procyanidin concentration for both species. Soil nitrate correlated with NEA only for P. aquilinum. Our results show that both species increased procyanidin production in the most acid soils, thereby reducing denitrification and decreasing nitrate loss, this process being more efficient for E. cinerea. However, P. aquilinum additionally increased nitrification, and this double control on nitrification and denitrification was very efficient to increase soil nitrate availability in the most acid soils. This may participate to the success of P. aquilinum invasions in heathlands. This shows that approaches for bracken control in heathlands should better account for belowground processes and, more generally, that biological denitrification inhibition by plants may be a widespread phenomenon influencing soil N dynamics in N‐poor environments.

Heather, Calluna vulgaris , is a key species of European dry heath and central determinant of its conservation status. The established Calluna life cycle concept describes four phases—pioneer, building, mature, and degeneration—distinguishable by growth and vitality characteristics of undisturbed plants grown from seeds. However, little is known about the life cycle and ageing of plants subjected to severe disturbance, although measures to this effect (burning, mowing) are common in heathland management. We studied the vitality of over 400 heather plants by examining multiple morphological (plant height, long shoot and inflorescence lengths, flowering activity), anatomical (growth rings) and environmental (management, nitrogen deposition, climate) attributes. We found Calluna vitality to be mainly determined by the aboveground stem age, and that severe disturbances promote vigorous vegetative regeneration. Ageing-related shifts in the habit and vitality of plants resprouting from stem-base buds is similar to that of seed-based plants, but the former revealed higher vitality when young, at the cost of a shorter life span. In contrast, plants originating from decumbent stems resemble building-stage plants but apparently lack the capacity to re-enter a cycle including stages other than degeneration-type. As a consequence, we supplemented the established heather life cycle concept with a post-disturbance regeneration cycle of plants derived from resprouting. We conclude that management of dry lowland heathlands should include rotational small-scale severe disturbance to support both seed germination and seedling establishment as well as vegetative regeneration chiefly of young heather plants capable of resprouting from buds near rootstock.

Temperature and nutrients are major limiting factors in subarctic tundra. Experimental manipulation of nutrient availability along elevational gradients (and thus temperature) can improve our understanding of ecological responses to climate change. However, no study to date has explored impacts of nutrient addition along a tundra elevational gradient, or across contrasting vegetation types along any elevational gradient. We set up a full factorial nitrogen (N) and phosphorus (P) fertilization experiment in each of two vegetation types (heath and meadow) at 500 m, 800 m, and 1000 m elevation in northern Swedish tundra. We predicted that plant and microbial communities in heath or at lower elevations would be more responsive to N addition while communities in meadow or at higher elevations would be more responsive to P addition, and that fertilizer effects would vary more with elevation for the heath than for the meadow. Although our results provided little support for these predictions, the relationship between nutrient limitation and elevation differed between vegetation types. Most plant and microbial properties were responsive to N and/or P fertilization, but responses often varied with elevation and/or vegetation type. For instance, vegetation density significantly increased with N + P fertilization relative to the other fertilizer treatments, and this increase was greatest at the lowest elevation for the heath but at the highest elevation for the meadow. Arbuscular mycorrhizae decreased with P fertilization at 500 m for the meadow, but with all fertilizer treatments in both vegetation types at 800 m. Fungal to bacterial ratios were enhanced by N + P fertilization for the two highest elevations in the meadow only. Additionally, microbial responses to fertilization were primarily direct rather than indirect via plant responses, pointing to a decoupled response of plant and microbial communities to nutrient addition and elevation. Because our study shows how two community types differ in their responses to fertilization and elevation, and because the temperature range across this gradient is ∼3°C, our study is informative about how nutrient limitation in tundra may be influenced by temperature shifts that are comparable to those expected under climate change during this century.

The extinction debt, delayed species extinctions following landscape degradation, is a widely discussed concept. But a consensus about the prevalence of extinctions debts is hindered by a multiplicity of methods and a lack of comparisons among habitats. We applied three contrasting species–area relationship methods to test for plant community extinction debts in three habitats which had different degradation histories over the last century: calcareous grassland, heathland and woodland. These methods differ in their data requirements, with the first two using information on past and current habitat area alongside current species richness, whilst the last method also requires data on past species richness. The most data‐intensive, and hence arguably most reliable method, identified extinction debts across all habitats for specialist species, whilst the other methods did not. All methods detected an extinction debt in calcareous grassland, which had undergone the most severe degradation. We conclude that some methods failed to detect an extinction debt, particularly in habitats that have undergone moderate degradation. Data on past species numbers are required for the most reliable method; as such data are rare, extinction debts may be under‐reported.

1. In England, the loss of lowland heathland, a habitat of global conservation importance, is primarily due to the invasion of birch and pine. This encroachment has been researched in depth from a plant perspective but little is known about the role of mycorrhizal fungi. In lowland heathlands the resident dwarf shrubs form ericoid mycorrhizas whereas invading trees form ectomycorrhizas. Therefore, tree encroachment into heathlands can be regarded as the replacement of a resident mycorrhizal community by an invading one. 2. This study examined how fungi form mycorrhizas with Betula and Pinus in lowland heathlands. We addressed the question of whether there are mycorrhizal fungi that mediate invasion using a molecular ecology approach to compare the mycorrhizal inoculum potential of soil at three levels of invasion (uninvaded heathland, invaded heathland and woodland) and the fungi forming mycorrhizas on tree seedlings and trees across diverse sites. 3. We show that in lowland heathlands: (i) seedlings have severely limited access to ectomycorrhizal fungi relative to woodlands, (ii) there are few keystone spore-dispersed ectomycorrhizal fungi that can mediate tree invasion, (iii) tree seedlings can remain non-mycorrhizal for at least one year when no inoculum is present, even near saplings, and (iv) mycorrhizal seedlings achieve greater biomass than non-mycorrhizal seedlings. Within uninvaded heathland we detected only Rhizopogon luteolus, Suillus variegatus, S. bovinus (Pinus symbionts) and Laccaria proxima (primarily a Betula symbiont). 4.Synthesis. Overall, ectomycorrhizal inoculum in lowland heathlands is rare; most tree seedlings growing in heathland soil are not mycorrhizal due to limited spore dispersal, poorly developed spore banks and weak common mycorrhizal networks. These seedlings can persist awaiting mycorrhization to boost their growth.