Physical damage to coastal dunes and ecological impacts caused by vehicle tracks associated with beach camping on sandy shores: a case study from Fraser Island,Australia

As coastal populations expand, demands for recreational opportunities on beaches and coastal dunes grow correspondingly. Although dunes are known to be sensitive to direct human disturbance and provide irreplaceable ecosystem services (e.g. erosion control, critical habitat and nesting sites), dunes serve as campsites for large numbers of people (∼90,000 p.a.) on the ocean-exposed shores of Fraser Island, Australia. Campsites are located in the established dunes and can only be accessed with 4WD vehicles along tracks cut directly from the beach through the foredunes. Here we quantified the extent of physical damage to foredunes caused by this practice, and tested whether human-induced physical changes to foredunes translate into biological effects. Of the 124 km of ocean-exposed beaches, 122 km (98%) are open to vehicles driven on the beaches, and camping zones cover 28.7 km or 23% of the dunes. A total of 235 vehicle tracks are cut across the foredunes at an average density of eight tracks per km of beach. These tracks have effectively destroyed one-fifth (20.2%) of the dune front in camping zones, deeply incising the dune-beach interface. There is evidence of accelerated erosion and shoreline retreat centred around vehicle tracks, resulting in a “scalloping” of the shoreline. No dune vegetation remains in the tracks and the abundance of ghost crabs (Ocypode spp.) is significantly reduced compared with the abutting dunes. Because current levels of environmental change caused by dune camping may not be compatible with the sustainable use of coastal resources and conservation obligations for the island (listed as a World Heritage Area and gazetted as a National Park), restoration and mitigation interventions are critical. These will require spatial prioritisation of effort, and we present a multi-criteria ranking method, based on quantitative measures of environmental damage and ecological attributes, to objectively target rehabilitation and conservation measures. Ultimately, coastal management needs to develop and implement strategies that reconcile demands for human recreation, including beach camping, with conservation of coastal dune ecosystems.     

沙地退化植被恢复过程中植被的空间异质性

通过野外取样和室内分析,应用地统计学分析方法研究了科尔沁沙地退化植被恢复过程中不同封育年限（0、11和20年）的流动沙丘的植被盖度和丰富度特征及其空间异质性规律。结果表明,随着流动沙丘的固定和封育年限的增加,植被盖度和丰富度逐渐增加。在流动沙丘植被恢复过程中,植被盖度和丰富度具有明显的空间自相关性,其空间自相关范围从封育0年的流动沙丘（46.05m和33.63m）、封育11的流动沙丘（21.63m和17.25m）到封育20年的流动沙丘（26.12m和24.18m）先减小后增加,但均未超出我们的研究尺度50m,表现出不同大小的斑块形式分布的小尺度分布格局。由半方差函数及其参数和空间分布格局图分析得出,随着沙丘植被的恢复,植被特征的空间异质性在所研究的尺度上表现出先增大（封育0年到封育11年）后减小（封育11年到封育20年）的变化特点。     

The impact of motorcycle traffic on soil and vegetation of stabilized coastal dunes, Israel

This study aimed to assess the response of soil and annual plant vegetation of stabilized coastal dunes in Israel, to varying intensities of off-road motorcycle (ORM) traffic, and to assess their resistance and resilience to such a disturbance. A standard experimental procedure that included 0, 20, 50, 100 and 200 ORM straight passes and 150 ORM turn passes was used. Plant ground cover, mean plant height, species richness, species diversity, soil penetrable depth, organic matter and moisture contents were measured on several dates within a period of 372 days after the experiment. Results have shown that: (1) ORM passes had a significant immediate impact on annual plants that increased with traffic intensity. The impact on the soil was detected only as an increase of penetrable depth. (2) The maximum impact on annual plants was observed in the wheel ruts and turn areas. The impact on the area between the wheel ruts and on the margins outside the wheel ruts was indirect and smaller. (3) Annual plant ground cover and mean plant height were less sensitive parameters than species richness and species diversity for determining the impact of ORM traffic on the area. (4) One year after the experiment, soil and annual plant vegetation in all passes were very similar to their pre-experimental condition. This indicates high resilience and recovery potential of the Mediterranean stabilized coastal dune ecosystem to ORM disturbance.     

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.     

Sustainability of sand dune restoration along the coast of the Tyrrhenian sea

This study reports the results of restoration management on sand dune environments along the coastal belt of the Castelporziano nature reserve (Rome, Italy) and the subsequent monitoring phases to test the sustainability of the ‘soft techniques’ applied. In the area concerned, over a length of ca. 3 km, 40 dunes were built up along with three belts located at <40 m, 40–70 m, and >70 m, respectively, from the shoreline. On each of 38 dunes 20 individuals ofAmmophila littoralis were planted; this species is one of the local autochthonous species considered particularly suitable for stabilizing sand dunes. After one year, two years and five years, respectively the changes in height and surface of each dune, the survival rates ofA. littoralis, and its changes in cover, the appearance of new shoots and the establishment of new species were observed. A progressive increase in species number, which five years after the restoration amounted to about 60% of those characterizing the natural dunes, was reported indicating a progressive trend towards populations similar to natural ones. In the colonization of new species there is a prevalence of theSporobolus-Elymetum farcti and theSalsolo Kali-Cakiletum maritimae association, while the species established successively refer to theEchinophoro spinosae-Ammophiletum arundinaceae association and theCrucianellion maritimae alliance as occurring in natural successions. This succession runs parallel to the natural dune colonization processes. In particular, the data regarding survival, cover and number of vegetative shoots indicate that the dune belt between 40 and 70 m from the sea is the one most suitable for restoration. Some changes in dune morphology was observed: the height of the artificial dunes tended to decrease considerably in the five years of observation, whereas a progressive increase in their surface area was observed. During the study period.A littoralis favoured the establishment of new species, but as yet exercises no action on increasing dune height.     

Temporal and spatial changes in plant dune diversity in urban resorts

We investigated the temporal and spatial changes in the floristic composition and abundance in sand dunes along a coastal strip in the province of Buenos Aires (Argentina). Firstly, we analysed changes over a span of 70 years, comparing 41 of our own inventories carried out in 14 beach resorts in 2005–2006 with 18 inventories made in the first half of the XXth century (1930–1940) in the same study area. We grouped sampled plots into four categories for comparison, taking into account the time period (1930–1940 or 2005–2006) and location at each end of the surveyed coastal strip in the northern or southern ‘tourist’ sectors. Secondly, we analysed the effect of afforestation with exotic trees on extant sand dune vegetation for the period 2005–2006. A total of 74 plant inventories were carried out in afforested and natural dunes at the same resorts. In both comparisons we contrasted plant richness, percentage plant cover, similarity and diversity. A cluster analysis was used to classify categories taking into account plant abundance and composition. Significant differences in total diversity were observed between sampling years, suggesting an increase in diversity at the present time due to urbanization and recreation related activities. Nevertheless, no single exotic plant species was clearly dominant across the sampled sites, suggesting that these coastal areas are, up to now, resistant to alien invasion. Diversity indexes discriminated by plant groups indicating different habitat conditions and exotics were shown to be more sensitive to existing changes than to total richness and cover.     

Floristic composition and vegetation ecology of the Malindi bay coastal dune field, Kenya

A short outline is given of the floristic composition, structure and distribution of coastal dune vegetation found at Malindi Bay, Kenya. The area was studied by air photo interpretation and field sampling to determine the relationship of plants to aeolian features. TWINSPAN classification was used to distinguish geomorphological units on the basis of their species composition. In this paper, an inventory and first quantitative analysis of vegetation distribution is presented. We identified 174 plant species from 62 families in the sand dunes and several plant communities are distinguished based on the species content and the connection with morphological units.Papilionaceae with 18 species andPoaceae with 17 species were the most represented families. A distinct zonal distribution of the plant communities was found. The most important plant species are the pioneer vegetation consisting ofHalopyrum mucronatum, Ipomoea pescaprae andScaevola plumieri. The woody shrub species which have colonized the established primary dunes and hummock dunes areCordia somaliensis, Pluchea discoridis, Tephrosia purpurea (dunensis). Succulent herbs were identified in the dune slacks and salt marsh that are moist and damp environments.     

Early development of vegetation in restored dune plant microhabitats on a nourished beach at Ocean City, New Jersey

Topography and vegetation of restored dunes on a developed barrier island were examined after a large-scale beach nourishment project. Restoration began in 1993 using sand-trapping fences andAmmophila breviligulata Fern. plantings. Subsequent growth of dunes was favored by installing new fences and suspending beach raking to accommodate nesting birds. Plant species richness, percent cover of vegetation, and height ofA. breviligulata were sampled in 1999 on seven shore perpendicular transects in six dune microhabitats (backdune, primary crest, mid-foredune, swale, seaward-most fenced ridge, incipient dune on the backbeach). A total of 26 plant taxa were found at all seven sites. Richness and percent cover were greatest in the backdune and crest, especially in locations that predated the 1992 nourishment. Richness was greater where fences enhanced stabilization. Fences initially compensate for time and space and allow vegetation to develop rapidly, but maintenance nourishment is required to protect against wave erosion and ensure long-term viability of habitat. An expanded environmental gradient is an option, where beach nourishment provides space for a species-rich crest and backdune to develop, while the incipient dune remains dynamic. Options where space is restricted include a dynamic, full-sized seaward section of a naturally functioning dune (truncated gradient) or a spatially restricted sampler of a wider natural dune (compressed gradient) maintained using fences. Expanded and truncated gradients may become self-sustaining and provide examples of natural cycles of change. Compressed gradients provide greater species richness and flood protection for the available space, but habitats are vulnerable to erosion, and resident views may be impaired.     

Inventory and conservation assessment for the management of coastal dunes,Veneto coasts,Italy

Management of coastal dunes on developed coasts could effectively take advantage of comprehensive and multitemporal georeferenced data collection, which offers the possibility to relate dune data with the natural and cultural characteristics of the beach and hinterland. The recent implementation of a coastal management geodatabase for the Veneto region provides the opportunity for improving knowledge on coastal dunes on developed littoral as well as a basis for appropriate future coastal planning in the study area. The geodatabase gathers data concerning different physical, evolutionary and human aspects of the coastal zone, with a special focus on coastal dunes. Established foredunes, human-altered dunes and relict dunes are irregularly distributed along 59 km, 38% of the entire coastal length. Their distribution and characteristics are the result of favourable natural conditions as well as long-lasting tourism exploitation (evaluated through an index of Land Use Pressure) and fragmentary and diversified uses of beaches (evaluated through an index of Tourism Pressure on the beach). At the same time, beach/dune nourishment intervention allowed the presence of artificial or sand fenced dunes along 17 km of coast. High dune elevation up to 8–10 m is promoted by the onshore exposition of the beach to dominant wind (from ENE), by stable-to-slowly negative sedimentary budget or by the re-activation of high relict foredunes in the case of shoreline retreat associated with strong negative budget. Present sedimentary budget (evaluated through the code ASPE – Accretion, Stable, Precarious, Erosive) is the tool used for dividing coasts in sedimentary compartments or cells. Past and present sedimentary budget and different human responses to erosive cases (hard and soft interventions) give the foredunes different means to form, grow, survive and evolve over time. The assessment of Human Impact and Active Management Practices on the dunes allows a first evaluation of the Management Effectiveness, which shows strong shortcomings for 81% of the dunes. The great variability of beach usage, human impact and management practises on the different dune stretches highlight the lack of effective and systematic management actions being correctly scheduled and performed.     

The importance of coastal sand dunes for terrestrial invertebrates in Wales and the UK,with particular reference to aculeate Hymenoptera (bees,wasps &; ants)

Although most UK sand dune systems are now fossilized, with little mobility and reducing amounts of bare sand, they support important populations and assemblages of terrestrial invertebrates. Offering open conditions, warm substrates and a range of habitats and habitat structures, they have become increasingly significant as other coastal habitats have been lost. In Wales, 680 Red Data Book and Nationally Scarce species have been recorded from dunes. 109 species in the UK are restricted to dunes, and in Wales there are an additional 145 species confined to dunes and 208 species strongly associated with dunes. Of these, 172 species are dependent upon bare and sparsely-vegetated sand, in grey dunes and early-successional dune grassland, at some stage of their life cycle, rising to 292 species if those associated with the strandline, foredunes, yellow dunes and pioneer dune slacks are included, equating to 63% of the 462 dune species. Bees and wasps are particularly well represented, with 278 species (68% of the Welsh fauna) recorded on Welsh dunes, including 17 obligates and 44 species with a strong dependence, 52 of which are associated with bare and sparsely-vegetated sand. Key to maintaining invertebrate populations on UK dunes is the provision of bare sand but in Wales, bare sand accounts for only 1.7% of the total sand dune resource. As a more appropriate bare sand threshold is likely to range between 10 and 30%, radical action is required to re-mobilize at least the key sand dune systems.     

Contrasting ecology of prograding coastal dunes on the northwest coast of Ireland

This paper examines the influence of dune morphological evolution on plant species diversity and composition on the Magheramore dune system, Donegal, north-west Ireland. It aims to demonstrate the need for understanding of local geomorphological factors and their affect on ecological processes for the enhanced conservation management of coastal dunes. Vegetation surveys and cluster analysis of species associations, in combination with GIS analysis focused on two prograding dune sites within the Magheramore system. Contrasting geomorphic evolution over similar timescales has imposed site-specific environmental conditions that have had a direct impact on the range of plant species communities present. Cluster analysis revealed 5 main species community types that defined specific dune habitats, the organisation of which is strongly influenced by geomorphic evolution. This study demonstrates the importance of geomorphic evolution and morphological heterogeneity in habitat and species diversity.     

Vegetation succession and lichen diversity on dry coastal calciumpoor 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. Nomenclature: Van der Meijden (1990) for higher plants; Touw & Rubers (1989) for mosses; Purvis et al. (1992) for lichens. A set of lichen specimens has been deposited at the herbarium of Dr. A. Aptroot (C.B.S. Baarn) — herbarium code ABL (Taxon 44: 258).     

The effect of grazing on biodiversity in coastal dune heathlands

The effect of sheep grazing on species richness, higher order diversity measures, inequality, species composition, functional diversity and allometric relationships at a coastal dune heathland site was investigated. After a prescribed fire in 2002, the site was divided into two parts, where one of the parts was unmanaged and the other part was fenced and grazed by sheep. Sheep grazing had a positive effect on species richness as well as a significant positive effect on the functional diversity at the coastal dune heathland site. Generally, the cover of dwarf shrubs was negatively affected by grazing, whereas the cover of sedges and grasses was positively affected by grazing. There is a need for comparative investigations of the effect of different management methods not only on floristic biodiversity, but on all relevant kinds of biodiversity as well as on soil structure, soil chemistry and habitat micro- and macrostructure. Consequently, we advocate the initiation of an international systematic investigation of the effect of different management methods.     

Coastal sand dune vegetation: An extreme case of species invasion

The coastal sand-dune flora of the Gulf and Caribbean region of Mexico was analyzed to understand differences in floristic composition and richness found along the coast. Each of the 655 species reported was classified according to its ecology and distribution range by checking herbaria specimens, literature and specialists. Three groups were formed: (a) species with predominantly coastal distribution; (b) ruderal or secondary species frequently found inland, common of disturbed areas such as roadsides, abandoned fields or forming part of secondary growths; (c) inland species frequently found in other vegetation types such as tropical dry or seasonal forest and grassland. A total of 71 coastal species, 237 ruderal/secondary and 336 species from other community types were found. The distribution of these groups was analyzed along 44 sites of the Gulf and Caribbean, in the different dune habitats and for the dominant growth forms. Coastal species are more widely distributed; they predominate in habitats with sand movement and the herbaceous component prevails. Ruderal/secondary species and especially those belonging to other vegetation types frequently appear in only one or two sites occupying more protected or stabilized habitats. The two latter groups considerably increase species richness of sand dune flora, but also pose interesting problems for dune conservation.     

A novelty coastal susceptibility assessment method: application to Valdelagrana area (SW Spain)

The main aim of this paper is to present a new methodology to determine coastal susceptibility to erosion and flooding processes by means of an index-based method. The proposed indices take into account physical parameters, such as dune and beach geomorphologic characteristics, shoreline evolution, local significant wave height and relative run-up. The coastal susceptibility has been estimated by elaboration of spatial input data into a GIS environment. The method has been tested in Valdelagrana area, a sandy spit located in SW Spain. The spit includes several morpho-sedimentary environments: sandy beach, discontinuous embryo dunes and foredunes, mud flats and wide areas of vegetated salt marshes. The Northernmost sector is densely urbanized whilst the rest is part of a natural protected area belonging to the Bahía de Cádiz Natural Park. The results obtained showed how the Southern part of the spit presented a high susceptibility due to an elevated erosion rate and the presence of low and discontinuous dune ridges.     

Lichens as indicators of a perturbation/stability gradient in the Asperillo dunes, SW spain

In the asperillo dune system, Southwest Spain, lichen vegetation covering the dune sand, has a low species diversity but is an important component of the perennial vegetation, providing stability, nutrients, and moisture to the soil layer. The Asperillo dunes harbour (1) natural ecosystems, (2) disturbed systems affected by forestry activities where the natural vegetation is eliminated, and (3) pine forest resulting from afforestation withPinus pinea since the end of the 1940s where the composition and abundance of the lichen flora has been drastically changed. Our hypothesis was that the lichen species composition and distribution are influenced by the type and period of human intervention. To test this hypothesis a 1.2 km × 1.1 km area, which includes zones with different degrees of conservation and perturbation, consisting of five different habitats, was selected: well preservedJuniperus woodland,Pinus pinea plantations with dense scrub, dense scrub,P. pinea plantations cleared from scrub, and pioneer scrub in recent pine afforestation. In each area, randomly located 5m × 5 m homogeneous plots were sampled. In each plot the cover of shrub and lichen species was recorded. The first results show that lichen cover is higher in the most preserved areas,Cladonia mediterranea being the dominant species under theJuniperus phoenicea canopy. The lichen community changes in composition and abundance depending on the time elapsed since the last perturbation.     

Trampling impact on vegetation of embryonic and stabilised sand dunes in Montenegro

Trampling is one of the human activities that are harmful for plant species and communities of sand dune ecosystems. The aim of this study was to compare the vegetation of embryonic and shifting Ammophila sand dunes with and without fencing to limit trampling. Fenced sand dunes appeared to be richer in species but differences were more prominent in embryonic sand dunes. Some species (Cakile maritima, Pancratium maritimum) were missing on trampled embryonic dunes. The positive impact of trampling exclusion on embryonic sand dunes was indicated by a lowered slope in a Whittaker graph as well as by rarefaction curves that showed higher species richness on the lower slope. Changes in the vegetation of more stabilised shifting Ammophila sand dunes due to trampling are not evident, although species composition is also impoverished. Fencing of parts of sand dunes proved to be an effective measure for vegetation conservation. In addition to physical exclusion of visitors, fences can also have symbolic value for raising public awareness.     

Cattle Grazing Impacts on Annual Forbs and Vegetation Composition of Mesic Grasslands in California

Abstract:   Livestock grazing represents a major human alteration of natural disturbance regimes in grasslands throughout the world, and its impacts on plant communities have been highly debated. We investigated the impact of cattle grazing on the California coastal prairie plant community with a focus on native annual forbs, a number of which are of conservation concern. In spring 2000 and 2001, we surveyed the vegetation community composition, vegetation structure, and soil chemical parameters at 25 paired grazed and ungrazed sites over a 670-km range of the ecosystem. Native annual forb species richness and cover were higher in grazed sites, and this effect was concomitant with decreased vegetation height and litter depth. Soil properties explained less of the variation. Exotic annual grass and forb cover were higher in grazed sites. Native grass cover and species richness did not differ in grazed and ungrazed sites, but cover and species richness of native perennial forbs were higher in ungrazed sites. Our results suggest that cattle grazing may be a valuable management tool with which to conserve native annual forbs in the ecosystem we studied but that grazing differentially affects the various life-history guilds. Therefore, land managers must focus on creating a matrix of disturbance regimes to maintain the suite of species native to these mesic grasslands. The results of this and other studies highlight the importance of considering the adaptation of vegetation communities to disturbance in making recommendations for grazing management.     

Site specific factors have an overriding impact on Baltic dune vegetation change under low to moderate N-deposition—a case study from Hiddensee island

At three coastal dune sites at the island of Hiddensee, north-east Germany, vegetation cover was mapped during 2002 and compared to vegetation surveys from the late 1980s and 1930s. Abiotic and biotic factors, which have been identified as being critical for coastal dunes in former studies such as disturbance, salt spray or nutrient availability, were measured. Grazing and land-use history were reviewed by literature and interviews. Tall graminoid communities, mainly Carex arenaria, are a common vegetation unit today. Development, distribution of these dominances and possible causes for its occurrence have not been analysed. Generally, older successional vegetation units increased and pioneer stages decreased from the 1930s until 2002. At the geologically youngest site, the southern dunes, grass encroachment by Carex arenaria was highest (ca. 50% cover in 2002), and age and density of trees lower than at the older, central dunes. Land-use changes such as decrease in grazing pressure, additional feeding of livestock, increase in coastal protection measures and subsequent decrease in shifting sands as well as varying availability of groundwater and amount of salt spray are relevant factors for vegetation changes in coastal dunes over the past 70 years. Site-specific land-use differences such as livestock density and land-use history have a stronger influence than atmospheric N-pollution on the vegetation composition of these acidic, coastal dunes under low to moderate N-deposition loads of 6–8 kg N ha⁻¹ yr⁻¹.     

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.