The reproductive potential and importance of key management aspects for successful Calluna vulgaris rejuvenation on abandoned Continental heaths

The abandonment of traditional pastoralism as well as the use of heath areas for military purposes has had a major impact on dry heaths in the Continental biogeographical region of Europe, causing severe degradation of its key species Calluna vulgaris (L.) HULL. The reproductive potential of this species in a Continental climate is assumed to be low, although there is yet no observational or experimental evidence for this. More knowledge is also needed about cost‐effective and sustainable measures to restore abandoned dry heaths in this biogeographical region, because traditional management options are often too expensive (e.g., sod‐cutting) or restricted due to environmental laws and the danger of unexploded ammunition (e.g., burning). Using as an example an 800 ha Continental heathland in Germany that has been abandoned for about two decades, we studied the reproductive potential (seed production, soil seed bank, and germination ability) of degenerate C. vulgaris stands. In addition, we conducted a comprehensive field experiment to test the effects of low‐intensity, year‐round grazing by Heck cattle and Konik horses as well as one‐time mowing and patchy exposure of bare soil on the generative rejuvenation (i.e., recruitment and survival) of degenerate C. vulgaris stands over 3 years. We used generalized linear mixed models for statistical analyses. Seed production of degenerate C. vulgaris stands was high as well as the germination ability of their seeds, being similar to Atlantic heathlands. However, soil seed‐bank densities were lower than those found in managed or abandoned Atlantic heaths. Overall seedling recruitment in the field was considerably lower in comparison with Atlantic heaths. Low‐intensity grazing or one‐time mowing did not induce a substantial increase in C. vulgaris recruitment, whereas an additional one‐time creation of bare soil patches or the one‐time creation of bare soil without subsequent management significantly facilitated seedling recruitment and survival in the first year. However, from the second year on, the positive effect of the creation of bare soil without subsequent management was no longer present. In the third year, survival of juveniles was significantly supported by low‐intensity grazing in combination with shallow soil disturbances as well as in combination with one‐time mowing and shallow soil disturbances, whereas mowing alone resulted in marginally significant lower survival. The extremely low seedling recruitment requires a careful choice of suitable management measures to promote the survival of sufficient numbers of Calluna individuals. Therefore, we recommend low‐intensity grazing with free‐ranging robust breeds and the combination of this with one‐time mowing as an effective means of supporting generative rejuvenation of C. vulgaris in degraded heaths. However, at the beginning of the restoration process, the creation of bare soil patches for seedling recruitment is crucial. For implementation into practice, we present different strategies to enhance the proportion of bare soil after long‐term abandonment of heaths when traditional management options are no longer feasible.     

Seedling recruitment of Succisella inflexa in fen meadows: Importance of seed and microsite availability

Abstract. The effects of different forms of land use on germination and establishment of the rare fen species Succisella inflexa were investigated in seed introduction experiments in a mown and an abandoned fen meadow in SE Germany. Treatments included abandonment, mowing in fall and mowing with creation of gaps in the moss and litter layer. Floating capacity of seeds was tested in order to estimate potential dispersal by water. On the mown meadow, gaps had a slightly positive effect on germination rates, while greatly increasing seedling survival until the next spring. At the abandoned site, litter inhibited germination, whereas mosses had a negative effect on germination and a positive effect on survival rates during the first year after germination. Both germination and seedling establishment were negatively affected by the presence of slug herbivores. On the abandoned site, no seedlings at all survived until the next spring. Even though seeds of Succisella inflexa were capable to float for several weeks and to germinate thereafter, the situation at the field sites indicates that longdistance dispersal is highly unlikely. Our results showed that not only direct effects of abandonment, such as accumulation of litter, may have led to poor germination and poor seedling establishment of the species. Additionally, indirect consequences of changes in land use, such as higher seedling herbivory by slugs and successional vegetation changes due to abandonment, were important in determining habitat quality and availability of microsites for seedling recruitment. Furthermore, early mowing imposed seed limitation on plant populations.     

Mechanisms of structural change derived from patterns of seedling emergence and mortality in a semi-natural meadow

Questions: Is seedling emergence limited by the set of viable seeds, by incompatibility between the phenology of seed shedding and timing of mowing, or by dry weather in germination periods? Does seedling mortality fluctuate with season and weather? Location: Negrentino, southern Alps, Switzerland. Methods: Fecundity estimates of the dominant grass Bromus erectus; highly frequent counts of spontaneous seedlings by species and calculation of a community-level average mortality rate across 5 years; species-level records of seed shedding date and measurements of seed mass; measurement of soil moisture. Results: B. erectus produced 143.9 viable seeds/m²/year while the density of its seedlings was a 55 times smaller fraction. Grasses had fewer seedlings than forbs and their phenology of seed shedding was less compatible with mowing date. Soil moisture was a strong determinant of seedling emergence in spring and less so in autumn. Average seedling mortality declined with age of the populations and reached a maximum in an extremely dry summer. In relatively wet summers establishment success was positively related to seed mass. Conclusion: Community structure is susceptible to drought through mechanisms that selectively reduce recruits of coexisting plant functional groups. We propose that (1) more frequent intense droughts tend to reduce species that depend on frequent recruitment from seed, hence favour long-lived clonally spreading species, (2) drought timing selects between species with different germination phenology and drought resistance, and (3) drought impacts can be mitigated by changing management regimes that affect seed shedding.     

The effects of sod cutting and additional liming on potential net nitrification in heathland soils

The effects of sod cutting, a common restoration measure to remove excess nutrients from grass-dominated heathlands, on nitrification were studied in dry and wet Dutch heathlands and in incubation experiments. In the field, soil ammonium and nitrate concentrations were measured after treatment by sod cutting, with or without additional liming. Potential net nitrification was measured by incubating soil samples of all treatments with extra ammonium in a climate chamber at pH 6. Potential net nitrification of heaths dominated by Molinia caerulea was significantly higher than that of dwarf-shrub dominated heaths. Sod cutting of the former areas significantly decreased potential net nitrification, whereas in the latter areas no differences were found. Liming of sod-cut soils greatly increased potential net nitrification and the accumulation of ammonium in the soil up to toxic concentrations could be prevented. Our results show that the combination of sod cutting and liming would create suitable soil conditions for the germination and establishment of endangered plant species of dry and wet heathlands. The success of restoration projects of these areas can thus be increased.     

Sod cutting and soil biota effects on seedling performance

Sod cutting (i.e. top soil removal) is a restoration management option for enhancing seedling establishment and for lowering the nutrient concentration in eutrophicated soils of nutrient-poor species-rich grasslands. Removal of the upper soil changes not only abiotic soil properties but may also affect the resident soil community. We investigated the effects of sod cutting on the establishment and performance of two endangered plant species (Cirsium dissectum and Succisa pratensis) while simultaneously manipulating the interaction between seedlings and soil biota. In intact grassland and sod-cut areas at two localities, seedlings were grown in plastic tubes. Half of the tubes had a filter that excluded roots but allowed entry of fungal hyphae and soil microorganisms. The other tubes were closed (i.e. no contact with the surrounding soil). In a greenhouse experiment we studied the effect of soil solutions (with or without fungal tissue) from three grasslands and three sod-cut areas on seedling growth. Sod cutting had a positive net effect on seedling growth for S. pratensis. Access to (mycorrhizal) fungi and other soil biota resulted in a negative impact on seedling growth of both plant species, both in grassland and sod-cut areas. The greenhouse experiment confirmed that the soil biota in these meadows reduced seedling growth. Although sod cutting did not mitigate negative plant-soil feedback, it enhanced seedling growth, presumably by decreasing competition for light. Sod cutting is therefore very useful when seedling establishment needs to be stimulated.     

Influence of management on the seed production and seed bank of calcareous fen species

Abstract. The dynamics of the seed production and seed bank of two dominant perennial species of calcareous fens were investigated in response to different mowing frequency, fallow or pasture. The production of fertile shoots responded strongly to management practices. On sites where mowing alternated with fallow, autumn mowing led to an increase of fertile shoot density in Schoenus ferrugineus and a decrease in Molinia caerulea in the subsequent season. Repeated annual mowing after long-term fallow raised the density in Schoenus and inhibited initiation of flowering shoots in Molinia. Seed set and viability were influenced mainly by climatic or other factors. Variation of seed production between sites and in consecutive years at the same site could thus be large. The relationship between total seed production and seed bank was weak, but it improved when germination rates were included in the calculation. As the seed bank was generally smaller than the viable seed production, low survival and a high turnover of seeds in the upper soil layers is assumed for both species. Implications for recruitment strategies are discussed.     

Long‐term influence of sod cutting depth on the restoration of degraded wet heaths

Temperate heaths have an unfavorable conservation status in most European biogeographical regions. Increasing nitrogen levels promote competitive grass species such as Molinia caerulea, which is a main threat to heathland conservation in Europe. This article investigates the long‐term influence of sod cutting and the resulting changes in soil properties on the heath composition, integrity, and structure. In 15 nature reserves across the northern half of Belgium, we used (1) a large number of plots (203); (2) a broad range of sod cut depths (2–40 cm), and (3) a temporal dimension that describes how long the effects of sod cutting persist (census up to 19 years after sod cutting). Multivariate analyses were used in order to explore the influence of sod cut depth and time after sod cutting on the soil and vegetation properties. There was a positive relationship between sod cut depth and soil pH and water level, and a negative relationship with Al³⁺, NH₄⁺, and total organic matter (TOM). However, only a limited number of typical (target) species appeared after sod cutting, and then only weakly. Most of the time they remained a minor component of the restored vegetation. Moreover, M. caerulea reappeared and its cover significantly increased during the years following sod cutting. Although we were able to show that sod cut depth has a differential effect on soil properties and vegetation recovery, it also appeared that sod cutting does not restore wet heaths in the long term when applied in regions with high nitrogen deposition.     

Taxonomic importance of seed morphology in Gentiana (Gentianaceae)

The seeds of Gentiana show high diversity in size, shape and surface pattern. Until now, seeds of only a limited number of taxa have been studied in detail and their utility in the subgeneric classification of Gentiana has not been evaluated. In this study we investigated seed micromorphology of the poorly known sections Pneumonanthe, Frigidae and Isomeria. In order to evaluate the relevance of seed sculpturing for taxonomy, we selected qualitative characters of seed shape and testa ornamentation of all documented sections and performed cluster analyses based on presence and absence of coded characters. Several new scanning electron microscopy pictures of seeds provided a number of valuable characters for the subgeneric differentiation of Gentiana. The results of the cluster analyses of seed characters generally support the sectional classification given by former authors. However, the European Gentiana asclepiadea does not group together with other taxa of section Pneumonanthe, but clusters near to the strictly European section Gentiana. Gentiana froelichii, endemic to the south‐east Alps and treated traditionally within section Frigidae, clusters together with taxa of the European section Ciminalis. Our results reveal that seed micromorphology supports the reassignment of some taxa incertae sedis within Gentiana. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 101–115.     

Fire increases aboveground biomass, seed production and recruitment success of Molinia caerulea in dry heathland

During the last decades, the perennial tussock grass Molinia caerulea has shown an increased abundance in European heathlands, most likely as a result of increased nitrogen deposition and altered management schemes. Because of its deciduous nature, Molinia produces large amounts of litter each year, which may affect the intensity and frequency of accidental fires in heathlands. These fires may influence plant population dynamics and heathland community organization through their effects on plant vital attributes and competitive interactions. In this study, fire-induced changes in competitive ability and invasiveness of Molinia through changes in biomass production, seed set and seed germination under both natural and laboratory conditions were investigated. We found that fire significantly increased aboveground biomass, seed set and germination of Molinia. Seed set was twice as high in burned compared to unburned heathland. Two years after fire, seedling densities in natural conditions were on average six times higher in burned than in unburned heathland, which resulted in increased abundance of Molinia after burning. The seed germination experiment indicated that seeds harvested from plants in burned heathland showed higher germination rates than those from unburned heathland. Hence, our results clearly demonstrate increased invasive spread of Molinia after large and intense fires. Active management guidelines are required to prevent further encroachment of Molinia and to lower the probability of large fires altering the heathland community in the future.     

Re-establishment of dominance by dwarf shrubs on grass heaths

Experimental fields were established at three sites on different soil types in Dutch heathlands, where grass species have become dominant, in order to find methods for re-establishment of an ericoid dwarf-shrub heath. Treatments included mowing, mowing and cutting for hay, ploughing, milling, sod cutting and burning.Re-establishment of ericoid dwarf shrubs was only observed after creation of gaps or patches of open soil. The results after nine years for two of the sites and six years for the other site indicate that sod cutting is the most suitable method provided that the mineral top soil is not removed or disturbed. More profound removal of the sod including mineral soil causes a delay in re-establishment of ericoid dwarf shrubs and an increase of grasses.Factors that may explain different responses to gap creation including availability of seeds, soil moisture, soil fertility and plant survival strategies are briefly discussed.Abbreviations Calluna = Calluna vulgaris - Deschampsia = Deschampsia flexuosa - Erica = Erica tetralix - Molinia = Molinia caerulea - Sarothammus = Sarothammus scoparius     

Effects of experimental and real land use on seedling recruitment of six fen species

Seedling emergence and early establishment of six fen species differing in seed mass and growth form were investigated under experimental land use with changed vegetation structure and under real land use in a calcareous fen. Seeds of all six species were sown in plots with different experimental land-use treatments: summer and autumn mowing with or without litter removal, trampling and abandonment. Additionally, emergence and survival of experimentally sown seeds was investigated under real land use on adjacent sites managed by mowing, grazing, intense trampling or abandonment.On abandoned plots and on plots without litter removal of the land-use experiment, emergence rates of all species were negatively affected either by high litter and moss cover or by tall canopy. No differences were found between autumn and summer mowing. Gap creation by experimental trampling did not increase germination rate. Under real land use, establishment of seedlings of most species was positively affected by litter cover and tall canopy. Trampling, in contrast, had a severe negative effect on seedling survival.The investigated species differed in their germination ability which was tested in the germination chamber and in their response to land use. Succisa pratensis with the highest seed mass germinated well in the chamber and in the field more or less regardless of land use. The low germination rate of Parnassia palustris in the germination chamber indicated a limitation of viable seeds. In the field, however, seedling emergence was additionally limited by microsite availability. Seeds of Serratula tinctoria and Primula farinosa germinated well in the germination chamber, but seedling recruitment in the field was hampered in the presence of a high litter or moss cover. Seeds of Tofieldia calyculata and Pinguicula vulgaris were strongly dependent on the availability of suitable microsites in the field. They hardly germinated under natural conditions, in spite of a high number of germinable seeds in the germination chamber.     

Effects of removal of organic matter on the productivity of heathlands

Abstract. Effects of milling, mowing and sod cutting on productivity in a Deschampsia grass heath and a Molinia-Deschampsia grass heath were studied from 1977 to 1986. The sum of above-ground biomass, litter and accumulated humified matter (TOM) in both types was ca. 70 ton/ha and the primary productivity 3–5 ton ha^-1yr^-1. Mulching did not affect the annual production. Sod cutting reduced the productivity to 1–2 ton ha^-1yr^-1; on the Molinia-Deschampsia site this reduction lasted for at least a decade, while the Deschampsia heath started to recover from sod cutting soon and increased in productivity again. Annual mowing both with and without removal of hay reduced the production as well, particularly on the Molinia-Deschampsia heath. Lower production was either the result of phosphorus depletion - Molinia-Deschampsia site - or nitrogen depletion - Deschampsia site. In the mowing treatments there was also a depletion of carbohydrate reserves.     

Plant species decline due to abandonment of meadows cannot easily be reversed by mowing. A case study from the southern Alps

Abstract. We resumed mowing in two plots of ca. 100 m² in an abandoned meadow dominated by Brachypodium pinnatum on the slope of Monte Generoso (Switzerland). We monitored species composition and hay yield using point quadrats and biomass samples. Species frequencies changed little during 10 yr (1988–1997) while hay yields showed large fluctuations according to mean relative humidity in April-June. We performed a seed-addition experiment to test whether the establishment of meadow species is limited by lack of diaspores or favourable microsites for germination and recruitment from the seed bank. We sowed ca. 12 000 seeds of 12 species originating from a nearby meadow individually in plots of a 4 × 6 unbalanced Latin square with four treatments, burning, mowing, mowing and removal of a layer of decayed organic matter, and a control. We monitored the fate of seedling individuals for 24 months. Seedlings of all species were established and survived for 12 months, 10 species survived during at least 24 months, some reached a reproductive stage. Species responded to different qualities of microsites provided by the different treatments thus required different regeneration niches. Spontaneous long-distance immigration was insignificant. We conclude that the former species composition of abandoned meadows cannot easily be restored by mowing alone because many plant species of meadows do not have persistent seed banks and immigration over distances of more than 25 m and successful establishment is very unlikely.     

Comparative ecology of seedling recruitment in an oligotrophic wet meadow

Abstract. For the regeneration niche to contribute to the maintenance of species diversity interspecific differences in sensitivity of seedling recruitment to environmental conditions is assumed. We experimentally tested differences between meadow species for the response of seed germination to chilling, and sensitivity of seedling recruitment to microscale heterogeneity. We also compared the dynamics of seedling recruitment in gaps. Seed germination was tested in standard laboratory germination tests, comparing control seeds with seeds chilled at +4 °C, and at –14°C for one month. Species responses varied from significant increases in germinability after chilling (e.g. Cirsium palustre, Betonica officinalis, Angelica sylvestris) to significant decreases (e.g. Hieracium umbellatum, Succisa pratensis, Selinum carvifolia). In some species, chilling at + 4 °C has a similar effect to chilling at –14 °C, in others the effect of chilling at + 4°C was intermediate, and in some, there was no effect of chilling at + 4°C, but an effect of chilling at –14°C. Different chilling temperatures also affect timing and speed of seed germination under greenhouse conditions. The dynamics of seed germination under field conditions was studied by sowing seeds into artificially created gaps and following their germination, both where the seedlings were removed after emergence and where they were not removed. Species differ in their germination dynamics: they all start late April, but then differentiate from an abrupt maximum and early finish of germination (in the second half of May), to prolonged germination without a marked maximum. Seedling removal increased the total number of germinated seeds, with a marked density dependence at this stage. Seeds were also sown into plots with treatments (1) gaps, sod stripped, above-ground vegetation removed; (2) mown, moss layer removed; (3) mown; (4) untouched control. Seedling emergence was monitored for 3 yr. Seedling recruitment decreased from treatments (1) to (4), but sensitivity differed between species. With increasing seed weight, the difference between gaps and other treatments decreased. The results show that there are considerable differences in seedling recruitment sensitivity between species.     

Reduced light availability and increased competition diminish the reproductive success of wet forest sedge Carex loliacea L.

Wet forest ecosystems in temperate regions have been heavily drained and logged, often with significant negative consequences for biodiversity in these habitats. Our research focused on population maintenance mechanisms of a declining wet forest sedge Carex loliacea L. We studied germination under different light regimes and seedling survival under different vegetation densities using an in situ removal experiment. For successful germination, seeds of C. loliacea need light; germination in reduced light conditions is depressed. The seeds of C. loliacea are able to accumulate a seed bank and exhibit seasonal dormancy cycles. Survival of seedlings strongly depends on competition with other plant species. Our results imply that changes in habitat conditions (draining, forest cutting) affect the successful generative reproduction of C. loliacea primarily via a change in light conditions, which is a strong factor both at the stage of germination and seedling growth. However, adult plants are able to persist over a much broader range of habitat conditions without detectable vitality loss.     

COMPARATIVE REPRODUCTIVE BIOLOGY OF ALPINE BIENNIAL AND PERENNIAL GENTIANS (GENTIANA: GENTIANACEAE) IN CALIFORNIA

Reproductive characteristics of three sympatric species of Gentiana exhibiting perennial and non-perennial life histories were studied in alpine meadows of the White Mountains of California during three consecutive years. High fruit and seed set and the production of a relatively large number of seeds characterized the alpine biennials, Gentiana tenella and G. prostrata. In contrast, fruit and seed set were considerably reduced and yearly seed production was relatively low in the alpine perennial, G. newberryi. Successful seed production in the biennial species was a result of low rates of flower predation, self-pollination, and the allocation of a comparatively high proportion of biomass to reproductive structures. Seed production in the perennial species was limited by high rates of flower predation, insufficient amounts of pollen reaching stigmas, an unusually late flowering period, and by the relatively small proportion of biomass allocated to reproduction.     

Effects of Frequent Mowing on Survival and Persistence of Forbs Seeded into a Species-Poor Grassland

Many early attempts at tallgrass prairie reconstruction failed to achieve the high species diversity of remnant prairies, and instead consist primarily of C₄ grasses. We hypothesized that frequent mowing of established prairie grasses could create sufficient gaps in the aboveground and belowground environment to allow for the establishment of native forbs from seed. We studied forb seedling establishment in a 25‐year‐old prairie planting in northern Iowa that was dominated by native warm‐season grasses. In winter 1999, 23 species of native forbs were broadcast into the recently burned sod at a rate of 350 viable seeds/m². Treatment plots were mowed weekly for either one or two growing seasons, and control plots were unmowed. Mowed plots had greater light availability than controls, especially when warm‐season grasses began to flower. Overwinter seedling mortality was 3% in mowed treatments compared to 29% in the controls. Forbs in mowed plots had significantly greater root and shoot mass than those in control plots in the first and second growing seasons but were not significantly more abundant. By the fourth growing season, however, forbs were twice as abundant in the mowed treatments. No lasting negative impacts of frequent mowing on the grass population were observed. Mowing a second year influenced species composition but did not change total seedling establishment. Experimental evidence is consistent with the idea that mowing reduced competition for light from large established grasses, allowing forb seedlings the opportunity to reach sufficient size to establish, survive, and flower in the second and subsequent years.     

Fire and regeneration: the role of seed banks in the dynamics of northern heathlands

Questions: How do species composition and abundance of soil seed bank and standing vegetation vary over the course of a post‐fire succession in northern heathlands? What is the role of seed banks – do they act as a refuge for early successional species or can they simply be seen as a spillover from the extant local vegetation? Location: Coastal Calluna heathlands, Western Norway. Methods: We analysed vegetation and seed bank along a 24‐year post‐fire chronosequence. Patterns in community composition, similarity and abundances were tested using multivariate analyses, Sørensen's index of similarity, vegetation cover (%) and seedling counts. Results: The total diversity of vegetation and seed bank were 60 and 54 vascular plant taxa, respectively, with 39 shared species, resulting in 68% similarity overall. Over 24 years, the heathland community progressed from open newly burned ground via species rich graminoid‐ and herb‐dominated vegetation to mature Calluna heath. Post‐fire succession was not reflected in the seed bank. The 10 most abundant species constituted 98% of the germinated seeds. The most abundant were Calluna vulgaris (49%; 12 018 seeds m^?2) and Erica tetralix (34%; 8 414 seeds m^?2). Calluna showed significantly higher germination the first 2 years following fire. Conclusions: Vegetation species richness, ranging from 23 to 46 species yr^?1, showed a unimodal pattern over the post‐fire succession. In contrast, the seed bank species richness, ranging from 21 to 31 species yr^?1, showed no trend. This suggests that the seed bank act as a refuge; providing a constant source of recruits for species that colonise newly burned areas. The traditional management regime has not depleted or destroyed the seed banks and continued management is needed to ensure sustainability of northern heathlands.     

Fire increases aboveground biomass,seed production and recruitment success of Molinia caerulea in dry heathland

During the last decades, the perennial tussock grass Molinia caerulea has shown an increased abundance in European heathlands, most likely as a result of increased nitrogen deposition and altered management schemes. Because of its deciduous nature, Molinia produces large amounts of litter each year, which may affect the intensity and frequency of accidental fires in heathlands. These fires may influence plant population dynamics and heathland community organization through their effects on plant vital attributes and competitive interactions. In this study, fire-induced changes in competitive ability and invasiveness of Molinia through changes in biomass production, seed set and seed germination under both natural and laboratory conditions were investigated. We found that fire significantly increased aboveground biomass, seed set and germination of Molinia. Seed set was twice as high in burned compared to unburned heathland. Two years after fire, seedling densities in natural conditions were on average six times higher in burned than in unburned heathland, which resulted in increased abundance of Molinia after burning. The seed germination experiment indicated that seeds harvested from plants in burned heathland showed higher germination rates than those from unburned heathland. Hence, our results clearly demonstrate increased invasive spread of Molinia after large and intense fires. Active management guidelines are required to prevent further encroachment of Molinia and to lower the probability of large fires altering the heathland community in the future.     

Effects of cutting and nitrogen deposition on biodiversity in Cantabrian heathlands

Questions: Does the diversity of heathland vegetation change when subjected to experimental disturbances such as cutting and nitrogen fertilization? Do changes in the vegetation structure negatively affect the regeneration of the dominant species Calluna vulgaris? Is cutting an alternative method of conserving the diversity and maintaining the structure of heathlands in the Cantabrian Mountains? Location: Calluna vulgaris heathlands on the southern slopes of the Cantabrian Mountain range, NW Spain. Methods: A total of 60 plots were treated with different combinations of cutting and twice the estimated atmospheric deposition of nitrogen (56 kg‐N.ha^?1.yr^?1). The changes in the cover values of the species present were monitored over a five year study period. The cover values were used to calculate abundance and species richness. Results: Fertilizing with nitrogen allows biodiversity to increase over time. However, the greatest biodiversity is associated with the cutting plus fertilization treatment, since cutting allows gaps to be opened that are easily colonized by pioneer annual species, while fertilization mainly favours an increase in the mean number of perennial herbs (graminoids and forbs). Increased perennial herb richness also corresponds to a rise in their cover values. The recovery of the dominant woody species in these communities, Calluna vulgaris, is not impeded by the increase in perennial herbs species' cover values. Conclusions: In the Calluna vulgaris heathlands studied, cutting plus fertilization allowed an increase in biodiversity over time. No displacement of the dominant woody species, Calluna vulgaris, is observed due to the presence of the perennial herbs. Cutting patches of heathland is recommended as a mechanism for maintaining high vegetation diversity, when grazing is not possible.