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.     

Impact of prescribed burning on the nutrient balance of heathlands with particular reference to nitrogen and phosphorus

Question: Can prescribed winter burning compensate atmospheric nutrient loads for dry heathlands? What effects does prescribed burning have on nutrient balances, particularly as regards the limiting nutrients N and P? Location: Lueneburg Heath, NW Germany. Methods: In two burning experiments (in 10/15 year old Calluna‐stands) nutrient balances (for N, Ca, K, Mg, P) were calculated by analysing nutrient inputs (atmospheric deposition, ash deposition), nutrient stores (above‐ground biomass, organic horizon) and nutrient outputs (biomass combustion, leaching). Results: Atmospheric nutrient deposition amounted to 22.8 kg.ha^‐1.a^‐1 for N and < 0.5 kg.ha^‐1.a^‐1 for P. Nutrient stores in the above‐ground biomass were 95/197 kg.ha^‐1 for N and 5/13 kg.ha^‐1 for P (first/second experiment, respectively). From these stores 90/53% (for N) and 25/14% (for P) were removed by burning. Effects of leaching on nutrient balances were low. In the first two years after burning, leaching rates of N increased by about 4/6 kg.ha^‐1, whereas leaching rates of P did not change significantly. Input/output‐ratios showed that prescribed burning leads to positive nutrient balances for N, Ca and Mg in the long term. For example, the amounts of N removed by prescribed burning are equivalent to ca. five years of atmospheric inputs. Applied in ten‐year cycles, this measure alone cannot prevent N accumulation in the long term. Conclusion: Regarding 10/15 year old Calluna‐heaths, we assume that prescribed burning cannot compensate for atmospheric N inputs, thus making long‐term changes in the nutritional state inevitable. Therefore, prescribed burning should be applied in combination with high‐intensity management measures.     

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.     

Interactive effects of clipping and nutrient availability on the compensatory growth of a grass species

Resource availability is an important factor affecting the capacity of compensatory growth after grazing. We performed a greenhouse experiment with Poa bulbosa, a small perennial grass of the Mediterranean and Central Asian grasslands, to test the importance of nutrient availability for compensatory growth after clipping. We also compared the results with predictions of the limited resource model (LRM). Plants were grown at low and high fertilization levels and subjected to a clipping treatment. Contrary to the LMR, we found that in Poa plants compensatory growth occurred under the high fertilization level, while it did not occur under the low level. The LMR predicts a higher tolerance for grazing in the stressful environment. Our plants showed a significant decrease in their relative growth rates (RGR) after clipping. Although the plants allocated a 32–188% greater fraction of the mass to lamina growth after clipping, this greater allocation to the leaves did not fully compensate for the initial reduction in leaf area ratio (LAR). A sensitivity analysis showed for the clipped plants under the high fertilization treatment, that changes in leaf allocation (f _lam) enabled the plants to compensate for a part of the potential loss caused by defoliation. Probably, the increased biomass allocation comes largely from the bulbs. We conclude that the inconsistency of the LRM with our results originates in the lack of compensatory mechanisms in the model. To better understand how environmental conditions affect tolerance to herbivory, the effects of compensatory growth should be taken into account.     

Differential effects of ammonium and nitrate on three heathland species

Arnica montana and Cirsium dissectum, typical of species-rich heathlands and acidic grasslands, have declined rapidly in The Netherlands in recent years. Field surveys suggest that the decline is caused by soil acidification as a result of enhanced atmospheric N and S deposition. Therefore, the survival, growth and development of these species were studied in a water culture experiment, using nutrient solutions which differed both in mineral nitrogen form and in ammonium concentration. For comparison, the performance of a third, acid tolerant species, Calluna vulgaris, was studied. The results showed that both Arnica and Cirsium performed better using nitrate than when using ammonium as a sole nitrogen source, whereas ammonium toxicity became apparent when ammonium concentrations were raised above 100 µM. Ammonium toxicity was expressed by an increase in mortality of Arnica plants with increasing ammonium concentrations and by a reduction of biomass in Arnica and Cirsium. Furthermore, cation concentrations in both roots and shoots decreased when ammonium was supplied as a nitrogen source. In contrast, Calluna showed optimal development when using ammonium as a sole nitrogen source. In this species, only root biomass was negatively affected by high ammonium concentrations. The ecological implications of these preferences are discussed in relation to soil acidification.     

Effects of light and nutrient availability on the growth,allocation, carbon/nitrogen balance,phenolic chemistry,and resistance to herbivory of two freshwater macrophytes

Phenotypic responses of Potamogeton amplifolius and Nuphar advena to different light (7% and 35% of surface irradiance) and nutrient environments were assessed with field manipulation experiments. Higher light and nutrient availability enhanced the growth of P. amplifolius by 154% and 255%, respectively. Additionally, biomass was allocated differently depending on the resource: high light availability resulted in a higher root/shoot ratio, whereas high nutrient availability resulted in a lower root/shoot ratio. Low light availability and high nutrient availability increased the nitrogen content of leaf tissue by 53% and 40% respectively, resulting in a 37% and 31% decrease in the C/N ratio. Root nitrogen content was also increased by low light and high nutrient availability, by 50% (P=0.0807) and 77% respectively, resulting in a 20% and 40% decrease in root C/N ratio. Leaf phenolics were significantly increased 72% by high light and 31% by high nutrient availability, but root phenolic concentrations were not altered significantly. None of these changes in tissue constituents resulted in altered palatability to crayfish. N. advena was killed by the same high nutrient treatment that stimulated growth in P. amplifolius, preventing assessment of phenotypic responses to nutrient availability. However, high light availability increased overall growth by 24%, but this was mainly due to increased growth of the rhizome (increased 100%), resulting in a higher root/shoot ratio. High light tended to increase the production of floating leaves (P=0.09) and significantly decreased the production of submersed leaves. High light availability decreased the nitrogen content by 15% and 25% and increased the phenolic concentration by 88% and 255% in floating and submersed leaves, respectively. These differences in leaf traits did not result in detectable differences in damage by herbivores.     

Lignin signatures show the effects of changes in heather and bracken cover on the composition of organic matter in a moorland soil profile

Bracken (Pteridium aquilinum) is aggressively displacing heather (Calluna vulgaris) on many moorlands in Britain. We investigated the use of lignin derivatives to identify the distribution of soil organic matter (SOM) derived from bracken in moorland soil profiles formed under heather. Phenylpropanoids extracted from recently senesced litters, roots and SOM, using alkaline CuO oxidation, showed distinct signatures for bracken and heather, with vanillyl moieties dominating bracken litter extracts and vanillyl and syringyl dominating heather litter extracts. Ratios of vanillyl and syringyl concentrations characterised the SOM derived from heather and bracken better than the concentrations of the individual moieties. The analysis showed up to a depth of 5 cm under pure bracken cover, and at the interface between heather and bracken, the SOM was largely derived from bracken litter but below that depth SOM was apparently derived from heather. The use of these methods to identify the plant origin of SOM not only enables understanding the effects of changing vegetation cover on organic matter dynamics in moorland soils but could also facilitate management techniques in moorland/heathland restoration which involve the removal of comparatively nutrient-rich SOM derived from bracken. Received: 30 November 1997 / Accepted: 11 April 1998     

Effects of enhanced UVB on populations of the phloem feeding insect Strophingia ericae (Homoptera: Psylloidea) on heather (Calluna vulgaris)

Heather plants (Calluna vulgaris (L.) Hull) were grown for two years (April 1994–October 1996) under ambient or enhanced ultraviolet-B radiation (UVB: 280–315 nm), provided as a modulated treatment simulating a 15% ozone depletion (seasonally adjusted). Populations of the psyllid (Strophingia ericae (Curtis)) were measured before treatment and at yearly intervals thereafter. Before treatment there was no significant difference in the psyllid populations between treatments, or between the experimental and source populations. Enhanced UVB progressively produced a reduction in S. ericae populations compared with controls over 27 months. Analyses of C, N, total water soluble phenolics, total free amino acids and measurements of leaf angles and distances between leaflets demonstrated no effects of UVB treatment. However, concentrations of the amino acid isoleucine were lower (28%) in C. vulgaris exposed to the enhanced UVB treatment. Over the duration of the experiment the psyllid population structure at Lancaster changed from that typical of the upland site of origin (two-year cycle with overlapping cohorts) to a one-year life cycle typical of lowland sites, but this was independent of UVB treatments. Reduced isoleucine might explain the negative effects of elevated UVB on psyllid population numbers, but the precise effects of UVB on host chemistry and morphology are unknown. The problem of interpreting herbivore responses to enhanced UVB treatments in the field is discussed.     

Effects of nutrient and CO2 availability on tolerance to herbivory in Brassica rapa

The ability of plants to recover from herbivore damage and maintain their fitness depends on physiological mechanisms that are affected by the availability of resources such as carbon and soil nutrients. In this study, we explored the effects of increased carbon and nutrient availability on the response of rapid cycling Brassica rapa to damage by the generalist herbivore, Trichoplusia ni (Noctuidae), in a greenhouse experiment. Using fruit mass as an estimate of plant fitness, we tested three physiological models, which predict either an increase or a decrease of tolerance to herbivory with increasing resource availability. We used leaf demography to examine some plausible mechanisms through which resource availability may affect tolerance. Our results contradict all models, and, rather, they support a more complicated view of the plasticity of resource uptake and allocation than the ones considered by the models tested. Fruit mass was negatively affected by herbivore damage only under elevated CO₂, and only for certain harvest dates. Increased CO₂ had no effect on the number of leaf births, but it decreased leaf longevity and the total number of leaves on a plant. Nutrient addition increased the number of leaf births, leaf longevity and the total number of leaves on a plant. We conclude that a shortening of the life span of the plants, brought about by elevated CO_2, was responsible for a higher susceptibility of plants to herbivore damage under high CO₂ concentration.     

Effects of habitat fragmentation and soil quality on reproduction in two heathland Genista species

Habitat fragmentation decreases plant population size and increases population isolation, as well as altering patterns of plant–animal interactions, all of which may reduce plant fitness. Here, we studied effects of habitat fragmentation (in terms of population size and isolation) and soil quality on the reproduction of two rare legume species, Genista anglica (13 populations) and Genista pilosa (14 populations), confined to remnants of acidic and nutrient‐poor Calluna heathlands. Single individuals of the Genista plants are impossible to distinguish; population size was therefore estimated according to the area occupied (referred to as population size hereafter). We collected soil samples in all heathland sites to determine content of soil water, C, N, P, Ca, K and Mg. In both species values of soil pH and C/N ratio, as well as content of soil P and base cations, reflected the highly acidic and nutrient‐poor environment of the heathlands. Population sizes were unrelated to soil quality. Although the two Genista species are similar in morphology and ecology, effects of explanatory variables on reproduction were largely inconsistent across species. In G. anglica, population size had a positive impact on all reproductive variables except germination rate, which, in contrast, was the only variable affected positively by population size in G. pilosa. In both species, mean total reproductive output, calculated as the product of total seed mass per shoot and total germination, increased with increasing water content and decreased with increasing P. In G. anglica, we found positive effects of the C/N ratio on all reproductive variables except mean single and total seed mass per shoot. In summary, in both species reproductive success per shoot decreased with increasing soil nutrient availability in the heathland sites. The infestation of two large populations of G. pilosa with the pre‐dispersal, seed‐predating weevil Apion compactum had no significant effect on reproduction of the populations.     

Influence of nutrient availability on the mechanisms of tolerance to herbivory in an annual grass, Avena barbata (Poaceae)

Tolerance, or the capacity of a genotype to survive and reproduce following herbivore damage, varies widely across the plant kingdom. One proximate cause of this variation is resource availability, which can influence tolerance through mechanisms such as growth rate and photosynthesis. We examined the effect of high and low soil nutrient levels on the relationship between tolerance and two of its underlying mechanisms, biomass regrowth and photosynthetic upregulation, among genotypes of the Mediterranean annual grass Avena barbata. Although defoliated plants did not reach the same biomass as controls, biomass regrowth was higher at high nutrients. However, increased seed abortion at high nutrients caused tolerance to be the same in both nutrient treatments. Increased seed abortion also uncoupled biomass regrowth from tolerance at high nutrients. We found no evidence for photosynthetic upregulation in defoliated compared to control plants in either nutrient treatment. However, tolerance was positively correlated with predefoliation photosynthetic efficiency at high nutrients. Thus, constitutive photosynthetic efficiency may be a better predictor of tolerance than photosynthetic responses following herbivory in A. barbata. More generally, our results highlight the possibility that the mechanisms of tolerance can differ across resource environments even if tolerance is the same.     

Effects of light availability on growth, architecture and nutrient content of the seagrass Zostera noltii Hornem

The growth vs. irradiance response of the seagrass Zostera noltii from Cadiz Bay Natural Park (southwestern Spain) was characterised. Plants were exposed along 14 days to different light treatments (1%, 7%, 42% and 100% surface irradiance, SI), using shade screens in an outdoor mesocosm. Growth at 100% SI (1.6 mg DW plant⁻¹ day⁻¹) was lower than that at 42% SI (2.4 mg DW plant⁻¹ day⁻¹), suggesting photoinhibition. The minimum light requirement estimated was 0.8 mol photons m⁻² day⁻¹ (2% SI). Light availability affected the pattern of plant development and the overall plant growth. The contribution of the apical shoots to the aboveground production was nearly constant (c.a. 1.13 cm plant⁻¹ day⁻¹) regardless of the light level (except at 1% SI). In contrast, recruitment and growth of lateral shoots arising from the main rhizome axes accounted for the observed differences in aboveground growth. Rhizome branching was only observed at 42% SI. The possibility of a light threshold for rhizome branching could explain the seasonality of shoot recruitment, as well as the observed decrease in shoot density along depth (or light) gradients in seagrass meadows. Carbon demands at low irradiances (1% and 7% SI) were partially met by mobilization of carbohydrate reserves (sucrose in belowground and starch in aboveground parts). Plant nitrogen content decreased with increasing light, especially in belowground parts, reaching critical levels for growth.     

Coping with low nutrient availability and inundation: root growth responses of three halophytic grass species from different elevations along a flooding gradient

We describe the responses of three halophytic grass species that dominate the low (Spartina anglica), middle (Puccinellia maritima) and high (Elymus pycnanthus) parts of a salt marsh, to soil conditions that are believed to favour contrasting root-growth strategies. Our hypotheses were: (1) individual lateral root length is enhanced by N limitations in the soil but restricted by oxygen limitations, (2) the density of root branching within a species is inversely related to the length of the lateral roots, and (3) species from high elevations (i.e. the driest parts of a marsh) are the most responsive to changing soil conditions. Plant growth responses and soil parameters showed that the contrasting but uniformly applied soil treatments were effective. All three species showed a small but significant shift towards a finer root diameter distribution when N was limiting, partly because of the finer diameters of the laterals (Elymus and Spartina) and partly because of increased length of individual 1st-order laterals (Elymus and Puccinellia). The increased length of the 1st-order laterals of Elymus and Puccinellia grown under low N indicates that the first part of hypothesis 1 may be true. However, lack of effect of flooding and reduced soil conditions lead us to reject the second part of hypothesis 1. Hypothesis 2 was rejected for these three halophytes, as the branch density of 1st- and 2nd-order laterals appears to be controlled by other factors than length of individual laterals. Hypothesis 3 may be true for specific root characteristics (e.g. length of individual 1st-order laterals), but cannot be generalised (e.g. branch density and topological index). In conclusion, the present data on root growth in contrasting but homogeneous soil conditions indicate that morphological responsiveness of the root systems of these halophytic grass species is limited, regardless of their location along the elevational gradient.     

Isolation and characterization of microsatellite loci from Lochmaea suturalis, the heather beetle (Coleoptera: Chrysomelidae), a pest in Europe and a biocontrol agent in New Zealand

The heather beetle Lochmaea suturalis which is native to northwest Europe has been released as a biocontrol agent for heather in New Zealand. We have isolated and optimized eight microsatellite loci from New Zealand beetles. These loci provide markers with high polymorphism ranging from four to 20 alleles per locus. Observed heterozygosity averaged 0.631 per locus. These results suggest the markers are useful for population studies that will contribute to assessment of L. suturalis as a biocontrol agent.