Xylem acoustic emissions and water relations of Calluna vulgaris L. at two climatological regions of Britain

The acoustic emission rate (as a result of xylem cavitation), stem water content, stomatal conductance, and leaf water potential of heather (Calluna vulgaris L. Hull) were compared over a period of 18 months at two contrasting sites in Britain. The sites were Thursely Common in south-east England, and Flanders Moss in central Scotland (average rainfall of 600 mm per year and 1500 mm per year, respectively). In the first year of study (1992) the natural difference between the sites was amplified by a severe drought affecting south-east England. The relative water content of the xylem (RWC) did not differ between the sites, and did not fluctuate seasonally, despite the drought at Thursely. Acoustic emissions were detected at both sites, but the incidence was low. The same range of foliage water potential and stomatal conductance was found at both sites. The leaf area to sapwood area ratio at the dry site was half that at the wet site, and it is suggested that this adjustment of the transpiring area relative to the water conducting area facilitates the close physiological state of the plants at the two sites. The possible occurrence of aerial refilling and freezing induced xylem cavitation is also discussed.     

Effects of fire on the vegetation of a lowland heathland in North-western Italy

This study focuses on the effect of fire on lowland heathlands at the extreme southern edge of their European distribution (Vauda Nature Reserve, NW Italy). Forty-nine plots (50 m radius) were surveyed between 1999 and 2006. Each year, fire occurrences were recorded and per cent cover of four vegetation types (grassland, heath, low shrubland, and tall shrubland) was estimated in each plot. Vascular plant species richness was also recorded in 255, 1 m² quadrats. After a fire, grassland vegetation expanded, but then declined rapidly as heath and shrubland recovered: 7 years after a fire, tall shrubland encroached on to more than 40% of the plots, and grassland declined from 50% to 20% cover. Between 1999 and 2006, Betula pendula shrubland greatly expanded, while grassland decreased over most of the Reserve, even where fire frequency was high. Tall shrubland had low plant diversity and was dominated by widespread species of lower conservation value. By contrast, early successional vegetation (grassland and low shrubland) had higher richness and more narrowly distributed species, indication that the development of tall shrubland causes significant species loss in the heathland. Italian lowland heathlands are characterized by high rates of shrubland encroachment that threatens both habitat and species diversity. Burning frequencies of once in 3–6 years seem appropriate in this habitat, but burning alone might not suffice without actions to increase herbivore grazing.     

Changes in heathland vegetation under goat grazing: effects of breed and stocking rate

Questions: How are heathland vegetation dynamics affected by different goat grazing management? Location: Cantabrian heathlands in Illano, Asturias, northern Spain. Methods: During 4 years, vegetation dynamics (structural composition, canopy height and floristic diversity) were studied under three goat grazing treatments with three replicates: high stocking rate (11.7 goats ha^?1) with a local Celtiberic breed, and high (15 goats ha^?1) and low (6.7 goats ha^?1) stocking rates with a commercial Cashmere breed. Results: The relative cover of woody plants, particularly heather species, decreased more while herbaceous cover increased more under local Celtiberic than under Cashmere breed grazing. Within Cashmere treatments, the cover and height of live shrubs decreased more and the herbaceous cover increased more under high than under low stocking rate. Redundancy analysis showed a significant effect of treatment × year interaction on floristic composition. Greater species richness was recorded under local goat grazing, but Shannon diversity index fell in the fourth year on these plots because of dominance by two grass species. Conclusions: Local Celtiberic goat grazing at such a high stocking rate (11.7 goats ha^?1) hinders the development of sustainable systems on these heathlands, both in environmental and productive terms, owing to the limitations in soil fertility. Nevertheless, Celtiberic goats could be useful for controlling excessive shrub encroachment and reducing fire hazard. Cashmere goat grazing at high stocking rate promoted the highest Shannon diversity by generating a better balance between woody and herbaceous plants, while shrub dominance was not altered under the low stocking rate.     

Influence de la concentration de la solution nutritive dans la phase de réinsertionin vivo des vitroplants d'Erica × darleyensis

Résumé La réinsertion de vitroplants d'Erica × darleyensis est suivie en chambre climatisée et sur perlite arrosée de solutions nutritives, en analysant pendant deux mois le développement des plantes. Quatre solutions nutritives ont été testées. Seule en varie les concentrations (A, A/2, A/4 et A/10); le rapport entre les différents éléments reste constant. Les vitroplants issus d'un milieu de multiplication gélosé A/2, transplantés sur les milieux de réinsertion subisent un choc nutritionnel: même les vitroplants transférésin vivo sur solution dont la concentration en macroéléments est le double de celle du milieu de multiplicationin vitro, enregistrent une diminution des teneurs minérales. L'influence de la nature des substrats de ces différents milieux (gélose ou perlite) semble prédominante à cet égard. Des milieux de réinsertion moyennement concentrés (A/2 ou A/4) constituent les meilleures références pour la précocité et l'optimisation de la production des rameaux. La production des rameaux est reliée à certains paramètres internes nutritionnels tels que: rapports élevés K⁺/(Ca²⁺+Mg²⁺) et Mn²⁺/Fe³⁺, et de hautes teneurs en azote et phosphore.

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Influence ofin vivo nutrient concentration inErica × darleyensis vitroplant transfer on perlite mediaI. Development and nutrient contents of transferred vitroplants

Development ofErica × darleyensis vitroplants was studied on perlite nutritive media in growth chamber conditions, during two months. Four nutritive solutions were examined. Only solution concentrations vary (solutions A, A/2, A/4 and A/10); the ratio between nutrients remains constant. Vitroplants from a A/2 agar-agar propagation medium, transferred on perlite media underwent a nutritional stress: even vitroplants transferred on a perlite medium whose concentration is twice that of the propagation medium, show a decrease in mineral content. The influence of the kinds of substrates (agar-agar or perlite) seems to be prevailing in this respect. Perlite media irrigated with mildly concentrated solutions (A/2 or A/4) are the best references for earliness and optimization of ramification. Branch production is correlated to internal nutritional parameters such as: high K⁺/(Ca²⁺+Mg²⁺) and Mn²⁺/Fe³⁺), a high nitrogen and phosphorus contents.

     

Sustainable natural remediation of abandoned tailings by metal-excluding heather (Calluna vulgaris) and gorse (Ulex europaeus), Carnon Valley, Cornwall, UK

This study was conducted to determine the uptake of elements (Ag, Al, As, Ca, Cd, Co, Cu, Fe, Mn, Mo, Ni, Pb, P, S, Sb, Tl, U, W, Zn) by native gorse (Ulex europaeus) and heather (Calluna vulgaris), growing on abandoned tailings, Carnon Valley, Cornwall, UK. The metalliferous tailings are particularly As-rich (0.11-0.59 wt% As) and contain acid-generating sulfides (pH 3.36-6.59). Since abandonment three decades ago, gorse and heather have colonized much of the exposed tailings surface. Biogeochemical analyses demonstrate that gorse and heather are opportunistic, pioneering metallophytes that have the ability to exclude metals and As from their above-ground biomass. Concentrations of trace elements in gorse and heather were evaluated in terms of maximum tolerable levels in the feed of rodents and horses. The analyses revealed that gorse and heather do not accumulate large quantities of trace elements (Al, As, Cd, Co, Cu, Ni, Pb, Zn) in their tissue, preventing harmful effects on rabbits feeding on them and transfers of trace metals and As into the developing wildlife food chains. This study demonstrates that (a) biogeochemical examinations of abandoned mined lands can reveal pioneering, metal-excluding plants; and (b) abandonment and benevolent neglect of mined lands can lead to the successful development of sustainable vegetation covers over mine wastes in the long term.     

Effects of browsing and grazing on cyclic succession in nutrient‐limited ecosystems

Abstract. This paper deals with browsing and grazing as forces driving cyclic succession. Between 1989 and 1994 reciprocal transitions between the dwarf shrub Calluna vulgaris and the grass Deschampsia flexuosa were monitored in permanent plots in a cattle grazed grass‐rich Dutch heathland on podsolic soils in which tree encroachment was prevented. Heather beetles killed Calluna in four of the nine plots during 1991/1992. The monitoring revealed reciprocal transitions and cycles between Calluna and Deschampsia on a subplot scale. Beetles and cattle had additional and complementary effects on the two competing species. Defoliation by beetles and trampling by cattle‐killed Calluna and favoured grass invasion. Grazing and gap creation by cattle in Deschampsia favoured the establishment and recovery of Calluna. Analysis of the causal mechanisms suggests that indirect, resource‐mediated herbivory effects may be as important for the replacement processes as direct effects of defoliation and trampling. Herbivory created differential light and nutrient levels in Calluna and Deschampsia gaps. Grazing and browsing improved the resource‐capturing abilities of Calluna and its resistance to herbivory and abiotic disturbances. The emerged Calluna‐Deschampsia cycle and its driving forces are summarized in a conceptual triangular resource‐mediated successional grazing cycle (RSGC) model, a limit cycle involving herbivore‐plant‐plant resource interactions. It offers a deterministic equilibrium model as alternative for stochastic transitions between the meta‐stable states with dominance of Calluna and Deschampsia respectively. The validity range of the RSGC model and its management implications are briefly discussed.     

Altitudinal gradients in seed dynamics of Calluna vulgaris in eastern Scotland

Abstract. Relationships between seed deposition, size of soil seed banks and some of the environmental factors affecting them were assessed for Calluna vulgaris throughout its altitudinal range (150–960 m a.s.l.) in eastern Scotland. Seed rain was assessed using pitfall traps, collected every 5–10 wk for 3 yr; germinability was determined by laboratory incubation. Seed bank size was estimated, once only, by counting seedlings emerging from soil cores kept for 50 wk in a glasshouse. Seed deposition varied annually, was related to parent plant cover but always declined with altitude, falling sharply above 600 m a.s.l. Seed bank size was more closely correlated with the proportion of organic matter in the soil than with the amount of seed rain. Seed bank sizes declined gradually with altitude but did not differ significantly between four altitudinal zones. The mean density of buried seeds was less than half the mean annual seed rain at sites below 300 m a.s.l. but was over 200 × greater than annual seed rain above 800 m, suggesting that seeds buried at high altitudes remain viable for much longer than those at lower altitudes.     

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.     

The role of catastrophic death of Calluna in heathland dynamics

In the Breckland heaths of East Anglia large patches of Calluna are often killed by adverse climatic conditions or insect attack. This paper studies one site, Cavenham Heath, where large areas of Calluna were killed between 1976–1979, and shows that older stands were most affected, and recovery was slowest compared to either younger stands, or stands with an uneven-aged structure. Moreover, where Calluna death was most severe, directional succession to birch woodland was accelerated. At this site, although Calluna regeneration has been previously assumed to be initiated by endogenous factors (i.e. by the life cycle of the Calluna), it is also clearly interrupted by exogenous factors. The implications of these results for heathland conservation are discussed, and it is suggested that intervention management by cutting and burning, to maintain large areas of Calluna in the building phase, may be the most appropriate long term strategy.This work was funded in part by the Nature Conservancy Council as part of its research programme into nature conservation. I thank Mr D. Malins and Mrs D. Reynolds for assistance in both the field and the laboratory, Ms S. Ide and Mr J. Pattin-gale for preparing the figures, and Professor C. H. Gimigham, Dr J. Miles and Dr I. C. Prentice for helpful comments on earlier drafts of this paper.