Competition between heather and grasses on Scottish moorlands: Interacting effects of nutrient enrichment and grazing regime

Abstract. Considerable losses and degradation of heathlands (in moorlands and lowlands) have been reported across Europe, with Calluna vulgaris (heather) being replaced by other species, often grasses. Increasing atmospheric nitrogen deposition and overgrazing have been suggested as the driving factors behind this change. This possibility was investigated in a study of the interacting effects of nutrient inputs and grazing on heather and three grass species (Nardus stricta, Deschampsia cespitosa and D. flexuosa) in the field, on a moorland in northeastern Scotland. In addition, the interacting effects of increasing nutrients and Calluna canopy height on N. stricta and D. cespitosa were studied using turves in an outdoor experimental area. In the field, fencing had a larger effect than fertilizer on the growth of all species, except for N. stricta, the species most unpalatable to herbivores. Fencing led to an increase in the height of the Calluna canopy, which may reduce light availability for the grasses. In the turf experiment, the height of the Calluna canopy affected the diameter of the grass tussocks and percentage of green matter (i.e. live leaf material), with plants under the more closed Calluna canopies being smaller. This study suggests that the slow‐growing, evergreen Calluna is a more effective competitor than the faster growing grasses when it has a tall, intact canopy, even at increased levels of nutrient supply. However, overgrazing promotes gap formation in the Calluna canopy, providing an opportunity for grasses to take advantage of increased nutrients. Thus the conservation of heather moorlands requires an understanding of the grazing level which allows Calluna to maintain sufficient canopy structure to outcompete grasses for light.     

Vegetation re‐establishment on land previously subject to control of Pteridium aquilinum by herbicide

Abstract. Vegetation development after conventional bracken control with herbicide in the uplands is often slow and rarely results in the development of the target community. Two experiments were established on contrasting sites to investigate cost‐effective methods for the re‐establishment of vegetation dominated by Calluna vulgaris (heather), which is the usual target community seen as the desired replacement for Pteridium aquilinum in the uplands by many land managers. At one site, where grazing intensity was low and shelter was sufficient to prevent rapid Pteridium litter loss, the necessary treatments to establish Calluna were to disturb the litter and add a source of Calluna seeds. A difference between continued Pteridium control or a relaxation of control was not observed over the time‐scale of the experiment. At the second site, where grazing intensity was high and shelter minimal, vegetation dominated by Rumex acetosella had developed on a thin but dense litter layer. To enhance Calluna regeneration it was necessary to prevent sheep grazing, disturb the remaining litter layer and add a source of Calluna propagules. A nurse crop hindered Calluna establishment due to its vigorous growth. At neither site was an initial application of fertilizer an aid to Calluna establishment. It is possible to devise methods for the re‐establishment of Calluna at sites where previous attempts at Pteridium control alone have not had the desired effect on vegetation development. However, the precise choice of methods used is dependent on the grazing management at the site. High levels of livestock grazing prevented Calluna establishment, but some trampling may aid litter breakdown. The ability to adjust stocking rates and access to the site by machinery are critical factors to take into account in producing an effective strategy for long‐term Pteridium control and its replacement by target vegetation.     

Effects of nutrient supply,light availability and herbivory on the growth of heather and three competing grass species

Grasses are becoming more abundant in areas in NE Scotland which until recently were dominated by heather (Calluna vulgaris). However, it is not clear if grasses are aggressive competitors which are now able to outcompete the dwarf shrub due to changes in environmental factors (such as grazing pressure and increasing nutrient inputs), or just opportunistic invaders, occupying gaps in the canopy which occur when heather reaches the degenerate stage. Experiments in turves and in field plots were carried out in order to investigate the performance of three grass species, Nardus stricta, Deschampsia cespitosa and Deschampsia flexuosa growing in competition with heather. These three species were selected because they differ in their nutrient requirements, palatability to herbivores and tolerance of shading. The grasses were planted in heather canopies of different structure, either turves of heather of different height and age, or moorland plots with or without heavy grazing by sheep and deer. Fertiliser (NPK) was applied to half the experimental plants. The growth of the grass species and the heather in response to the fertiliser and grazing treatments was measured, together with the light levels penetrating the canopy and received by the grass plants.Results indicated that heather was likely to be outcompeted by grasses only when there are gaps in the canopy, resulting either from heavy grazing or from the heather being in the mature or degenerate phase. Fertiliser enhanced plant growth whereas fencing out herbivores led to strong competition for light as the heather canopy closed. It is concluded that grasses require gaps in the canopy to successfully invade heather moorland, or they tend to be shaded out. Thus better management of heather moorlands to maintain a dense canopy structure may help to preserve heather cover even under increasing nutrient inputs.     

Conservation and restoration of Nardus grasslands in the Swiss northern Alps

Aim: Species‐rich Nardus grasslands are high nature‐value habitats. In Switzerland, many of these grasslands are degraded even though they have been under protection since the 1980s. Degradation shows two divergent trends: Nardus grasslands are either dominated by Nardus stricta or by eutrophic plants, both trends leading to the disappearance of typical Nardus grassland species. With this study, we aim to identify the factors that could be adjusted to conserve the integrity of this habitat. Location: Bernese Alps, Switzerland. Methods: In 2016, we investigated the underlying causes of this degradation process by assessing vegetation composition in 48 Nardus grasslands located in the Swiss northern Alps of canton Bern and linking it to soil, management and environmental variables. To explore the effect of the degradation on higher trophic levels, orthopteran species richness and densities were assessed. Results: Results show that Nardus meadows (mown) are rarely degraded compared to Nardus pastures (grazed). Within pastures, eutrophic plants are most abundant on small pastures with low soil carbon/nitrogen ratio, indicating high nutrient availability. Nardus stricta dominance is most problematic on north‐exposed slopes and in summer pastures. A plausible driver of both degradation trends is the grazing management regime: within small pastures at low elevation where the grazing periods are short but intense, soil carbon/nitrogen ratio is low because of high dung deposition, thus the eutrophic species become dominant. Contrastingly, on large summer pastures with low‐intensity and long‐term grazing, N. stricta becomes dominant due to selective grazing. Both degradation trends show a negative impact on the orthopteran density. Conclusion: Species‐rich Nardus grasslands are a precious alpine habitat for specialised plant species and orthopterans. With an extensive mowing regime or a more controlled grazing regime that homogenises intensity in time and space, species‐rich Nardus grasslands can be conserved in Switzerland.     

Management of heather for game and livestock

MOSS, R., 1989. Management of heather for game and livestock. Heather provides food and cover for moorland herbivores. Interactions between soils, drainage, climate, muirburn and grazing affect the species composition, structure and growth of moorland swards. Together, these aspects of a sward determine which and how many herbivores it can support. Grazing and burning maintain much heather ground which would otherwise revert to scrub or woodland. Heavy grazing by ungulates can turn a heathery sward into a graminaceous one, so reducing numbers of grouse and mountain hares. On good soils, invading graminoids are usually nutritious grasses and the carrying capacity for ungulates can increase. On poor soils, grazing and frequent burning may lead to swards dominated by poor quality grasses and sedges so that the carrying capacity for ungulates declines. One way of reinstating heather dominance is to remove livestock; where there are no heather plants left, other techniques may be necessary.     

Woodlands on farms in southern New South Wales: A longer-term assessment of vegetation changes after fencing

Summary Fencing incentive programmes have been widely used throughout Australia to assist landholders to fence remnant woodland vegetation, to control grazing and improve native vegetation condition. This study investigated vegetation and soil condition in remnant woodlands fenced for 7–9 years in the Murray catchment area in southern New South Wales. Surveys were undertaken at 42 sites, where vegetation condition was assessed in paired fenced and unfenced sites. Semi‐structured interviews were also conducted with landholders to gather management information. Woodlands surveyed were Yellow Box/Blakely's Red Gum (Eucalyptus melliodora/E. blakelyi, 15 sites), Grey Box (E. microcarpa, 13 sites) and White Cypress Pine (Callitris glaucophylla, 14 sites). Fencing resulted in a range of responses which were highly variable between sites and vegetation types. In general, fenced sites had greater tree regeneration, cover of native perennial grasses, less cover of exotic annual grasses and weeds, and less soil compaction than unfenced sites. However, there was greater tree recruitment in remnants to the west of the study area, and tree recruitment was positively correlated with time since fencing. Within sites, tree recruitment tended to occur in more open areas with a good cover of native perennial grasses, as compared to sites with a dense tree canopy, or dominated by exotic annuals grasses or weeds. Forty‐eight per cent of fenced sites had no tree regeneration. There was a significant decline in native perennial grasses, and increase of several unpalatable weeds in many fenced areas, suggesting certain ecological barriers may be preventing further recovery. However, drought conditions and associated grazing are the most likely cause of this trend. A range of grazing strategies was implemented in fenced sites which require further research as a conservation management tool. Continued long‐term monitoring is essential to detect key threats to endangered woodland remnants.     

Early regeneration of Calluna heathland under various fertilization treatments

Summary A sod-cutting and fertilization experiment was performed on a Calluna-dominated heathland in The Netherlands to determine appropriate management regimes for Calluna regeneration, and to further understand the nutrient responses of heathland species. Replicated permanent plots were analysed by multivariate techniques. Sod-cutting alone caused Calluna regeneration from its soil seed bank. A single fertilization at the start of the experiment caused initial vegetation differences which disappeared after a few years as the nutrients were lost from the system, except that one application of nitrogen enhanced the rate of Calluna regeneration. Repeated fertilization caused large differences in the vegetation: repeated nitrogen enhanced several bryophyte species while greatly inhibiting Calluna, repeated phosphate partly inhibited Calluna while greatly favouring several lichen species, and the most striking result of repeated calcium was also an increase in bryophytes, but the species were different from those favoured by nitrogen. Treatments which inhibited Calluna tended to increase species diversity because of the lessened Calluna dominance.     

The influence of ammonium nitrate on the root growth and ericoid mycorrhizal colonization of Calluna vulgaris (L.) Hull from a Danish heathland

Conversion of European heathlands to grassland has been reported as a response to increased nutrient availability, especially of nitrogen; a direct effect upon mycorrhizal colonization has been proposed as an likely explanation.This hypothesis was tested in a random block experiment with four blocks and four replicates on a Danish inland heath, Hjelm Hede. Ammonium nitrate was applied (0, 35, 50 and 70 kg N ha^–1 year^–1) to a stand of Calluna vulgaris (L.) Hull four times annually for 2 years. Calluna roots were sampled on four occasions in the 2nd year of the nitrogen treatment. The extent of ericoid mycorrhizal colonization was determined by direct observation of the roots using a line-intersection method. The nitrogen content of the current-year shoots of Calluna increased when they were treated with nitrogen. Nitrogen fertilization had no significant effects on ericoid mycorrhizal colonization of Calluna nor on root biomass. The seasonal variation in mycorrhizal colonization of the Calluna roots was highly significant. The spatial variability of mycorrhizal colonization, both in replicated plots and in the two contrasted soil horizons – the mor layer and the bleached sand – within the plots, were considerable. I conclude that heather decline under enhanced nitrogen input is unlikely to be caused by a direct impact on the ericoid mycorrhizae of Calluna. Received: 1 December 1998 / Accepted: 7 December 1999     

藏北古露高寒草地生态系统对短期围封的响应

过度放牧导致高寒草地生态系统退化,围封是生态保护和恢复的管理手段。以青藏高原那曲县古露镇过牧退化高寒草地为对象,系统分析了高寒草地生态系统的植被特征及土壤理化特性、土壤酶活性、土壤微生物生物量和群落结构对围封的响应。结果表明,短期围封后,(1)植被平均高度、盖度和地上生物量均有极显著增加(P0.01),而生物多样性指数则显著降低(P0.01);(2)土壤的水溶性有机碳含量、土壤物理结构(沙土与粉土的比例)及pH有显著变化(P0.05);(3)土壤酶活性没有明显改善;(4)土壤微生物生物量(细菌、放线菌、真菌)均呈显著增加(P0.05);(5)土壤中细菌的多样性有增加的趋势,其群落组成在门水平上也发生了变化;(6)Manteltest分析显示与土壤细菌群落结构的呈正相关性的环境因子主要为土壤有机碳含量(TOC)、总氮含量(TN)、碳磷比(C/P)与氮磷比(N/P)(P0.05)。这表明围栏封育有利于藏北草地植被、土壤理化特性的恢复,还能维持土壤微生物多样性,促进高寒草地生态系统的可持续发展。     

Further observations on a Callunetum intensively grazed by mountain hares (Lepus timidus L.)

Summary

Heavy utilisation by mountain hares checked the growth of Calluna so that mean height was only 12 cm thirty years after the 0.2 ha area was burnt. Cover and biomass increased gradually over the thirteen years studied, from 59 to 85% and 300 to 580 g/m² respectively. Differences in sward height were maintained, the greater utilisation of short heather being offset by greater growth. Compared to ruminant-grazed Calluneta, the hare-grazed sward had a distinctive, compact form, and biomass was great relative to cover and height because of the many shoots. Non-ericoid angiosperms had little cover.     

氮水添加对放牧背景下荒漠草原生产力的影响

水分与氮素作为干旱和半干旱草原生产力的共同限制性因子在退化草原的生态快速修复过程中备受关注。以不同放牧强度背景下的短花针茅荒漠草原为研究对象,开展围封模拟放牧利用实验,同时添加氮素和水分。通过分析历史放牧强度与年份对生产力的影响,以及添加氮素和水分对不同功能群植物生物量的作用,探讨放牧强度对短花针茅草原生产力的内在作用机制,以及如何实现荒漠草原资源合理开发和可持续利用。研究结果显示,降雨量与放牧强度决定着短花针茅草原的植物群落结构。氮素和水分添加可分别提升11%-29%和12%-32%的群落地上生物量,且二者存在显著的交互作用。不同功能群植物的地上生物量对氮素与水分添加的响应存在差异,多年生丛生禾草对氮素和水分添加响应最敏感。氮素与水分添加可显著提高多年生丛生禾草的地上生物量,但与自然降水量相关。氮素添加对地上生物量的影响在正常降雨和稍旱年份作用显著,而水分添加在干旱年份作用显著。在正常降雨年份,以半灌木植物为优势种的轻度放牧背景以添加水分对提升生产力最宜,以多年生丛生禾草和半灌木为共优种的中度放牧背景和以多年生丛生禾草为优势种的重度放牧以同时添加水分和氮素对提升生产力最为宜;在干旱年份不同放牧强度背景下均以同时添加水分和氮素对提升生产力最为宜。我们的结果表明了养分与资源的改善有利于退化短花针茅草原的快速恢复和可持续生产。     

Convergent grazing responses of different‐sized ungulates to low forage quality in a wet savanna

We studied how grazing intensity by small and mid‐sized ungulate grazers varied with nutritional quality and grass species composition in wet oligotrophic tallgrass savanna of coastal northern Tanzania. Average grazing intensity was low (3–15% by cover), and most grass species were scarcely used by herbivores. Only two grasses, Panicum infestum and Digitaria milanjiana, had nitrogen and phosphorus concentrations that were consistently above the minimum levels (e.g. nitrogen concentrations <7 mg g^?1) required by the three commonest grazers, Bohor reedbuck, waterbuck and wildebeest. The best predictors of grazing intensity were cover of P. infestum (the most abundant grass, with a mean cover of 15%) and canopy height of ungrazed vegetation. Models did not contain separate predictors for nutritional quality, presumably because quality varied mainly at the grass species level and therefore was fully represented by the variable ‘cover of P. infestum’. Given that the three grazers differed greatly in body size and muzzle width (parameters known to influence nutrient requirements and the ability of grazers to feed selectively at the smallest spatial scale), we expected there to be strong resource partitioning that would be detectable in terms of grazing strategies and feeding sites. However, apart from minor differences in canopy height, greenness and diameter of grazed patches (albeit consistent with our expectations), feeding stations of the three grazers were similar and strongly dominated by P. infestum. We conclude that the low quality of herbage in wet oligotrophic savannas restricts foraging choices, which produces a characteristic yet impoverished grazing community that exhibits only limited resource partitioning.     

Experimental evidence from Cantabrian mountain heathlands suggests new recommendations for management of Vaccinium myrtillus L

Background: Management actions in mountain heathlands oriented to increasing the cover of Vaccinium myrtillus promote mowing of Calluna vulgaris to avoid competition. However, such action ignores the fact that plant–plant interactions range from competition to facilitation under different stress conditions.

Aims: To test whether the interactions between these two species are related to competition or facilitation, which would change the perspective for their management in mountain heathlands.

Methods: A total of 40 experimental plots located in a montane heathland (northern Spain) were selected. Calluna or Vaccinium plants were either cut at ground level or not cut, and plots were fertilized with nitrogen or not in a three-factorial design. Vegetative and reproductive shoot mass and length of the current year’s shoots were estimated in two consecutive years for both species. Herbivory on Vaccinium by browsing ungulates was compared in control plots and plots where Calluna had been cut 2 yrs ago.

Results: The two study species co-occurred in 72.4% of the sampled plots, although Calluna was more abundant. Vaccinium growth was greater in plots with Calluna than in plots where Calluna had been cut. Browsing on Vaccinium was also higher in plots were Calluna had been cut, and the effect of fertilization was only significant on the reproductive biomass of Calluna.

Conclusions: Contrary to initial proposals, our results suggest that Vaccinium does not always benefit from the cutting of Calluna in heathlands. Thus, the management of Cantabrian mountain heathlands should consider maintaining Calluna in order to facilitate the growth of Vaccinium.     

Long‐term livestock exclusion facilitates native woody plant encroachment in a sandy semiarid rangeland

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Field responses to added organic matter, arbuscular mycorrhizal fungi, and fertilizer in reclamation of taconite iron ore tailing

A three season study was conducted to determine the effect of added composted yard waste, arbuscular mycorrhizal (AM) fungi, and fertilizer on plant cover, standing crop biomass, species composition, AM fungal infectivity and spore density in coarse taconite iron ore tailing plots seeded with a mixture of native prairie grasses. Plant cover and biomass, percent seeded species, mycorrhizal infectivity and spore density were greatly increased by additions of composted yard waste. After three seasons, total plant cover was also greater in plots with added fertilizer. Third season plant cover was also greater in plots amended with the higher rate (44.8 Mg ha^{–1) of compost than the moderate rate (22.4 Mg ha-1}). Field inoculation with AM fungi also increased plant cover during the second season and infectivity during the first two seasons. Seeded native species, consisting mostly of the cover species Elymus canadensis, dominated plot vegetation during the second and third seasons. Dispersal of AM fungal propagules into nonmycorrhizal plots occurred rapidly and increased infectivity in compost-amended plots during the third season. In plots with less than 10% plant cover, AM fungal infectivity of inoculated plots was greatly reduced after the second season. The high level of plant cover and the trend of increasing proportion of mycorrhizal-dependent warm-season grasses, along with increases in infectivity, forecast the establishment of a sustainable native grass community that will meet reclamation goals.     

Further evidence for the importance of residual leaf protein complexes in litter decomposition and the supply of nitrogen for plant growth

Summary The nitrogen absorbed by birch seedlings grown in sand culture has been used as a measure of the resistance to decomposition of complexes formed in vitro by interaction between protein and water-extractable leaf constituents ofCalluna vulgaris, Chamaenerion angustifolium andCircaea lutetiana. The resistance to decomposition of these complexes measured in this way isCalluna vulgaris >Chamaenerion angustifolium >Circaea lutetiana. This result seems to be in agreement with the soil-forming and field characteristics of these species and especially the raw humus-forming tendencies ofCalluna vulgaris.The nitrogen released, as measured by the nitrogen absorbed by birch seedlings, fromCalluna raw humus (H-layer material) and the model protein —Calluna leaf extractives complex is very nearly proportional to the widely different amounts of total nitrogen initially added to the root environment thereby suggesting that the organic nitrogen of the model complex and of theCalluna raw humus have similar properties.The amount of nitrogen absorbed by the birch seedlings when unaltered protein was added to the sand culture was less than when protein —Circaea lutetiana leaf extractives complex was added. This is ascribed to too rapid release and loss of nitrogen before it could be absorbed by the birch seedling roots and suggests a role for some leaf protein complexes in the conservation of litter nitrogen. Some possible effects of protein complexes in relation to the digestion of fresh and dry leaf material of various plant species by animals are discussed.Information from archeological observations and from investigations into the tanning of proteins is considered in relation to possible factors influencing the decomposition of protein complexes of litter and it seems probable that a less acid reaction,e.g. such as is brought about by the addition of adequate amounts of calcareous material, will assist the mobilization of the nitrogen of raw-humus-forming litter.The contention that the protein-precipitating substances of leaves are of fundamental importance in soil processes, especially raw humus formation and the supply of nutrients for plant growth, is considered to be supported by experimental approaches from different directions.     

Effects of rain, nitrogen, fire and grazing on tree recruitment and early survival in bush-encroached savanna, South Africa

Moisture, nutrients, fire and herbivory are the principal factors governing tree–grass cover ratios of savannas. We investigated tree (Acacia mellifera) recruitment after fire and under conditions of maximum-recorded rainfall, nitrogen addition and grazing in a completely-crossed field experiment. We employed a similar garden experiment with the exception of the fire treatment. Tree germination in the field was extremely low, probably due to below-average natural rainfall in plots that only received natural rain, and insufficient watering frequency in irrigated plots. Due to low germination in the field experiment, no treatment significantly affected tree recruitment. In the garden experiment, frequent watering, nutrient control (i.e. no nitrogen addition) and grazing enhanced tree recruitment with significant interactions between rain, nitrogen and grazing. We infer that above-average rainfall years with frequent rainfall events are required for mass tree recruitment. Grass defoliation makes space and resources available for tree seedlings. Nitrogen enrichment increases the competitive ability of fast-growing grasses more than that of the N₂-fixing tree component. In contrast to conventional wisdom that grazing alone causes encroachment, we suggest that there are complex interactions between the above-mentioned factors and ‘triggering’ events such as unusually high rainfall.     

Jährliche Schwankungen der Individuenzahl in einer nordwestdeutschen trockenen Heide II

Summary The vegetation of a permanent quadrat situated in a Northwest German Calluno-Genistetum typicum community has been analysed every year from 1955 to 1959 (cf. Vegetatio X, p. s53–56, 1961) and again from 1960 to 1964.In the winter of 1955/56, when there were temperatures of –27° and an incomplete snow-cover, Calluna vulgaris had died in large amount. Nevertheless it increased again during the following years. The dry year of 1959 again caused a 35% reduction of this heather, but by 1960/61 a good recovering of Calluna was observed. Again from 1961 to 1964 Calluna decreased from 85% to 30%, but we have no satisfactory explanation for this fact.Probably the decline from 1959 to 1960 of the young plants of Quercus robur has been caused by the cutting off, the reduction from 1962 to 1964 by sheep grazing. The increase from 1960 to 1961 may have been caused by sowing by jays. The sudden appearance of Betula verrucosa and B. pubescens (1961) and their remarkable increase is the result of the invasion of large numbers of seeds and of the small vegetation-cover of the soil, perhaps also the result of the damaging of the cover of heathhumus by the sheep. Certainly the other higher plants have been imported by rabbits or sheep. They partially disappeared again after a few years.Perhaps the dryness of the year 1959 also caused the retrogression of the moss from 1958 (50%) to 1959. Maybe the sheep caused the further diminution of the moss and the decline of the cladonias. As expected, the macrofungi and the thalli of Parmelia physodes f. minor appeared during wet summers in especially great number, while dry periods caused their decay.     

Exclusion of livestock grazing and wood collection in dryland savannah: an effect on long‐term vegetation succession

Sahelian savannah faces increasing pressure from human activities, leading to its degradation. The aim of this study was to investigate the possibility of restoration of dryland savannah vegetation by the elimination of disturbance factors on the ecosystem. Is degraded dryland savannah vegetation able to be restored by means of natural succession? What is the timescale for its recovery? The study took place in the Bandia Reserve, 65 km south‐east of Dakar (Senegal), a unique site with two successional stages due to the elimination of uncontrolled exploitation. The vegetation structure of 15 years (15YRS) and 5 years (5YRS) after fencing was compared with vegetation exposed to continuous livestock grazing and wood collection outside the fenced area. Calculated by redundancy analysis, a significant effect of selected areas on the cover of all species was revealed and successional stage explained more than 45% of data variability. Perennial forbs, annual forbs and perennial grasses achieved the highest cover in 5YRS, woody species in 15YRS and annual grasses in the area outside of the fenced reserve. The dominant woody species Acacia seyal, A. ataxacantha, A. nilotica subsp. adstringens and Balanites aegyptiaca reconstituted the dense formation of Acacia bushland by means of natural succession in the 15YRS area.