Potential impact of a toxic dinoflagellate (Alexandrium excavatum) bloom on survival of fish and crustacean larvae

During the last 20 yr the western half of the Dutch Wadden Sea has undergone significant eutrophication: concentrations of P and N compounds and planktonic algae have roughly doubled, as has primary production. Though oxygen levels are often low in summer, anoxic areas are small and rare due to strong tidal mixing. During the 1970 to 1990 period, macrozoobenthos was sampled annually at 15 stations at Balgzand, a 50-km² tidal-flat area in the westernmost part of the Wadden Sea. Not only did the estimates of total numbers, biomass, and production double during these two decades, but significant changes in the composition of the benthic community were observed, too: (1) the numerical proportion of polychaetes increased at the expense of molluscs and crustaceans, (2) the overall mean weight per individual of the macrozoobenthos decreased (numbers of individuals of small-sized species increased more rapidly than those of large-sized species), and (3) though absolute numbers and biomass of all feeding types increased, the share of carnivores declined and that of deposit feeders increased; the proportion of suspension feeders showed little change. This study refers to true macrobenthos only (1-mm sieve) and further excludes two taxa (Corophium spp. andHydrobia ulvae) which occasionally exercised an undue influence on numbers. Mass mortalities caused by low oxygen concentrations were of a small-scale nature only. Total number of species fluctuated without a clear trend. As a consequence of the increasing numerical densities, trends in species numbers were slightly increasing when expressed per unit of area and slightly decreasing when estimated per 100 individuals (by rarefaction).     

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.     

A new method for detecting pollution effects on marine macrobenthic communities

A method is described by which the pollution status of a marine macrobenthic community may be assessed without reference to a temporal or spatial series of control samples. Theoretical considerations suggest that the distribution of numbers of individuals among species should behave differently from the distribution of biomass among species when influenced by pollution-induced disturbance. Combined k-dominance plots for species biomass and numbers take three possible forms representing unpolluted, moderately polluted and grossly polluted conditions, one curve acting as an internal control against which the other can be compared. Field data from unpolluted communities and from a well documented temporal pollution gradient support the model, but further empirical testing is required.     

Relative Effects of Disturbance on Red Imported Fire Ants and Native Ant Species in a Longleaf Pine Ecosystem

Abstract: The degree to which changes in community composition mediate the probability of colonization and spread of non‐native species is not well understood, especially in animal communities. High species richness may hinder the establishment of non‐native species. Distinguishing between this scenario and cases in which non‐native species become established in intact (lacking extensive anthropogenic soil disturbance) communities and subsequently diminish the abundance and richness of native species is challenging on the basis of observation alone. The red imported fire ant (Solenopsis invicta), an invasive species that occurs throughout much of the southeastern United States, is such an example. Rather than competitively displacing native species, fire ants may become established only in disturbed areas in which native species richness and abundance are already reduced. We used insecticide to reduce the abundance of native ants and fire ants in four experimental plots. We then observed the reassembly and reestablishment of the ants in these plots for 1 year after treatment. The abundance of fire ants in treated plots did not differ from abundance in control plots 1 year after treatment. Likewise, the abundance of native ants increased to levels comparable to those in control plots after 1 year. Our findings suggest that factors other than large reductions in ant abundance and species density (number of species per unit area) may affect the establishment of fire ants and that the response of native ants and fire ants to disturbance can be comparable.     

Detection of pollution effects on marine macrobenthos: further evaluation of the species abundance/biomass method

Further empirical data are presented to evaluate the method of detecting pollution-induced disturbance in marine benthic communities by a comparison of the distribution of numbers of individuals among species with the distribution of biomass among species. A suggested abbreviated name for the technique is the ABC method (abundance biomass comparison). Application of the technique to new data shows that it is a sensitive indicator of natural physical and biological disturbance as well as pollution-induced disturbance over both spatial and temporal scales. Changes in the configuration of ABC plots during ecological succession are the reverse of those resulting from increased pollution levels. The technique should also be applicable to intertidal sediments, where physical disturbance of the sediment by waves does not appear to preclude its utility.     

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.     

Seasonal and annual variability in an epifaunal community in the German Bight

While the benthic infauna of the North Sea has been studied intensively over the past decades, few studies have focused on the larger mobile epifauna. Studies carried out to date have described the distribution of epifaunal communities over the whole of the North Sea, but variability within the identified communities, which occurs on a much smaller scale, has so far remained unstudied. This is the first study to describe seasonal and annual variability of an epifaunal assemblage in the German Bight area, where environmental conditions are highly variable. The benthic community was sampled with a 2 m beam trawl from 1998 to 2001. The echinoderms Ophiura albida and Asterias rubens and the crustacean Pagurus bernhardus were the dominant species caught throughout the study period. Overall the species composition of the catches was relatively consistent, while abundances of dominant species fluctuated considerably between sampling periods. Differences between sampling periods were not only influenced by the abundances of dominant species, but also by less dominant species such as Ophiura ophiura, Astropecten irregularis, Corystes cassivelaunus, Crangon crangon and Aphorrahis pespelicani. The abundances of these species varied annually and seasonally in the assemblage. Clear differences between summer and winter in the species composition, abundance and biomass were identified. Annual and seasonal changes were most likely linked to migratory movements of epifauna into and out of the area under investigation. Temporal changes in species composition and abundance correlated best with water temperatures, while the spatial distribution of the total biomass over the whole sampling period was correlated with sediment characteristics. Anthropogenic influences such as fishing activity and chronic large-scale eutrophication are thought to have influenced the community on a long-term basis, but have been considered unlikely causes for the short-term variability described by this study.Communicated by J.P. Thorpe, Port Erin     

Morphological evidence for vertical transmission of symbiotic bacteria in the viviparous sponge<Emphasis Type="Italic"> Halisarca dujardini</Emphasis> Johnston (Porifera,Demospongiae, Halisarcida)

All stages of vertical transmission of symbiotic bacteria, from the penetration into oocytes to the formation of rhagon, were investigated in the White Sea (Arctic) representatives of Halisarca dujardini Johnston (Demospongiae). Small populations of free-living specific symbiotic bacteria inhabit the mesohyl of H. dujardini. They are represented by a single morphotype of small spiral gram-positive bacteria. Vertical transmission of symbiotic bacteria between generations in sponges may occur in different ways. In the case of H. dujardini the bacteria penetrate into growing oocytes by endocytosis. A part of the bacteria plays a trophic role for oocytes and the other part remains undigested in membrane-bound vacuoles within the cytoplasm. In cleaving embryos bacteria are situated between the blastomeres or in the vacuoles. In the blastula all bacteria are disposed in the blastocoel. The symbionts are situated in intercellular spaces in free-swimming larvae and during metamorphosis. Symbiotic bacteria do not play any trophic role in the period of embryonic and postembryonic development of H. dujardini. No signs of destruction and digestion of bacteria were revealed at any stage of development.Communicated by O. Kinne, Oldendorf/Luhe     

Consequences of Buffelgrass Pasture Development for Primary Productivity,Perennial Plant Richness,and Vegetation Structure in the Drylands of Sonora,Mexico

Abstract: In large parts of northern Mexico native plant communities are being converted to non‐native buffelgrass (Pennisetum ciliare) pastures, and this conversion could fundamentally alter primary productivity and species richness. In Sonora, Mexico land conversion is occurring at a regional scale along a rainfall‐driven gradient of primary productivity, across which native plant communities transition from desert scrub to thorn scrub. We used a paired sampling design to compare a satellite‐derived index of primary productivity, richness of perennial plant species, and canopy‐height profiles of native plant communities with buffelgrass pastures. We sampled species richness across a gradient of primary productivity in desert scrub and thorn scrub vegetation to examine the influence of site productivity on the outcomes of land conversion. We also examined the influence of pasture age on species richness of perennial plants. Index values of primary productivity were lower in buffelgrass pastures than in native vegetation, which suggests a reduction in primary productivity. Land conversion reduced species richness by approximately 50% at local and regional scales, reduced tree and shrub cover by 78%, and reduced canopy height. Land conversion disproportionately reduced shrub species richness, which reflects the common practice among Sonoran ranchers of conserving certain tree and cactus species. Site productivity did not affect the outcomes of land conversion. The age of a buffelgrass pasture was unrelated to species richness within the pasture, which suggests that passive recovery of species richness to preconversion levels is unlikely. Our findings demonstrate that land conversion can result in large losses of plant species richness at local and regional scales and in substantial changes to primary productivity and vegetation structure, which casts doubt on the feasibility of restoring native plant communities without active intervention on the part of land managers.     

Aquatic plant community invasibility and scale-dependent patterns in native and invasive species richness

Invasive species richness often is negatively correlated with native species richness at the small spatial scale of sampling plots, but positively correlated in larger areas. The pattern at small scales has been interpreted as evidence that native plants can competitively exclude invasive species. Large-scale patterns have been understood to result from environmental heterogeneity, among other causes. We investigated species richness patterns among submerged and floating-leaved aquatic plants (87 native species and eight invasives) in 103 temperate lakes in Connecticut (northeastern USA) and found neither a consistently negative relationship at small (3-m2) scales, nor a positive relationship at large scales. Native species richness at sampling locations was uncorrelated with invasive species richness in 37 of the 60 lakes where invasive plants occurred; richness was negatively correlated in 16 lakes and positively correlated in seven. No correlation between native and invasive species richness was found at larger spatial scales (whole lakes and counties). Increases in richness with area were uncorrelated with abiotic heterogeneity. Logistic regression showed that the probability of occurrence of five invasive species increased in sampling locations (3 m2, n = 2980 samples) where native plants occurred, indicating that native plant species richness provided no resistance against invasion. However, the probability of three invasive species' occurrence declined as native plant density increased, indicating that density, if not species richness, provided some resistance with these species. Density had no effect on occurrence of three other invasive species. Based on these results, native species may resist invasion at small spatial scales only in communities where density is high (i.e., in communities where competition among individuals contributes to community structure). Most hydrophyte communities, however, appear to be maintained in a nonequilibrial condition by stress and/or disturbance. Therefore, most aquatic plant communities in temperate lakes are likely to be vulnerable to invasion.     

Modelling the effects of eutrophication, mitigation measures and an extreme flood event on estuarine benthic food webs

Human-mediated and natural disturbances such as nutrient enrichment, habitat modification, and flood events often result in significant shifts in species composition and abundance that translate into changes in the food web structure. Six mass-balanced models were developed using the “Ecopath with Ecosim” software package to assess changes in benthic food web properties in the Mondego estuarine ecosystem (Portugal). Field, laboratory and literature information were used to construct the models. The main study objective was to assess at 2 sites (a Zostera meadow and a bare sediment area) the effects of: (1) a period of anthropogenic enrichment, which led to excessive production of organic matter in the form of algal blooms (1993/1994); (2) the implementation of mitigation measures, following a long period of eutrophication (1999/2000); and (3) a centenary flood (winter 2000/2001). Different numbers of compartments were identified at each site and in each time period. In general, the Zostera site, due to its complex community, showed a higher number of compartments and a higher level of system activity (i.e. sum of consumptions, respiration, flow to detritus, production, total system throughput, net primary production and system omnivory index). The differences at the two sites in the three time periods in the breakdown of throughput were mainly due to differences in the biomass of the primary producers (higher primary production at the Zostera site). Consumption, respiration and flow to detritus were dominated by the grazers Hydrobia ulvae and Scrobicularia plana at the Zostera and bare sediment sites respectively. At both sites, after recovery measures were implemented there was an increase in S. plana and Hediste diversicolor biomass, consumption, respiration and flows to detritus, and a decrease in H. ulvae biomass and associated flows, which increased again after the flood event. The mass-balanced models showed that the trophic structure of the benthic communities in Mondego estuary was affected differently by each disturbance event. Interestingly, in our study a high system throughput seems to be associated with higher stress levels, which contradicts the idea that higher system activity is always a sign of healthier conditions.     

Niche overlap in mud snails (hydrobiidae): freezing tolerance

Tolerance to freezing was measured in four species of hydrobiid snails exposed to temperatures from 0° to-8°C and salinities from 0 to 30 S four up to 7 d. Experiments showed the following increasing order of tolerance to freezing: Potamopyrgus jenkinsi (Smith), Hydrobia neglecta Muus, H. ulvae (Pennant) and H. ventrosa (Montagu). Survival decreased with increasing salinity in P. jenkinsi, while the opposite was found in Hydrobia species. H. ulvae survived only slightly better than H. neglecta, while H. ventrosa was very tolerant to freezing even at low salinities in contrast to the other species of Hydrobia. H. neglecta suffered higher winter mortality than H. ventrosa during the severe winter of 1985 in an estuary.     

Batrachochytrium dendrobatidis and the Collapse of Anuran Species Richness and Abundance in the Upper Manu National Park,Southeastern Peru

Abstract: Amphibians are declining worldwide, but these declines have been particularly dramatic in tropical mountains, where high endemism and vulnerability to an introduced fungal pathogen, Batrachochytrium dendrobatidis (Bd), is associated with amphibian extinctions. We surveyed frogs in the Peruvian Andes in montane forests along a steep elevational gradient (1200–3700 m). We used visual encounter surveys to sample stream‐dwelling and arboreal species and leaf‐litter plots to sample terrestrial‐breeding species. We compared species richness and abundance among the wet seasons of 1999, 2008, and 2009. Despite similar sampling effort among years, the number of species (46 in 1999) declined by 47% between 1999 and 2008 and by 38% between 1999 and 2009. When we combined the number of species we found in 2008 and 2009, the decline from 1999 was 36%. Declines of stream‐dwelling and arboreal species (a reduction in species richness of 55%) were much greater than declines of terrestrial‐breeding species (reduction of 20% in 2008 and 24% in 2009). Similarly, abundances of stream‐dwelling and arboreal frogs were lower in the combined 2008–2009 period than in 1999, whereas densities of frogs in leaf‐litter plots did not differ among survey years. These declines may be associated with the infection of frogs with Bd. B. dendrobatidis prevalence correlated significantly with the proportion of species that were absent from the 2008 and 2009 surveys along the elevational gradient. Our results suggest Bd may have arrived at the site between 1999 and 2007, which is consistent with the hypothesis that this pathogen is spreading in epidemic waves along the Andean cordilleras. Our results also indicate a rapid decline of frog species richness and abundance in our study area, a national park that contains many endemic amphibian species and is high in amphibian species richness.     

A new apparatus for force measurement in marine bioadhesion

In an experimental study on the effect of parrotfish (probably Scarus taeniurus) grazing on the structure of benthic reef communities, fishes in densities of 0.6 to 1.5 parrotfish per m² or 9 to 17 g wet weight of fish per m² of feeding surface were found to have an optimum effect, resulting in the greatest benthic species richness and biomass on 2-dimensional surfaces. The presence of refuges (3-dimensional habitats), however, has a greater impact on bemthic community structure (number of species and biomass) than does just the density of parrotfish in such an experimental system. Coral recruitment is enhanced by the presence of refuges and, like coralline algae, is more successful under increased grazing pressure. These optimum densities of parrotfishes relate well to observed field densities where, in a collection from a Hawaiian patch reef, there were 1.1 fish or 10.8 g wet weight of parrotfish per square meter of collection area. The success of coralline algae and corals under high grazing pressure may have important consequences for the stability and structure of modern coral reefs.     

贺兰山高山草甸生物多样性和地上生物量的关系

在对贺兰山高山草甸进行群落调查的基础上,研究高山草甸生物多样性和地上生物量与环境因子之间的关系,进而分析生物多样性和地上生物量的关系.结果表明:(1)地上生物量主要与土层深度成正相关关系.(2)海拔高度与生物多样性成负相关关系,而其它影响物种丰富度或Shannon指数的环境因子仅在个别群落类型中起作用.(3)生物多样性与地上生物量主要呈单峰曲线关系.     

Allocation of nitrogen and carbon in an estuarine salt marsh in Portugal

Above and below-ground biomass and nitrogen and carbon composition ofSpartina maritima, Halimione portulacoides andArthrocnemum perenne, dominating species in plant communities of the lower, middle and higher salt marsh, respectively, were compared in an estuarine salt marsh in Portugal. Plant and soil nitrogen and carbon pools were estimated. For all three species root biomass was significantly higher (70–92% of total biomass) than above-ground biomass. The percentage of root biomass was related to the location of the plants in the marsh: higher values were found in plants growing in the lower salt marsh where the sediment was more unstable and subject to tidal action, which stresses the role of the roots as an anchor. For all three species nitrogen concentrations were highest in leaves, reflecting the photosynthetic role of the tissue. For carbon higher concentrations were found in the stems, with the exception ofS. maritima. In general, lower nitrogen concentrations were found in summer, which can be explained by dilution processes due to plant growth. For both nitrogen and carbon, higher concentrations were found in the soil surface layers. Higher soil nitrogen and carbon levels were associated with higher organic matter contents. Most of the nitrogen in the salt marsh occurred in the sediments (0–40 cm) and only ca. 5.7–13.3% of the total was found in the plants. The greater portion (76.5%–86%) of carbon was found in the sediment.     

Particle size-selection of two deposit feeders: the amphipod Corophium volutator and the prosobranch Hydrobia ulvae

The feeding biology of the deposit-feeding amphipod Corophium volutator is compared to that of the coexisting, deposit-feeding prosobranch Hydrobia ulvae. Regarding ingestion of particles, both forms show size selection which alone can explain their coexistence. Particle size-selection also explains some qualitative differences in the composition of the food of the two forms; thus, diatoms play a relatively larger role in the diet of H. ulvae than in the diet of c. volutator, whereas bacteria are probably relatively more important for the latter. Results of experiments with feeding of C-14 labelled microorganisms are in accordance with the findings on particle size-distribution of the gut contents, and show that (1) C. volutator can only utilize bacteria adsorbed to particles within the size range 4 to 63 (this is why the presence of clay and silt particles in the sediment are necessary for efficient feeding of this amphipod); (2) C. volutator can utilize bacteria suspended in the water pumped through its burrow for respiration if silt and clay particles are present in the sediment. (3) H. ulvae can utilize large particles, and also browses on surfaces, and some evidence is brought forward that it also utilizes mucus for trapping microorganisms. The coexistence of deposit-feeding animals is discussed. It is concluded that the number of coexisting, closely related species is usually small, and that their resource partitioning is probably mainly based on particle-size selectivity. In the case of unrelated forms (e.g. H. ulvae and C. volutator, a number of behavioural, physiological and morphological differences, and also the widespread ability of deposit feeders to utilize alternative feeding mechanisms may also lead to resource partitioning. Thus, there are often several niche dimensions related to feeding allowing a certain diversity of coexisting deposit feeders.     

Relearning the ABC: taxonomic changes and abundance/biomass relationships in disturbed benthic communities

For marine macrobenthic communities, a shift from higher biomass dominance with increasing levels of disturbance can be determined by the abundance/biomass comparison (ABC) method. This response results from (i) a shift in the proportions of different phyla present in communities, some phyla having larger-bodied species than others, and (ii) a shift in the relative distributions of abundance and biomass among species within the Annelida (specifically Polychaeta) but not within any of the other major phyla (Mollusca, Crustacea, Echinodermata). The shift within polychaetes reflects the substitution of largerbodied by smaller-bodied species, and not a change in the average size of individuals within a species. In most instances the phyletic changes reinforce the trend in species substitutions within the polychaetes, to produce the overall ABC response, but in some cases they may work against each other. Indications of pollution or disturbance detected by this method should be viewed with caution if the species responsible for the polluted configurations are not polychaetes. These observations provide an aid to interpretation of the ABC plots, especially in some situations where they have been deemed to give a false impression of the disturbance status of a community.     

Assessment of forest vegetation and conservation priorities of communities in part of Nanda Devi Biosphere Reserve,West Himalaya. Part I

In most protected areas of the Indian Himalayan region site/habitat characteristics, community diversity and distribution pattern, vegetation composition (richness of native and endemic species), structural patterns, economic importance of forest communities and community priorities have rarely been studied. Therefore, the present study has focused on these in the buffer zone of Nanda Devi Biosphere Reserve. Seventy-six woody species (trees: 24; shrubs: 52) and 13 forest communities have been recorded between 2300–3800 m asl. Tree density ranged from 533–1220 ind ha-¹, tree basal area from 14.68-80.28 m²ha-¹ and shrub density from 1490–6695 ind ha-¹. Mean density of trees was significantly lower in temperate forests in comparison to subalpine forests. Richness of trees ranged from 3–18 and shrubs from 5–29. Species diversity (H') of trees ranged from 0.45-2.08 and shrubs from 0.90-3.14. In the temperate zone, species richness and altitude had significant positive correlations whereas in the subalpine zone the two variables were negatively correlated. The native species were high in the area (> 65% species) and in communities (> 70% species), and was highest for the Picea smithiana-Pinus wallichiana mixed community, whereas the maximum numbers of natives and endemic species were recorded in the Pinus wallichiana community. The density and richness of non-natives were found to be significantly lower in comparison to the natives. Economic importance and conservation value of the communities were assessed and communities prioritized. Monitoring of the identified habitats, species, populations and communities, and development of appropriate strategies for their conservation and management are suggested.     

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.