The seedbanks of the Breckland heaths and heath grasslands, eastern England, and their relationship to the vegetation and the effects of management

The vegetation and the associated germinable seedbank of a range of heath and grass heath communities in Breckland, eastern England, were compared. The composition of vegetation and seedbank were similar for heather-dominated communities due to the predominance of Calluna vulgaris . Recent rotovation of previously heather-dominated communities invaded by Deschampsia flexuosa reduced the number of emerging C. vulgaris seedlings by more than 80%. The composition of seedbanks of calcicolous grasslands differed considerably from the vegetation, with a significant contribution from many ruderal species not present in the vegetation and an absence of important species such as Pilosella officinarum. Acidic grasslands were intermediate between calcicolous grasslands and heather-dominated heathlands, with seedbanks characterized by high densities of Rumex acetosella and a number of ruderal species that were absent or rare in the vegetation. The composition of the seedbank of more mesotrophic grasslands suggests that the imposition of some form of disturbance may recreate more open grasslands typical of Breckland.     

Differential Indirect Effects of Excluding Livestock and Rabbits from Chalk Heath on the Associated Leafhopper (Hemiptera: Auchenorrhyncha) Fauna

Preliminary results are presented of sampling the leafhopper assemblages on a field experiment designed to examine the differential effects of rabbits and livestock (mainly sheep) on the vegetation of chalk heath in southern England. Experimental plots that excluded livestock either allowed entry by rabbits or excluded them. Results were compared with those from plots grazed by both livestock and rabbits. After 7 years, exclusion of grazing herbivores had resulted in predictable increases in vegetation height, but no major changes were detected in the species composition of the vegetation. As expected, ungrazed plots had higher species richness and greater abundances of several individual leafhopper species. However, plots grazed only by rabbits had a leafhopper assemblage that was distinct from either ungrazed or mixed grazing plots. It is suggested that rabbit grazing may have subtle effects on grassland invertebrate assemblages that are not necessarily predictable from an examination of the species composition of the vegetation. Chalk heath vegetation contains an unusual mixture of calcicole and calcifuge plant species, but the leafhopper assemblage included a restricted number of calcareous grassland specialist species and only one species strongly associated with acidic grasslands; most leafhoppers recorded were generalist grassland species.     

Experimental wet heath translocation in Dorset, England

Heathland has become fragmented throughout its European range with many countries losing 60-90% of their heathland area during the 20th century. A small, isolated area of wet heathland within permitted ball clay workings in Dorset (England) was translocated in October 1993 to an engineered receptor cell fed by groundwater and direct precipitation in order to mitigate the otherwise total loss of the wet heath. The aims were to rescue the wet heathland and to re-establish a community type similar to the wet heath that originally occurred at the donor site prior to the recent development of vegetation communities and birch colonisation associated with drier conditions. Vegetation monitoring over 12 years up to 2005 indicated that the experimental translocation successfully fulfilled the aims for a period of 7 years when the hydrological regime of the receptor site was actively managed. There was development of a mosaic of wet heath and mire vegetation community types including Sphagnum lawns and open bog pools. However, when active management ceased, the receptor site became drier and species were lost. Multivariate analysis revealed a trajectory of progressive rapid chronological change during 1994 and 1995 followed by a period of relative stability between 1996 and 2000 which coincided with greater species density. Subsequently up to 2005, the plant community became less species-rich and the plant community reverted to a species structure similar to that which occurred in 1994/1995. The importance of hydrological regime for plant community structure was tested using fixed-quadrat data and the water level measurements made in the period 1995-1998. Canonical Correspondence Analysis (CCA) multivariate analysis revealed a hydrological gradient with a significant seasonal effect of water table depth in the first half of the year and close association of the least wet quadrat location and non-mire species. Overall, the results support the use of translocation as a means of rescuing wet heath habitat which would otherwise be lost due to change in land use. There is a risk of failure when translocating a complex ecosystem such as wet heath but ecological engineering provides an insight into defining the risk. Our results demonstrated that an ecologically acceptable outcome for a wet heath translocation can be achieved with careful design and implementation and sensitive management of the receptor site. However, it is essential to secure long-term hydrological and vegetation management in order to provide a sustainable outcome for long-term survival of a target wet heath community.     

The contribution of grazing to plant diversity in alpine grassland and heath

Abstract Grazing on transplants of a grass, a forb and a tree was examined in low-diversity grassland and more diverse heath in Australia's Snowy Mountains. Transplants were surrounded by 2 mm mesh netting. In one grassland plot, grazers (probably soil invertebrates) attacked 40–90% of tree and forb seedlings but no grass seedlings. In heath, which had about half the grass cover of grassland, grazers consumed grasses but not trees or forbs. The results suggest that grazers can depress diversity in grassland by attacking species other than grass. In heath, they may promote diversity by attacking only grass and releasing other species from competition.     

Effects of fire regime on plant species richness and composition differ among forest,woodland and heath vegetation

Question

Do the effects of fire regimes on plant species richness and composition differ among floristically similar vegetation types?

Location

Booderee National Park, south‐eastern Australia.

Methods

We completed floristic surveys of 87 sites in Sydney Coastal dry sclerophyll vegetation, where fire history records have been maintained for over 55 years. We tested for associations between different aspects of the recent fire history and plant species richness and composition, and whether these relationships were consistent among structurally defined forest, woodland and heath vegetation types.

Results

The relationship between fire regime variables and plant species richness and composition differed among vegetation types, despite the three vegetation types having similar species pools. Fire frequency was positively related to species richness in woodland, negatively related to species richness in heath, and unrelated to species richness in forest. These different relationships were explained by differences in the associations between fire history and species traits among vegetation types. The negative relationship between fire frequency and species richness in heath vegetation was underpinned by reduced occurrence of resprouting species at high fire frequency sites (more than four fires in 55 years). However, in forest and woodland vegetation, resprouting species were not negatively associated with fire frequency.

Conclusions

We hypothesize that differing relationships among vegetation types were underpinned by differences in fire behaviour, and/or biotic and abiotic conditions, leading to differences in plant species mortality and post‐fire recovery among vegetation types. Our findings suggest that even when there is a high proportion of shared species between vegetation types, fires can have very different effects on vegetation communities, depending on the structural vegetation type. Both research and management of fire regimes may therefore benefit from considering vegetation types as separate management units.     

Influence of soil and microclimate on species composition and grass encroachment in heath succession

Aims and Methods Mostly due to land use changes, European heathlands have become increasingly rare. In addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses and a loss in species diversity. Despite many investigations, information about the precise environmental parameters that determine the development and maintenance of heathland vegetation is still insufficient. In order to determine the environmental factors that control heath succession and grass encroachment, and to develop appropriate management schemes, we studied the influence of several soil and microclimate parameters on species composition and vegetation characteristics in five successional stages in a coastal heathland on the island of Hiddensee, north-east Germany, where the encroachment of Carex arenaria has become a major problem.Important findings We recorded the highest plant species richness in grey dune and birch forest plots, while the encroachment of C. arenaria let to a significant decline in plant species richness. The most important environmental factors influencing species richness and distribution of single species were microclimate, soil moisture, soil pH and the C/N ratio. While many studies reported the importance of differences in nutrient availability, we found no significant correlations between soil nutrient availability and vegetation pattern. Environmental conditions in dense C. arenaria stands, especially soil properties (e.g. soil pH), showed great differences in comparison to the other successional stages. However, no correlations between the encroachment of C. arenaria and single environmental factors were found. Our results show that not only soil nutrients are important abiotic factors in heaths but that also microclimate and soil moisture play an important role and that many factors are involved in heath succession and in the promotion of grass encroachment. Management plans for the conservation and restoration of heathlands should therefore focus on the specific site conditions and should take several abiotic and biotic factors into account.     

Heathland dynamics in biotically disturbed areas: on the role of some features enhancing heath success

It is well accepted that intense anthropogenic activities threaten European heathland communities. The negative impact of high intensities of livestock grazing on heath survival has been demonstrated in many studies, however, it does not seem a straightforward consequence in some cases: in the southern Pyrenees, the existence of large heathland patches in intensively grazed areas suggests that other factors interact and affect heath success. In order to gain insight into the factors affecting heath success, this paper analyses the pattern of occurrence of ericaceous shrubs—particularly Erica vagans—and their effects on soils when compared to herbaceous vegetation, under a complex biotic disturbance regime. Survey plots were established in grassland and heathland communities disturbed by both livestock and by burrowing animals on soils with different levels of basic resources. Relative cover of the species on and off molehills was studied. In heathland plots, ericaceous shrubs were more common on old molehills than off them. This pattern was strongest in the poorest soils. Below established heath shrubs, independently of the initial properties of the soils, some basic resources were more available than in intershrub areas. In grazed ecosystems, where continuous trampling and removal of biomass occurs, the heath canopy provides a site for accumulation of litter and insects, which contribute to the build up of a resource-rich layer. We conclude that successful establishment on ubiquitous molehills and suitable, localised changes in the soil under shrubs are some of the factors that explain the success of ericaceous shrubs in complex biotically disturbed areas.     

不同放牧强度下的内蒙古草原蝗虫群落动态

本文研究了内蒙古草原蝗虫群落在不同放牧强度下的动态变化。植物和蝗虫的取样涉及3种植物群落类型。每一种群落类型又根据放牧强度的不同,划分为5个放牧梯度。随着放牧强度的增加,蝗虫群落的组成和结构均发生了明显的变化。使用贴近度分析及择近原则的方法,阐明了蝗虫种的优势度变化及替代种的替代次序。通过灰色关联及系统聚类分析方法的结合,分析了11个蝗虫种对不同放牧强度的反应。对应性分析和多元逐步回归分析说明,蝗虫群落的变化是与某些植被因子和土壤因子密切相关,特别是植物种类数、生物量、高度以及土壤硬度和含水量等。某些蝗虫种的生态指示意义也在文中进行了讨论。     

The age of Calluna stands moderates post‐fire regeneration rate and trends in northern Calluna heathlands

Questions: Does stand age influence the direction and rate of post‐fire successional dynamics in coastal Calluna heaths and can old degraded heath vegetation be restored through reintroduction of fire? Location: Coastal heaths in the Tarva archipelago, central Norway. Methods: We investigated revegetation dynamics after experimental fires set in young (8 years since last fire) and old (>50 years since last fire) grazed heath stands. A repeated measures design was used, with floristic data recorded in permanent plots in the post‐fire successions (n=12) over a 7‐year period. The data were analysed using multivariate ordination techniques (PCA, RDA and PRC) and mixed effects models. Results: The age of Calluna stands strongly influenced post‐fire succession, different trends due to age explained 10.4% of variation in floristic data. Young heath showed faster succession towards pre‐fire community composition than old heath, and this could partially be explained by succession‐related factors: young heath had lower cover of mosses and lichens in the pre‐burned vegetation, and lower cover of litter early in succession. Young heath had a less pronounced overall community response to fire than old heath. Vegetative regeneration of C. vulgaris was absent in both old and young heath, but Calluna still re‐established as the dominant species within 5–7 years in both young and old stands. Regeneration dynamics were also affected by habitat conditions, different trends due to habitat explained 6% of variation. Conclusions: Our study demonstrates that old stands do develop characteristic heathland vegetation and structure after fire, and while potential invasives into the system such as trees and rhizomatous species are present, they do not impair Calluna regeneration or vegetation development towards the target heathland community composition and structure. Further, as our young stands are only in their second fire rotation after restoration, we suggest that characteristic dynamics of managed heathlands can re‐establish relatively rapidly, even in severely degenerated sites (>50 years since last fire). Site‐specific factors also need to be considered. We conclude that there is restoration potential in old heaths, despite slow dynamics in the first rotation.     

A hierarchical model for analysing the stability of vegetation patterns created by grazing in temperate pastures

Questions: Does vegetation structure display any stability over the grazing season and in two successive years, and is there any correlation between the stability of these spatial patterns and local sward composition? Location: An upland grassland in the French Massif Central. Method: The mosaic of short and tall vegetation stands considered as grazed and ungrazed patches respectively is modeled as the realization of a Boolean process. This method does not require any arbitrarily set sward‐height thresholds to discriminate between grazed and ungrazed areas, or the use of additional variables such as defoliation indexes. The model was validated by comparing empirical and simulated sward‐height distributions and semi‐variograms. Results: The model discriminated between grazed and ungrazed patches at both a fine (1 m²) and a larger (500 m²) scale. Selective grazing on legumes and forbs and avoidance of reproductive grass could partly explain the stability of fine‐scale grazing patterns in lightly grazed plots. In these plots, the model revealed an inter‐annual stability of large‐scale grazing patterns at the time peak biomass occurred. At the end of the grazing season, lightly grazed plots showed fluctuating patch boundaries while heavily grazed plots showed a certain degree of patch stability. Conclusion: The model presented here reveals that selective grazing at the bite scale could lead to the creation of relatively stable patches within the pasture. Locally maintaining short cover heights would result in divergent within‐plot vegetation dynamics, and thus favor the functional diversity of vegetation.     

Fire,aridity and seed banks. What does seed bank composition reveal about community processes in fire‐prone desert?

Questions: The relationship between fire, aridity and seed banks is poorly understood in plant community ecology. We tested whether there was a close correspondence between the seed bank and standing vegetation composition with time‐since‐fire in a desert. We also examined whether longer‐lived species showed seed limitation relative to more ephemeral species, as this could influence grass‐woody ratios in a major biome. Location: Dune hummock grasslands/shrublands of central Australia. Methods: The effects of time‐since‐fire on floristic and functional group composition were examined by comparing plots unburned since 1984 against plots that had been burned in 2002. Three methods were used to quantify seed abundances: a germination trial using heat and smoke application, a flotation method, and a sieving method. Results: Seed bank densities were very low (<3000 m^?2). Species similarity between the seed bank and standing vegetation was high at sites recently burned (0.86) and low in sites long‐since burned (0.52). The relative abundance of ephemeral species in the seed bank peaked in recently burned plots, but the relative abundance of seeds of woody species did not match the pattern of abundance in the standing vegetation. Remarkably, the dominant perennial grasses and woody species were either absent from the seed bank or present at extremely low abundances. Discussion: Differences in the relative abundance of ephemeral species between standing vegetation and seed bank relate to the post‐fire succession process. The small soil pool of seed from woody species may be explained by allocation to belowground carbohydrate storage over seed production. Field observations suggest, however, that production of strongly dormant seed can be prolific and that high levels of seed predation make this system strongly seed‐limited. The discovery of this seed bank syndrome indicates that shifts in grass‐woody ratios can be driven by the juxtaposition of unpredictable seed rain and fire events in these desert dunes. However, estimates of grass‐woody ratios due to changing fire regimes will be difficult to predict.     

Response of the moss Racomitrium lanuginosum to changes in sheep grazing and snow‐lie due to a snow‐fence

Question: What are the responses of Racomitrium lanuginosum moss to altered snow‐lie and sheep use? Location: ACarexbigelowii‐Racomitrium lanuginosum heath on a Scottish montane plateau affected since 1986 by a fenced ski corridor. Methods: Permanent quadrats were set up along transects 45 m long perpendicular to the snow‐fence. Cover was assessed over a 12‐year period from 1990. Pellet‐group clearance counts provided data on sheep usage between 1990 and 1996. Snow‐lie was mapped in the springs of 1991–1996. Results: The snow‐fence created a gradient in sheep use and altered the duration of snow‐lie. At the start of monitoring Racomitrium cover was lower immediately adjacent to the fence, and after 12 years its cover was significantly reduced within 10 m of the fence. Further away from the fence Racomitrium cover was relatively stable. The loss of Racomitrium was correlated both with increased snow‐lie and heavier sheep usage. Grass cover increased near the fence and was related to sheep use. Dicranum fuscescens responded differently to Racomitrium, increasing significantly near the fence. Conclusions: We found that changes in snow‐lie and grazing pressure quickly brought about vegetation change in this montane ecosystem. Racomitrium was the most sensitive species to the changes in grazing and snow‐lie caused by the fence, having the biggest initial changes. Loss of Racomitrium permitted increases of species more resistant to grazing including Dicranum fuscescens and grasses.     

DYNAMICS OF GRASSHOPPER COMMUNITIES UNDER DIFFERENT GRAZING INTENSITIES IN INNER MONGOLIAN STEPPES

Abstract Vegetation and grasshopper community variables were monitored on the natural steppes grazed by livestock in Inner Mongolia, China. The sites sampled belong to 3 vegetation types laid separately in 3 continuums from different grazing intensities. Each continuum consisted of 5 plots along grazing gradients. Within the plots, the species composition, population number and population calorific value of grasshoppers that was converted from their biomass varied sensitively with increase of grazing intensities. The methods of neartude and choosing rules of fuzzy mathematics were used to analyze successive populations of grasshoppers. By combining the gray relation with hierarchic cluster analysis, eleven grasshopper species were divided into 3 grazing guilds based on their responses to grazing. The results from detrended correspondence and the multivariate regression analysis proved that there were close successive relationships among the grasshopper species that varied with the changes of plant species, biomass, height and coverage. The importance of some grasshopper indicators to grassland change is discussed.     

Recovery of subalpine dwarf shrub heath after neighbour removal and fertilization

We tested if subalpine heath vegetation in northern Italy recovered after experimental perturbation of soil nutrient availability (fertilization) and species composition (removal of co-dominant dwarf shrubs). Species cover was assessed non-destructively before the start of the treatments (1995), at the end of the treatments (1999) and 4 years after the treatments ended (2003). Shrub removal had rather modest effects on heath vegetation, except for mosses which decreased significantly in removal plots. Fertilization decreased the cover of shrubs, mosses, and some graminoids but increased the cover of Festuca rubra. Fertilization converted heath to grassland, but the response of graminoid species was individualistic. Fertilization decreased vascular species richness and evenness, probably through negative effects of shading and litter accumulation on plant growth or recruitment. The vegetation had not recovered completely 4 years after the perturbations had stopped. This suggests that, in contrast to rapid responses to species removal and fertilization, recovery from these perturbations was rather slow, presumably because recovery was affected by long-term biotic interactions and species controls on ecosystem properties.     

Relationships between vegetation and bird community composition in grasslands of the Serengeti

The Serengeti–Mara ecosystem holds one of the largest natural grasslands of the world, which is well known for its large herds of mammals. However, the bird community structure of these grasslands has hardly ever been studied. For the first time, a large‐scale study on grassland bird communities has been conducted in all typical open grasslands of the Serengeti National Park. We used ten grassland plots representing a gradient of increasing vegetation height and also including shrubs and trees to analyse the influence of vegetation structure on the composition of grassland bird communities. Three communities of breeding birds were identified in relation to (a) short grass, (b) intermediate and long grass and (c) wooded grasslands. The bird communities of intermediate, long and wooded grassland were very similar, because of identical dominant bird species. Our results suggest that breeding birds of East African grasslands exhibit two contrasting habitat relationships: (1) birds that are restricted to short grass, and (2) species that are tolerant to vegetational shifts from intermediate grass via long grass to early stages of woody vegetation. Because the vegetation is often driven beyond this range in managed tropical grasslands through high grazing pressure or through shrub encroachment, we expect that the long‐grass bird communities will not generally resist anthropogenic disturbance.     

Seedling emergence after sod cutting in grass heath

Abstract. Experimental plots in grass heath, previously subjected to various sod cutting treatments, contained a seed store of ericoid dwarf shrubs, but seeds of grasses of the former grass heath were not found. The density of emerging dwarf-shrub seedlings in the plots was determined mainly by soil moisture, but with density of seeds stored in the soil and soil acidity as modifying factors. It is concluded that the restoration of a dwarf-shrub heathby sod cutting is relatively successful, as compared to other measures, because the grass seeds become obsolete.     

Effects of livestock breed and grazing pressure on ground‐dwelling arthropods in Cantabrian heathlands

Abstract 1. The effects of different goat breeds and grazing pressures on epigeal arachnids (Lycosidae and Opiliones) and ground beetles (Carabidae) were assessed in a heath–gorse shrubland located in the north of the Iberian Peninsula. Three treatments (low grazing pressure with Cashmere breed, high grazing pressure with Cashmere breed, and high grazing pressure with local Celtiberic breed) with three replicates of each treatment were allocated randomly to nine plots. 2. Arthropod fauna abundance, diversity, and species richness were estimated using pitfall trapping over 3 years (2003, 2004, and 2005). Vegetation cover, composition, and height were assessed along transects and above‐ground plant biomass was sampled in quadrats. 3. Lower grazing pressure led to lower herbaceous cover and higher heather cover than high grazing pressure. Overall, it did not enhance arthropod diversity or abundance, but the abundance and species richness of carabids, and abundance of lycosids like Pardosa pullata, were higher under high stocking rates, whereas Opiliones were favoured by the low stocking rates. Although it has been assumed that traditional breeds enhance biodiversity, the present study found no significant differences for both arthropod fauna and vegetation between the goat breeds. 4. Arthropod–vegetation relationships were examined by constrained ordination. Fauna assemblages followed a gradient from closed canopy shrubland areas, in the lowest grazing pressure, to grassier open canopy areas resulting from higher grazing pressure, revealing the importance of grazing management as a driver of arthropod community structure of the heathland.     

Predicted changes in vegetation structure affect the susceptibility to invasion of bryophyte-dominated subarctic heath

Background and Aims

A meta-analysis of global change experiments in arctic tundra sites suggests that plant productivity and the cover of shrubs, grasses and dead plant material (i.e. litter) will increase and the cover of bryophytes will decrease in response to higher air temperatures. However, little is known about which effects these changes in vegetation structure will have on seedling recruitment of species and invasibility of arctic ecosystems.

Methods

A field experiment was done in a bryophyte-dominated, species-rich subarctic heath by manipulating the cover of bryophytes and litter in a factorial design. Three phases of seedling recruitment (seedling emergence, summer seedling survival, first-year recruitment) of the grass Anthoxanthum alpinum and the shrub Betula nana were analysed after they were sown into the experimental plots.

Key Results

Bryophyte and litter removal significantly increased seedling emergence of both species but the effects of manipulations of vegetation structure varied strongly for the later phases of recruitment. Summer survival and first-year recruitment were significantly higher in Anthoxanthum. Although bryophyte removal generally increased summer survival and recruitment, seedlings of Betula showed high mortality in early August on plots where bryophytes had been removed.

Conclusions

Large species-specific variation and significant effects of experimental manipulations on seedling recruitment suggest that changes in vegetation structure as a consequence of global warming will affect the abundance of grasses and shrubs, the species composition and the susceptibility to invasion of subarctic heath vegetation.     

Influence of Grazing on Soil Seed Banks Determines the Restoration Potential of Aboveground Vegetation in a Semi‐arid Savanna of Ethiopia

Species composition, number of emerging seedlings, species diversity and functional group of the soil seed banks, and the influence of grazing on the similarity between the soil seed banks and aboveground vegetation, were studied in 2008 and 2009 in a semi‐arid savanna of Ethiopia. We tested whether the availability of persistent seeds in the soil could drive the transition from a degraded system under heavy grazing to healthy vegetation with ample perennial grasses. A total of 77 species emerged from the soil seed bank samples: 21 annual grasses, 12 perennial grasses, 4 herbaceous legumes, 39 forbs, and 1 woody species. Perennial grass species dominated the lightly grazed sites, whereas the heavily grazed sites were dominated by annual forbs. Heavy grazing reduced the number of seeds that can germinate in the seed bank. Species richness in the seed bank was, however, not affected by grazing. With increasing soil depth, the seed density and its species richness declined. There was a higher similarity in species composition between the soil seed bank and aboveground vegetation at the lightly grazed sites compared with the heavily grazed sites. The mean similarity between the seed banks and aboveground vegetation was relatively low, indicating the effect of heavy grazing. Moreover, seeds of perennial grasses were less abundant in the soil seed banks under heavy grazing. We concluded that restoration of grass and woody species from the soil seed banks in the heavily grazed areas could not be successful in semi‐arid savannas of Ethiopia.