Changes in landscape and vegetation of coastal dunes in northwest Europe: a review

In coastal dunes, landscape changes are a rule, rather than an exception. This paper gives an overview of changes in landscape and vegetation with a focus on the past century. The history of dunes is characterised by phases of sand drift, alternated with geomorphological stability. The historical development of dune woodland during these stable phases has been documented for sites all over Europe. Vegetation reconstructions of historical open dune habitats however is very difficult due to limited preservation of fossil remains. People have drastically altered coastal dune landscapes through centuries of exploitation and more recently development of the coast. Historical land use has generally pushed vegetation back into a semi-natural state. During roughly the past century a tendency of increasing fixation and succession is observed on coastal dunes throughout northwest Europe. Six causes of change are discussed. 1) Changes in land use, mainly abandonment of agricultural practices, have led to the development of late successional stages such as scrub and woodland. 2) Crashing rabbit populations due to myxomatosis in the 1950s caused vigorous grass growth and probably stimulated scrub development. 3) A general tendency of landscape fixation is observed due to both natural and anthropogenic factors. 4) Eutrophication, mainly due to atmospheric nitrogen deposition is clearly linked to grass encroachment on acidic but also on some calcareous dunes. 5) The impact of climate change on vegetation is still unclear but probably lengthening of growing season and maybe enhanced CO₂ concentrations have led to an acceleration of succession. 6) A general anthropogenisation of the landscape occurs with rapid spread of non-native species as an important consequence. The reconstruction of a natural reference landscape is considered largely unattainable because of irreversible changes and the long tradition of human impact, in many cases since the development of the dunes. Two contradictory elements need reconciliation. First, the general acceleration of succession and scrub and woodland development in particular is partly caused by a decreased anthropogenic interference in the landscape and deserves more appreciation. Second, most biodiversity values are largely linked to open, early succession dune habitats and are threatened by the same tendency. Apart from internal nature management, in which grazing plays an important part, re-mobilisation of stable, senescent dunes is an important challenge for dune management.     

Ecology, management and monitoring of grey dunes in Flanders

Grey dunes are a priority habitat type of the European Union Habitats Directive and demand special attention for conservation and management. Knowledge of the ecology of coastal grey dunes can contribute to this policy. Dune grassland succession is initiated by fixation and driven by the complex of soil formation (humus accumulation) and vegetation development. Leaching and mobilization of CaCO₃. which are important in nutrient dynamics, complicate the picture. At present, grass- and scrub encroachment greatly overrules these fine scaled soil processes and causes substantial loss of regional biodiversity. Belgium has an international responsibility in grey dune conservation because of the limited range of its characteristic vegetation, flora and fauna. As biomass removal seems essential in grassland preservation, grazing is an important management tool. Evaluation of management measures focuses on biodiversity measurements on the levels of landscape, community and species.     

Recovery of Danish coastal dune vegetation after a wildfire

The initial recovery of vegetation after a wildfire in a coastal dune area in NW Jutland, Denmark, was studied over a 5-yr period by means of permanent plots representing various dune communities along a topographical gradient. The impact of the fire varied with the position of the plots. Fens and south-facing dunes were little affected while dune heath plots were severely affected including loss of the O-horizon. Post-fire conditions included presence of remaining soil organic matter, a soil seed bank and surviving below-ground plant parts. The soil surface remained stable during the study period. The initial five years of recovery comprised of an initial three-year recruitment phase during which cover and number of species increased and the quantitative species composition changed markedly, followed by two years of a declining rate of change. 38 species of vascular plants were recorded, 35 are regular components in dune, dune heath and heath fen and were recruited from the seed bank, from locally dispersed seeds and/or by sprouting from surviving vegetative parts. The remaining three species were ‘aliens’, dispersed from sources outside the area. Crustose lichens had an important role in the initial recovery by stabilizing the surface and probably inhibiting seed germination, whereas mosses mostly had a subordinate role. The seral position of the plots, as well as the expected time needed for full recovery of pre-fire vegetation, vary with topography and initial soil conditions. Five years after the fire the fen and the south-facing dune probably need less than a decade for full recovery. The remaining plots are judged to be relatively early seral; their full recovery into mature dry or moist dune heath vegetation and O-horizon is expected to need several centuries.     

Nutrient deficiency in dune slack pioneer vegetation: a review

A review of results of fertilization experiments in wet dune slacks is presented. In most cases the above-ground biomass appeared to be limited by nitrogen availability. Primary phosphorus limitation was assessed only once in a dune slack where sod cutting had been applied very recently. In most other case studies phosphorus limits biomass production after nitrogen deficiency was lifted. Potassium availability is of minor importance for biomass production in this type of ecosystem. Singular nitrogen additions led to increased dominance ofCarex andJuncus species as well as perennial grasses, such asAgrostis stolonifera andCalamagrostis epigejos. A combined addition of nitrogen and phosphorus led to total dominance of grasses, while the characteristic basiphilous pioneer species (including mosses) decreased or even disappeared. Certain mechanisms are considered which may maintain nutrient availability in slacks with basiphilous pioneer vegetation at a low level, despite of the accumulation of nutrients in the developing organic soil layer. Some implications for management and further research are discussed.     

Vegetation development influenced by grazing in the coastal dunes near The Hague, The Netherlands

In 1990, grazing was introduced in a section of Meijendel, a coastal sand dune system near The Hague, The Netherlands. After five years an evaluation was made of the effects of grazing on vegetation development. Three transects were established, two in grazed areas and one in an ungrazed area. Field survey data were classified by means of TWIN-SPAN, ordinated with Detrended Correspondence Analysis and the resulting vegetation types interpreted according to Westhoff & den Held (1969). All associations were found in both the grazed and the ungrazed areas, but at the subassociation and variant level some communities appeared to be restricted to the grazed area. These variants were five grassland variants characterized by disturbance indicators such asSenecio sylvaticus andCynoglossum officinale. The total number of plant species in the 19 permanent plots, which had been observed to have been decreasing since 1960, showed a considerable increase after the introduction of horses and cows in 1990. A marked decrease in the cover ofCalamagrostis epigejos andCarex arenaria since 1990 was evident, while in some plots species such asRibes rubrum andViburnum opulus increased considerably. A series of false-colour aerial photographs were used to compare vegetation structure in the three transects between 1990 and 1995. In the grazed area the tall grass vegetation had almost totally disappeared, whereas the areas of open sand. sand with moss and lichens, and low grass vegetation had increased and the pattern had become more fine-grained. In the ungrazed area the area covered by low grass vegetation had increased at the expense of the area of sand with moss and lichens and the pattern had become more coarse-grained.     

Factors affecting vegetation establishment and development in a sand dune chronosequence at Newborough Warren,North Wales

Newborough Warren is a large calcareous west coast UK dune system, which has experienced rapid vegetation spread in the last 70 years. Information from two high resolution chronosequences for dry and wet dune habitats, 0–145 years, was used to answer the following questions: Does climate influence colonisation of vegetation on bare sand? What are the timescales and sequences of successional change in the vegetation? Analysis of aerial photographs showed that stabilisation of the dune system since 1945 has occurred in three main phases. The onset of stabilisation predated myxomatosis by 10 years; while stabilisation virtually halted during the period 1964–1978. Periods of rapid stabilisation were coincident with higher values of Talbot’s Mobility index (M)?>?0.3. Successional development was apparent in both dry and wet habitats. Fixed dune grassland started to replace earlier successional communities at around 40 years, and could persist to 145 years. Linear succession in dune slacks was less apparent, but a separation between communities typically regarded as ‘younger’ and ‘older’ occurred at around 40 years. Species richness in dry dune habitats increased with age to a maximum on soils around 60 years old, then declined again. Species richness was unrelated to age or soil development in wet dune slacks. The influence of climate suggests that conservation managers can only operate within the constraints imposed by natural climatic conditions. Vegetation growth and soil development are closely linked and maintaining some open areas is key to preventing soil development and over-stabilisation.     

Raising groundwater differentially affects mineralization and plant species abundance in dune slacks.

The experience with restoring high water levels (i.e., rewetting) within restoration ecology is limited, and information on changes in soil nutrient supply is scarce. A reduction in nutrient supply is needed to restore the desired oligotrophic vegetation. We determined the effects of restoration of high water levels on decomposition and net carbon (C), nitrogen (N), and phosphorus (P) mineralization rates in wet dune slacks and its consequences for the relative abundance of eutrophic vs. oligotrophic species in the vegetation. This was done by analyzing these variables for valleys that experienced a large groundwater rise vs. valleys that had a small groundwater rise but the same current water level. In addition, the influences of underlying factors (waterlogging, vegetation dieback, and soil dynamics prior to groundwater rise) were separated in a transplantation experiment. Short-term effects of large groundwater rise were a massive dieback of vegetation, increased thickness of the fermentation layer, increased microbial decomposition activity, increased C mineralization, and decreased net N mineralization. Net P mineralization was not affected. The relative abundance of oligotrophic vs. eutrophic species was greater at large groundwater rise. Changes in decomposition and mineralization by large groundwater rise were, however, not caused by the vegetation dieback, but due to previous soil conditions. Soils experiencing waterlogged conditions for 3-4 years or more prior to large groundwater rise had lower C and higher net N mineralization rates at waterlogged conditions than soils that had experienced aerobic conditions, presumably due to differences in labile soil C contents. Contrary to expectations induced by previously determined nutrient pulses and measured vegetation dieback, large groundwater rise resulted in lower soil nutrient supply rates and more oligotrophic vegetation. If these trends continue on the longer term, restoration of high water levels may be effective in restoration ecology to establish oligotrophic, wet vegetation in dune slacks.     

Vegetation dynamics in succession process from Stipa bungeana to Artemisia ordosica in the Ordos Plateau北大核心CSCD

Shrub invasion has a serious effect on the structure and function of grassland ecosystems and understanding vegetation dynamics is of great significance to control shrub invasion and recover shrub invaded grassland. In the Ordos Plateau, we selected representative communities in transition process from Stipa bungeana to Artemisia ordosica. By sampling, cutting, and root-drilling methods, plant coverage, density, biomass, litter, root, and species diversity of different communities were investigated and analyzed. The results showed the following: (1) the succession process had six vegetation types, S. bungeana communities, S. bungeana + Cleistogenes squarrosa communities, S. bungeana + Artemisia ordosica communities, A. ordosica + C. squarrosa communities, A. ordosica + Lespedeza davurica communities, and A. ordosica communities. (2) The community coverage decreased initially, and then increased. Whereas, the total density decreased initially, then increased, and then decreased. The aboveground, underground, and total biomasses, and the dry weight of litter showed an increasing trend. (3) The coverage, density, and biomass of S. bungeana decreased gradually, whereas A. ordosica showed an opposite trend. (4) With the increase in soil depth, the dry weight of root showed a decreasing trend. The roots were mainly distributed in the 0-30 cm soil layer. At VI stage, the root distribution of Artemisia community initially increased, and then decreased, and the root depth reached 80-90 cm. (5) The species richness, Simpson, Shannon-Winner, and Pielou evenness indexes initially increased, and then decreased. In summary, shrub encroachment is severe in the Ordos Plateau. Although the community coverage, biomass, and biodiversity during the moderate shrub encroachment stage were high, the shrub-invaded grassland should be restored to S. bungeana grassland due to the decreased grazing value of grassland after shrub invasion. Keywords. © 2018 Science Press. All rights reserved.     

Seasonal variations in plant species effects on soil N and P dynamics

It is well established that plant species influence ecosystem processes, but we have little ability to predict which vegetation changes will alter ecosystems, or how the effects of a given species might vary seasonally. We established monocultures of eight plant species in a California grassland in order to determine the plant traits that account for species impacts on nitrogen and phosphorus cycling. Plant species differed in their effects on net N mineralization and nitrification rates, and the patterns of species differences varied seasonally. Soil PO4- and microbial P were more strongly affected by slope position than by species. Although most studies focus on litter chemistry as the main determinant of plant species effects on nutrient cycling, this study showed that plant species affected biogeochemical cycling through many traits, including direct traits (litter chemistry and biomass, live-tissue chemistry and biomass) and indirect traits (plant modification of soil bioavailable C and soil microclimate). In fact, species significantly altered N and P cycling even without litter inputs. It became particularly critical to consider the effects of these multiple traits in order to account for seasonal changes in plant species effects on ecosystems. For example, species effects on potential rates of net N mineralization were most strongly influenced by soil bioavailable C in the fall and by litter chemistry in the winter and spring. Under field conditions, species effects on soil microclimate influenced rates of mineralization and nitrification, with species effects on soil temperature being critical in the fall and species effects on soil moisture being important in the dry spring. Overall, this study clearly demonstrated that in order to gain a mechanistic, predictive understanding of plant species effects on ecosystems, it is critical to look beyond plant litter chemistry and to incorporate the effects of multiple plant traits on ecosystems.     

Vegetation succession and lichen diversity on dry coastal calcium-poor dunes and the impact of management experiments

The negative impact of grass and moss encroachment on the botanical diversity of West European coastal dunes attracted increasing attention in the 1990s. This paper focuses on moss encroachment during primary succession in the xeroseries. Until the mid-1970s, vegetation types rich in species of the lichen generaCladonia andCladina were found on the fixed,Corynephorus canescens-dominated, so-called grey dunes all over the island of Terschelling, The Netherlands. In addition, species ofHypogymnia, Parmelia andUsnea, which usually grow on trees, occurred here terrestrially on moss carpets or bare sand. These vegetation types are still present on the Noordsvaarder, a nature reserve in the western part of the island. They occur on parts of seven dune ridges parallel to the coast and form a chronosequence in which age increases with distance from the sea. Our study found the highest lichen diversity on the second and third dune ridges in a stage of primary succession that can be assigned to theViolo-Corynephoretum. The changes from lichen-rich to moss-dominated stadia were significantly related to soil development and acidification in connection with the ageing of the dune soil. The superficial cutting of sods in moss-encroached vegetation appeared to be unsuccessful as a management technique for restoring the biodiversity of cryptogams. Our findings suggest that the best option for maintaining lichen vegetation in theViolo-Corynephoretum is the blow-in of sand with a subneutral or neutral pH from reactivated and natural blowouts or from foredunes, with increasing lime content respectively.     

Response of vegetation and soil ecosystem to mowing and sod removal in the coastal dunes ‘Zwanenwater’, the Netherlands

This paper investigates the effects of mowing and sod removal on vegetation, soil mesofauna and soil profile, and the restoration of these features in the years following sod removal. The sampling site is located in a primary wet dune slack in the northern part of the province of North-Holland. The original vegetation is a heathland withEmpetrum nigrum andCalluna vulgaris, underlain by a Gleyic Arenosol with an O, Ah and (B) horizon development. Above-ground, the vegetation in the dune slack has been mown since 1940. The sod was removed from restricted areas in the slack at various times in the past (1980, 1985, 1987 and 1991). All three sources of data point to adaptation to wet conditions after mowing and sod removal. The vegetation of the mown area is related to the vegetation in the original heathland, although some species appear to have disappeared. Below-ground, mowing drastically reduces the number and depth of occurrence of microarthropoda. Restriction of depth applies also to the Enchytraeidae. Soil profile development is restricted to an Ah-AC-Cr sequence. Species diversity both above and below-ground is relatively high in plots where the sod has recently been cut, due to the rapid colonization by the first pioneer species. A time series for the vegetation is difficult to establish because hydrological conditions interfere with years since sod removal. Soil profile evolution after sod cutting is poor but consistent, from an AC-Cr sequence since 1991, to an O-Ah-Cr sequence since 1980. The management practices were set up with the intention to interrupt the succession to give pioneer species a chance. Neither the vegetation, nor the soil fauna or soil profile have fully recovered during the 13 yr since the first sod removal. So the goal has been reached.     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species.     

Plant winners and losers during grassland N-eutrophication differ in biomass allocation and mycorrhizas

Human activities release tremendous amounts of nitrogenous compounds into the atmosphere. Wet and dry deposition distributes this airborne nitrogen (N) on otherwise pristine ecosystems. This eutrophication process significantly alters the species composition of native grasslands; generally a few nitrophilic plant species become dominant while many other species disappear. The functional equilibrium model predicts that, compared to species that decline in response to N enrichment, nitrophilic grass species should respond to N enrichment with greater biomass allocation aboveground and reduced allocation to roots and mycorrhizas. The mycorrhizal feedback hypothesis states that the composition of mycorrhizal fungal communities may influence the composition of plant communities, and it predicts that N enrichment may generate reciprocal shifts in the species composition of mycorrhizal fungi and plants. We tested these hypotheses with experiments that compared biomass allocation and mycorrhizal function of four grass ecotypes (three species), two that gained and two that lost biomass and cover in response to long-term N enrichment experiments at Cedar Creek and Konza Long-Term Ecological Research grasslands. Local grass ecotypes were grown in soil from their respective sites and inoculated with whole-soil inoculum collected from either fertilized (FERT) or unfertilized (UNFERT) plots. Our results strongly support the functional equilibrium model. In both grassland systems the nitrophilic grass species grew taller, allocated more biomass to shoots than to roots, and formed fewer mycorrhizas compared to the grass species that it replaced. Our results did not fully support the hypothesis that N-induced changes in the mycorrhizal fungal community were drivers of the plant community shifts that accompany N eutrophication. The FERT and UNFERT soil inoculum influenced the growth of the grasses differently, but this varied with site and grass ecotype in both expected and unexpected ways suggesting that ambient soil fertility or other factors may be interacting with mycorrhizal feedbacks.     

植被恢复的生态效应研究进展

植被在水土保持、水源涵养及生态系统的固碳过程中起着重要的作用。植被恢复是指运用生态学原理,通过保护现有植被、封山育林或营造人工林、灌、草植被,修复或重建被毁坏或被破坏的森林和其他自然生态系统,恢复其生物多样性及其生态系统功能。目前,植被的自然及人工恢复是改善脆弱生态系统及退化生态系统生态环境现状最有效的措施。植被在恢复过程中对地上植被生态系统,物种多样性的恢复有着重要影响,同时通过凋落物及根系的输入,可以有效改善地下生态系统,增加土壤的养分含量、改善土壤的物理结构、增加土壤生物的生物量及活性。文章以地上及地下生态系统为出发点,综述了植被恢复过程中自然及人工恢复过程中不同的植被类型、不同的恢复时间下植物物种组成和多样性、土壤理化性质及土壤微生物群落的变化。植被的自然及人工恢复在一定程度上均能增加植物物种的多样性,随着恢复年限的增加物种的组成发生改变且多样性呈增加趋势,但一些特殊环境下不当的人工恢复可造成植被演替向退化方向发展,降低生物多样性。不同的植被类型由于其生长方式的不同对土壤理化性质和土壤微生物的影响存在差异,随着恢复年限的增长,土壤理化指标及微生物学指标呈现先增加而后趋于平稳的状态。针对已有的研究进展,提出在未来的研究过程中,一方面应该增加更多的对比研究,对不同环境下,不同的恢复物种,不同的恢复方式进行更深入地探讨;另外一方面应增加不同尺度的研究,现有的研究多集中在样地尺度,未来应在更大尺度上进行分析;再者,地上及地下生态系统之间的相互关系及影响机理一直是土壤学科研究的热点,植被恢复过程中应增加更多该方面的机理研究。     

Vegetation roles on changes of soil properties and microbial activities in a coastal sand dune. A case study

The vegetation effects on changes of soil physicochemical properties and microbial activities in the costal sand dune were investigated to understand the roles of vegetation on sand dune ecosystem. Eight sites from six vegetation zones and two bare zones in the dune front, dune crest, and dune back regions were selected. Soil microbial enzyme activities of β-glucosidase, acid phosphatase, arylsulfatase, and dehydrogenase, and soil physicochemical properties of each site were evaluated. The results showed that all the enzyme activities were higher in the mixed vegetation sites with native sand dune plants and naturalized plants and in Pinus thunbergii community site both located in the dune back regions where the accumulation of organic matter and nitrogen were more prominent. The results demonstrated that soil organic matter and nutrients are the primary determinants of the microbial activity in sand dune where are exposed to a gradient of physicochemical stress such as high salinity, moisture and salt spray. Therefore, the conservation of vegetation that generates more soil organic matter and nutrients is important factor in controlling the soil microbial activities and biogeochemical cycles in the coastal sand dune systems.     

The biogeochemical cycles of phosphorus: A review of local and global consequences of the atmospheric input

P‐input from the atmosphere is, in many oligotrophic ecosystems, substantial for the biomass production; in some regions biomass formation may depend fully or partially on the phosphorus input from the atmosphere. As a consequence, phosphorus must be considered as an element participating in cycles involving the atmosphere, like sulfur and nitrogen. Dust and aerosols containing phosphorus are transported worldwide, linking even distant regions. Human activities enhance the amounts of P distributed. Since the concentrations of P in the atmospheric dry and wet input are usually very low, special care in sampling and analysis is a prerequisite to obtain reproducible data. Some values in the literature may be questionable.     

三江源国家公园不同草地土壤微生物功能基因的差异性分析

土壤微生物在陆地生物地球化学循环过程中起着非常重要的作用。为了探索青藏高原高寒草地类型地上植被特性和地下土壤环境与土壤微生物功能基因之间关系,以三江源国家公园高寒草原、高寒沼泽化草甸及高寒草甸3种典型草地类型为研究对象,利用基因芯片(GeoChip 5.0)技术测定其微生物功能基因丰度,并分析它们之间的差异及影响因素。结果表明:(1)3种草地类型地上群落结构和地下土壤环境存在差异性,其中高寒草原物种多样性指数、pH值较高,沼泽化草甸中土壤含水量、微生物量碳、地上生物量、土壤速效氮含量较高,高寒草甸中则是土壤微生物量氮含量较高;(2)3种高寒草地类型的碳循环、氮循环、磷循环、有机修复的土壤微生物功能基因丰度存在显著差异,其中这些功能基因的丰度在高寒沼泽化草甸最高,高寒草甸、高寒草原次之;(3)地上植物物种多样性虽对功能基因丰度变化的解释率(r2)在57.1%-61.2%之间,但统计学上不显著(P>0.05),而微生物基因丰度随地上生物量的增加而增加,且解释率(r2)为77.5%-80.0%(P<0.05)。在pH、土壤含水量、土壤微生物量等地下土壤环境因子中,pH对功能基因丰度存在显著影响(P<0.01)解释率在83.4%-87.5%间,且土壤微生物功能基因丰度随土壤pH的增加而降低;土壤含水量、土壤微生物量对土壤微生物功能基因丰度的解释率分别为81.9%-83.1%(P<0.05)和76.8%-86.2%(P<0.05),微生物功能基因丰度随这两者含量的增加呈上升趋势。进一步运用RDA分析发现,pH、土壤微生物量、地上生物量是影响微生物功能基因丰度的主要因子,其中土壤微生物量是土壤有机质的重要组成部分,土壤有机质又是通过地上植被凋落物沉积所得到的。因此,地上植被特性的自上而下控制因子影响了土壤环境中自下而上的控制因子,间接的影响了微生物功能基因丰度。由此得出,地上植被特性和地下土壤环境因子共同作用控制了微生物功能基因丰度使其出现差异性。     

The impact of grazing on spider communities in a mesophytic calcareous dune grassland

During 1994–1995 and 1997–1998 spiders were sampled with pitfall traps in a botanically rich, mesophytic, calcareous dune grassland in Belgium. As a consequence of intensive cattle grazing, vegetation variation in a large part of the area had diminished. The study area was also patchily grazed by rabbits. Community analysis with TWINSPAN revealed five distinct spider communities. Ecological differentiation was best explained by combination of the habitat variables: distance from grazed or non-grazed vegetation,Rosa pimpinellifolia cover and grass cover in both summer and winter. Species diversity was highest in the border zone between the cattle-grazed and non cattle-grazed sites. Correlation of the most abundant spider species with the vegetation determinants explains the ecological differentiation between the spider communities. Species were classified into seven major groups that reflect the species’ habitat preferences. The group showing clear association with non cattle-grazed, tall vegetation consists of common species. Characteristic species for the intensively cattle-grazed sites are common aeronauts and rare species such asWalckenaeria stylifrons, Mastigusa arietina, Ceratinopsis romana andPardosa monticola. The latter are shown to be dependent on ungrazed vegetation for juvenile development and overwintering. Intensive grazing results in homogeneous short vegetation, which can only be colonized by ‘open ground’ species with a well-developed dispersal capacity, or by species which are not dependent on litter-rich situations for juvenile development. An extensive cattle grazing regime results in a patchy mosaic grassland where, in addition to the above mentioned groups of species, other species survive by migrating between the buffered litter rich ungrazed vegetation and the short vegetation. Additionally, some typical and rare species prefer the transition zone between the grazed and the ungrazed vegetation because they are associated with specific habitat structures or inhabiting ant-species. Nomenclature: Roberts (1987, 1995) forAraneae; van der Meijden et al. (1990) for vascular plants; Corly et al. (1981) for bryophytes; Schaminée et al. (1996) for vegetation associations.     

Coastal dune stabilization in Wales and requirements for rejuvenation

Welsh coastal dune systems have become increasingly vegetated in recent decades. Several rare species of plants and invertebrates have declined dramatically in abundance, and in some areas lost entirely. Of the ten dune habitats and species recognized as being features of European importance within the Welsh Natura 2000 sites, nine are currently in Unfavourable condition on at least one site. The decline in active aeolian processes has also reduced the geomorphological interest of the sites, several of which were designated as Geological Conservation Review sites principally on the basis of their physical processes and landforms. The decline in bare sand area between the 1940-50s and 2009 has been quantified at twelve Welsh dune sites using aerial photography and GIS. The decline ranged from 41 % at Gronant Dunes and Talacre Warren to 97 % at Kenfig Burrows, with an average of 81 %. Morfa Dyffryn had the highest remaining percentage of bare sand in 2009 (20 %), with 30–40 % coverage of mobile dune and pioneer communities, while seven sites had < 5 % bare sand. Dune stabilization over the past 60 years has been favoured by a number of factors, including less windy conditions, higher temperatures and longer growing season, increased atmospheric nitrogen deposition, a reduction in grazing intensity, and dune management policies aimed at controlling mobile sand. Climate change projections suggest that, in the next 50 to 100 years, Wales and adjoining areas are likely to experience higher temperatures and higher rainfall, especially in winter, and a further slight reduction in wind speeds. Without intervention, dune and dune slack habitats are likely to be increasingly replaced by fixed dune grassland and scrub, resulting in the extinction of rare plants, invertebrates and other species which require open, mobile conditions. Several intervention options exist, ranging in scale and potential impact. Increased livestock grazing, re-introduction of rabbits, scrub clearance, turf stripping and the creation of shallow ‘scrapes’ can be beneficial but will not by themselves create self-sustaining mobile dunes. In order to have any chance of achieving any significant impact, larger-scale intervention measures, involving large-scale vegetation removal and sand-re-profiling, will be required. At least in the short-term, maintenance measures will be required to prevent vegetation re-growth, and the challenge will be to encourage the development of mobile dune features which will be naturally mobile in the medium to longer term.     

Grassland restoration with and without fire: evidence from a tree-removal experiment

Forest encroachment threatens the biological diversity of grasslands globally. Positive feedbacks can reinforce the process, affecting soils and ground vegetation, ultimately leading to replacement of grassland by forest species. We tested whether restoration treatments (tree removal, with or without fire) reversed effects of nearly two centuries of encroachment by Abies grandis and Pinus contorta into dry, montane meadows in the Cascade Range, Oregon, USA. In nine, 1-ha plots containing a patchy mosaic of meadow openings and forests of varying age (20 to > 140 yr), we compared three treatments affecting the ground vegetation: control (no trees removed), unburned (trees removed, slash burned in piles leaving 90% of the area unburned), and burned (trees removed, slash broadcast burned). We quantified changes over 3-4 years in soils, abundance and richness of species with differing habitat associations (meadow, forest, and ruderal), and recruitment of conifers. Except for a transient increase in available N (especially in burn scars), effects of burning on soils were minimal due, in part, to mixing by gophers. Tree removal greatly benefited meadow species at the expense of forest herbs. Cover and richness of meadow species increased by 47% and 38% of initial values in unburned plots, but changed minimally in burned plots. In contrast, cover and richness of forest herbs declined by 44% and 26% in unburned plots and by 79% and 58% in burned plots. Ruderal species and conifer seedlings were uncommon in both treatments. Although vegetation was consumed beneath burn piles, meadow species recovered significantly after three years. Long-term tree presence did not preclude recovery of meadow species; in fact, colonization was greater in older than in younger forests. In sum, temporal trends were positive for most indicators, suggesting strong potential for restoration. Contrary to conventional wisdom, tree removal without fire may be sufficient to shift the balance from forest to meadow species. In meadows characterized by historically infrequent fire, small-scale disturbances and competitive interactions may be more critical to ecosystem maintenance and restoration. Managers facing the worldwide phenomenon of tree invasion should critically evaluate the ecological vs. operational need for fire in ecosystem restoration.