Herbivore and pollinator responses to grassland management intensity along experimental changes in plant species richness

Agricultural intensification is a major driver of global environmental change. Disentangling the relative impact of losses in plant species richness and intensified management on higher trophic level organisms is important for conservation recommendations.We established different management regimes along an experimental gradient of plant species richness within “The Jena-Experiment” in Germany and quantified herbivory as well as grasshoppers and pollinators. Herbivory, grasshopper density and species richness and frequency of flower visiting pollinators were recorded two times in each of four subplots of altogether 80 plots differing in plant species richness. Each of the four subplots was subject to four different levels of fertilizer application and mowing to simulate very low, low, high and very high land use intensity.Fertilization and mowing significantly affected plant–herbivore interactions but plant species richness had no discernible effect. Grasshoppers were most abundant at high intensity subplots (3.1 individuals per m²) and least abundant on very low intensity subplots (1.3 individuals per m²). Leaf damage caused by herbivores was highest in even the very high intensity subplots (3.7%) with four mowing events per year and high amounts of fertilizer application and lowest on subplots with a low management intensity (2.4%) comprising two mowing events per year but no fertilizer application. In contrast, pollinators benefited most from lower management intensities, with only one or two mowing events and no fertilizer addition. In addition, higher numbers of flowering plant species and increased blossom cover was associated with enhanced pollinator species richness and flower visitation.Our results indicate that even in grasslands with high plant species richness, mowing and fertilization are more important drivers of herbivory and flower visitation by pollinators. Management with no more than two cuts per year and without fertilizer application in our grasslands balanced the ecosystem functions of increased pollination and decreased herbivory.     

Effect of low-intensity grazing on the species-rich vegetation of traditionally mown subalpine meadows

Subalpine meadows, which traditionally were mown every other year, are particularly rich in plant species, especially forbs. Near Davos (Switzerland) we compared the vegetation of mown sites with that of sites grazed for up to 50 years by non-lactating cows. We recorded an overall mean of 51.5 plant species per 4×4 m plot. Among grazed sites, evenness decreased with time since conversion to grazing (−0.11 in 50 years; P<0.05), suggesting progressive vegetation change, which may eventually result in the loss of species. Ground cover by forbs tended to be higher in mown than in grazed sites (by 7.2%; P<0.1). The proportion of not-clonally colonising perennial species decreased after conversion to grazing (−7.72%; in 50 years, P<0.05), while the cover by graminoid species increased (+14.2% in 50 years; P<0.1). More intensively grazed sites had a lower cover of dwarf shrubs and higher cover of legume species (P<0.05). Because grazing negatively affects both botanical richness and agricultural quality, mowing of traditionally mown subalpine meadows should be maintained, and recently grazed meadows should be reconverted to mowing.     

Nutrient impoverishment and limitation of productivity after 20 years of conservation management in wet grasslands of north-western Germany

European wet grasslands are characterized by high diversity of plant and animal species but are threatened by intensive land use. Although preservation or restoration of species-rich wet grasslands requires low nutrient availability that could be achieved by long-term management, studies monitoring nutrient removal are lacking. Our objective was to assess the long-term effect of management (mowing twice a year without or with PK fertilization for 20 years) on (i) productivity and nutrient removal with the harvest, (ii) the type of nutrient limitation, and (iii) plant species richness in wet grasslands in north-western Germany considering the differences between organic and mineral soils.Initially low nutrient availability in soil led to decreased productivity and base cation removal with harvest particularly on mineral soils after six years of mowing twice a year without fertilization. On mineral soils, N:K ratios indicated limitation of plant growth by K. On organic soils, neither productivity nor K removal with the harvest changed with time suggesting additional K input probably caused by rising groundwater. On organic soils, K:P ratios and a significant decrease of productivity with increasing N:P ratios suggested P limitation. Plant species richness was maintained or even slightly increased by mowing twice a year without fertilization but mainly comprised species that were already present at the study sites.Productivity and N, P, K, and Mg removal with the harvest was significantly increased by mowing twice a year with PK fertilization while species richness was maintained. After 10 years, N:K ratios indicate K limitation even for mowing twice a year with PK fertilization. In case of initially low nutrient availability in soil, cautious PK fertilization and mowing can be recommended to meet demands of agriculture and nature conservation.     

Soil phosphorus and tree cover modify the effects of livestock grazing on plant species richness in Australian grassy woodland

The effects of grazing on the richness of understorey plant communities are predicted to vary along gradients of resources and tree cover. In temperate Australia livestock management has involved phosphorus addition and tree removal but little research has examined how the effects of grazing on plant species richness may vary with these management regimes. Patterns of understorey plant species richness were examined in 519, 0.09 ha quadrats in grazed pastures and remnant grassy forests and woodlands in southern Australia. Sheep grazing was the primary land use and sites varied widely in grazing frequency and density, tree cover and phosphorus fertiliser history. Using an information theoretic approach the available data provides strong evidence that the effect of grazing on total species richness varies according to available phosphorus and tree cover. Intermittent grazing and no grazing were associated with high total and native plant richness, but only at low phosphorus concentrations. Phosphorus was strongly negatively correlated with richness, particularly at low grazing frequency. Total species richness was positively correlated with tree cover except under frequent grazing at high stocking rates, suggesting that heavy grazing eliminates spatial and temporal heterogeneity imposed by trees. Native plant species richness was negatively correlated with a history of cultivation, positively correlated with tree cover and varied according to landscape position and geological substrate. Frequent high density grazing, particularly when associated with clearing, cultivation and fertiliser addition, was associated with the persistence of very few native plant species. In contrast, the richness of exotic plant species was relatively invariant and performance of the best model was low. While several studies have highlighted the importance of the grazed and cleared matrix for the conservation of native plant species, this benefit may be limited in landscapes where intensive grazing management systems dominate. Strong evidence for interactions between grazing, phosphorus and tree cover suggest that failure to consider other land use practices associated with grazing management systems could lead to erroneous conclusions regarding vegetation responses to livestock grazing.     

Invasive alien goldenrods negatively affect grassland bird communities in Eastern Europe

Invasive alien plants constitute a major threat to local biodiversity. Moreover, their appearance often coincides with land use change. Many endangered groups of organisms suffer from habitat loss, which is often a consequence of the invasion of alien plants. This paper examines how invasive alien goldenrods Solidago spp. affect the richness of endangered grassland bird species and numbers of breeding pairs in abandoned meadows in southern Poland. Meadows invaded by goldenrods (n = 15) had lower bird species richness and a lower number of breeding pairs than meadows in which goldenrods were absent (n = 15). Two important factors positively influencing bird species richness were meadow area and shrub density (the latter only in meadows without goldenrods). Moreover, both the index of potential food density (number of butterflies) and plant species richness were much lower in meadows invaded by goldenrods than in meadows without these plants. Urgent action aimed at preventing the invasion of alien goldenrods in abandoned meadows is needed and such measures should involve regular mowing. Our results also highlight the fact that land abandonment in Central and Eastern Europe is not necessarily as beneficial for biodiversity as it is commonly believed and it may even lead to a decrease in the populations of several bird species in the agricultural landscape.     

Factors affecting persistence of terrestrial orchids in wet meadows and implications for their conservation in a changing agricultural landscape

We examined the occurrence of Dactylorhiza majalis, the most abundant terrestrial orchid species growing in rapidly disappearing wet meadows, at 50 historical sites for 3 years. We aimed to find the most frequent reasons for its recent extinction at many sites. We found that the main reasons for its extinction were absence of mowing, intensive fertilisation and washouts of fertilisers from fields nearby. At extant sites, we studied its biometric characteristics and composition of surrounding vegetation, to determine factors affecting its persistence. Bad performance of persisting populations of this species was associated with prevalence of grasses, low May temperatures and absence of mowing. This confirms, at metapopulation level, what has previously been observed at the level of individual populations. We suggest that the system of agricultural subsidies in the country should change towards more sensitive allocation of funds to those farmers, who will adopt the appropriate management of wet meadows and their surroundings.     

Effects of soil properties and humidity regimes on semi-natural meadow productivity

ABSTRACT

The research has been conducted for four years in five phytocenoses affected by the processes of naturalization. Two of them are located in non-flooded, relatively abandoned meadows and the remaining - flooded meadows in different parts of the Nemunas delta (Western Lithuania). The aim of the research was to estimate the influence of ecological conditions (humidity regimes, soil agrochemical parameters) on semi-natural meadow productivity. Humidity regimes and nitrogen amount in the soil influenced dry matter (DM) yield of meadow phytocesosis. The biggest DM yield (8.64 ± 0.77 t ha^-1) was found in flooded meadow in the central part of the Nemunas delta in Terric Histosol (HSs) soil, which had moderate acidity and was rich in nitrogen. The economic value depended on the prevailing plant species and was the highest in non-flooded meadows. Botanical composition and plant species number depended on agrochemical characteristics of the soil and management activities: the greatest plant species diversity and the best plant population stability were established in flooded meadow in the riverside part of the Nemunas delta behind a levee in soil rich in phosphorus and potassium and low acidity. A similar number of plant species and similar ecological stability parameters were found in non-flooded meadow, whose soil was low in nutrients, but there was intensified farming.     

The influence of plant diversity on people’s perception and aesthetic appreciation of grassland vegetation

The conservation of biodiversity critically depends on the value that humans attach to it. Apart from an ecological and economic value, an aesthetic value has often been assigned to biodiversity. However, it is not known whether lay people appreciate the diversity of species and not just certain individual species or nature as a whole. We studied in a series of experiments and field studies people’s perception and appreciation of species diversity. We presented meadow-like arrays of different species richness and evenness but random species composition to lay people and asked them to estimate plant species richness and rank the arrays by attractiveness. The experiments were complemented by two larger-scale field studies using natural meadows. Both in the experiments and the field studies the mean perception of species richness by people increased with true species richness, but was slightly overestimated at low and increasingly underestimated at high diversity levels. Lay people’s aesthetic appreciation of both the experimental grassland arrays and the natural meadows increased with true species richness. Communities consisting of the same number of species were perceived to be more species-rich and were appreciated more when their evenness was high. Our results demonstrate that plant diversity in itself is attractive to humans. The current reduction of the diversity of grasslands due to intensification of management may thus reduce the attractiveness of regions where grasslands are a dominant feature of the landscape. This could have negative consequences for tourism and may add an economic argument for the conservation of biodiversity in grasslands.     

Short-term successional change does not predict long-term conservation value of managed arable field margins

Field margins have been widely advocated as a means of integrating agronomic and biodiversity objectives and are included in agri-environment schemes across Europe. However, information on the long-term development of field margin plant communities remains limited. We describe a long-term experiment on the effects of field margin management on biodiversity and weed species. Swards were established by natural regeneration or sowing a grass and wildflower seed mixture, and treatments manipulated the frequency and timing of mowing, application of herbicide and leaving of hay. Vegetation was monitored to evaluate the extent to which early conclusions remained valid after 13 years. Although early successional trends suggested that naturally regenerated swards would rapidly become dominated by pernicious perennial weeds, and that sown swards would exclude such species, neither was true in the longer term. Sown swards were eventually invaded by unsown perennials, but they remained distinct from naturally regenerated swards. Plant species richness declined throughout the experiment. Annuals were lost most rapidly from sown swards but, under natural regeneration, loss could be modified by mowing. Perennial species initially increased during natural regeneration before stabilising. In sown swards they declined under all treatments. Species richness in naturally regenerating swards was promoted initially by mowing twice annually. After 13 years, timing and frequency of mowing had no significant effect on species richness although it still influenced sward composition. Leaving cut hay lying produced species-poor swards. We conclude that the choice of establishment and management methods for arable field margins significantly affects the long-term conservation value of the swards.     

Old cultural traditions, in addition to land use and topography, are shaping plant diversity of grasslands in the Alps

Socio-economically motivated land use changes are a major threat for species diversity of grasslands throughout the world. Here, we comprehensively explore how plant species diversity of grasslands in the species-rich cultural landscape of the Swiss Alps depends on recent land use changes, and, neglected in previous studies, on old cultural traditions. We studied diversity in 216 grassland parcels at three altitudinal levels in 12 villages of three cultural traditions (Romanic, Germanic, and Walser). In valleys of Romanic villages more different parcel types tended to occur than in those of Germanic and Walser villages, suggesting that socio-economic differences among cultural traditions still play a role in shaping landscape diversity. Moreover, at the village level, higher man-made landscape diversity was associated with higher plant species richness. All observed changes in land use reduced the farmers’ workload. Plant species richness was lower in fertilized than in unfertilized parcels and in abandoned compared with used parcels. Grazing slightly reduced species richness compared with mowing among unfertilized parcels, while in fertilized parcels it had a positive influence. The highest species diversity was found in mown unfertilized subalpine grasslands. Nevertheless, moderate grazing of former meadows can be a valuable alternative to abandonment. We conclude that the ongoing changes in land use reduce plant species richness within parcels and at the landscape level. To preserve plant species diversity at the landscape level a high diversity of land use types has to be maintained.     

An indicator species approach for result-orientated subsidies of ecological services in grasslands - A study in Northwestern Germany

Agri-environmental subsidies for grasslands of high conservation interest are usually associated with fairly strict management rules, but have not always been successful in terms of the preservation of species. A new approach now links the subsidies paid to the ecological value of a grassland site: the farmers are paid for keeping up a high plant species richness, and they can control the species richness of their sites by using vascular plants as indicator species. The method was tested in different grassland communities in six regions of Lower Saxony in Northwestern Germany, mainly addressing the question whether the number of selected indicator species was correlated with the total species richness of vascular plants and with the number of endangered species.Field work was carried out in 2004. Total species richness of vascular plants was recorded in all grassland sites, and the number of indicators was counted along two transects, usually being the diagonals of a rectangular site. Each transect was divided into three segments in which all indicators growing within 1 m distance from the transect line were recorded. In total 43 indicators were selected, mostly representing species that are fairly easy to recognise.The number of indicators was significantly positively correlated with the total number of species in all regions, also when correcting for autocorrelation (except in one region). When using grassland area as a co-variable, the same results were obtained, supporting the low significance of grassland size for the predictive ability of the method. In all regions but one, the mean number of indicators per segment was also significantly positively related to the number of red-listed and near-threatened species in the grasslands. Grassland size and transect length were found to be of minor importance for the application of the method.In conclusion, the study shows that the indicator approach is suitable for identifying those grassland sites that have a high plant species richness and a high number of endangered species. It may thus form the basis for result-orientated subsidies in grasslands.     

The impact of mowing as a management strategy for wet meadows on spider (Araneae) communities

Meadows are often managed by mowing to prevent succession and brushwood encroachment. This management practice is efficient to maintain plant diversity, but the effect on arthropod groups is less well known. We sampled spiders in the Grande Cariçaie (Switzerland), a site of nature conservation importance, in two types of wet meadows, an unmanaged site and one 2-years-old mown conditions. Numbers of individuals for the most abundant families, diversity measures, and indicator taxa were compared among vegetation and treatment types. The results indicate that the less mobile spiders and species linked to litter or dead reeds, including rare species, are reduced by mowing. Present management consists in triennial mowing of 2-4 ha non-contiguous sectors. We investigate the conflict between the need to mow the meadows to maintain them, and the negative effect of mowing on spider communities. We therefore propose a new management scheme aimed at maintain the vegetation while lessening its negative effects on spider communities by providing refuges.     

Plant species richness and composition in managed grasslands: The relative importance of field management and environmental factors

Decline of grassland diversity throughout Europe within the last decades is threatening biological diversity and is a major conservation problem. There is an urgent need to determine the underlying factors that control vascular plant species richness and composition in managed grasslands. In this study, 117 grasslands were sampled using standardised methods. Explanatory variables were recorded for each grassland site, reflecting the local field management, site-specific environmental conditions and large-scale spatial trends. Using variation partitioning methods, we determined the pure and shared effects of these three sets of explanatory variables on the plant species richness and composition in grasslands. Most of the explained variation in plant species richness was related to the joint effect of local field management and environmental variables. However, the applied variation partitioning approach revealed that the pure effect of spatial variables contributed relatively little to explaining variation in both the plant species richness and species composition. The largest fractions of explained variation in plant species composition were accounted for by the pure effects of environmental and local field management variables. Moreover, the results revealed that the main mechanisms by which these sets of explanatory variables affect plant species vary according to the type of management regime under study. From our findings we could conclude that particularly a reduction of nitrogen fertilisation on meadows and grazing at a low stocking rate on pastures can help to conserve biodiversity.     

Effects of soil properties and land management on the structure of grassland snail assemblages in SW Germany

Quantitative analyses of the snail assemblages of four a priori defined categories of grassland sites, characterized by soil moisture, land use and management, were conducted in SW Germany. Topsoil pH and calcium content were determined for each of 39 sites. The objectives were to study the effects of these habitat characteristics on snail density, species richness and quantitative structure of the land snail assemblages. With one exception, no significant relationships between soil pH (as a proxy for Ca supply) and the total numbers of individuals and species at the four site categories were observed. Snail density and species richness on groundwater soils, represented by rarely mown sedge stands and regularly mown meadows, were clearly higher than on intermediate moisture soils (as defined by the climatic water balance) of regularly mown meadows, indicating that soil moisture was the most important factor accounting for these differences. Furthermore, we observed specific conditions associated with the presence of trees in the category of meadow orchards on intermediate moisture soils. There, numbers of individuals and species were significantly higher than in the meadows on intermediate moisture soils. A posteriori cluster analysis of quantitative similarity of all snail assemblages showed that soil moisture accounts for the major differences between the site categories with respect to snail density, species richness and quantitative composition of the grassland snail assemblages. As these results are in contrast to the findings of many woodland studies, we suggest that the occurrence or abundance of certain species in the site categories are explained by land management, especially by frequent disturbances at the soil surface caused by mowing.     

盐碱草甸植被退化对土壤硝化作用强度的影响

为了解不同退化阶段盐碱草甸草原土壤硝化作用强度特征及其影响因素,采用空间代替时间的方法,以松嫩平原盐碱草甸草原植被退化过程中4种典型植物群落为对象,以未做处理为参照,设置刈割、施氮和刈割同时施氮3种处理,测定了土壤的硝化作用强度(NI)、pH值、电导率(EC)、含水量(SMC)和有效磷(OP)、硝态氮(NO^-₃-N)、铵态氮(NH+4-N)及全氮(TN)的含量。结果显示:(1)土壤NI与pH值、电导率、含水量、OP和NO^-₃-N呈极显著正相关关系(p<0.01);通过逐步回归分析的方法得出土壤NI的重要影响因子,重要影响因子对土壤NI影响强弱表现为:pH值>有效磷含量>含水量>硝态氮含量,并推断土壤硝态氮含量可作为土壤NI的一个重要表征参数。(2)未作处理时,星星草群落与碱蓬群落土壤硝化作用强度分别为13.4,13.5 mg/(kg·h),显著高于羊草群落和退化羊草群落的5.0,2.5 mg/(kg·h),刈割和施氮处理分别使星星草群落土壤NI提高96.88%,253.77%,混合处理使其提高413.70%,显著高于其他3种植物群落,由此可见,在人为刈割和施氮肥的干预下,星星草群落土壤铵态氮可能更易转变为硝态氮,氮素流失的风险也更大,因此可认为星星草群落处于盐碱草甸退化过程中的关键阶段。     

高寒草甸不同类型草地土壤养分与物种多样性——生产力关系

研究了高寒草甸不同类型草地土壤养分与多样性—生产力之间的关系,即物种多样性对生产力的效应如何受到资源供给率等因素的影响。结果表明:以莎草类为优势种的藏嵩草沼泽化草甸群落其总生物量(包括地上和地下生物量)最高(13,196.96±719.69gm-2)、小嵩草草甸和金露梅灌丛群落为中等水平(2,869.58±147.52gm-2、2,672.94±122.49gm-2)、矮嵩草草甸群落为最低(2,153.08±141.95gm-2)。在藏嵩草沼泽化草甸群落中,总生物量和物种丰富度呈显著负相关(P<0.05);地上生物量与土壤有机质、土壤含水量和群落盖度显著正相关(P<0.05);地下生物量和土壤含水量显著正相关(P<0.05)。在矮嵩草草甸、小嵩草草甸、金露梅灌丛群落中,地上生物量与土壤有机质和土壤总氮显著正相关(P<0.05)。以上结果说明生物量的分布与土壤营养和水分变化相一致。在矮嵩草草甸、小嵩草草甸和金露梅灌丛中,多样性有随土壤养分的增加而增加的趋势;在藏嵩草沼泽化草甸中,则呈现负相关的关系。     

Subsequent nitrogen utilisation and soil water distribution as affected by forage radish cover crop and nitrogen fertiliser in a corn silage production system

Forage radish is a unique winter cover crop that is relatively new but becoming rapidly adopted in temperate, humid North America. Little is known about how the use of this cover crop may influence subsequent nitrogen availability, soil water accumulation in the soil profile in corn silage production system. In this present work, the average nitrogen uptake by silage corn increased significantly by 11.6% in cover plots compared with the no-cover control plots. The recovery efficiency and agronomic efficiency of applied nitrogen in silage corn declined in cover plots compared to no-cover plots. The average soil water storage in cover plots was significantly higher than in the control after corn sowing and at the harvest stage. With increasing nitrogen application level, the average corn grain yield increased significantly at 56 and 112 kg N ha^?1 by 13.1% and 39.8%, respectively. Planting a forage radish cover crop can facilitate growth of silage corn and markedly improve total nitrogen uptake of corn. Consideration should be given to nitrogen application rate and also to avoiding excessive nitrogen input in the subsequent crop following a cover crop, thereby truly improving subsequent fertiliser use efficiency.     

Neglect of mowing and manuring leads to slower nitrogen cycling in subalpine grasslands

Nitrogen (N) availability in grasslands varies with agricultural land use. Traditional management regimes of mowing for hay and manuring in subalpine meadows maintain plant communities with exploitative functional strategies suited to fertile soils with fast turnover of nutrients. We investigated whether the neglect of traditional practices has led to a reduction in N availability in two parallel ecosystems (terraced and unterraced fields). Soil nitrate and ammonium contents were assessed using soil cores and ion exchange resins over a 1-year period, and assays of microbial nitrifying and denitrifying enzyme activities, made early in the growing season. A large difference in pH between the two ecosystems, caused by historical ploughing, facilitated greater N availability in terraced than unterraced fields. Abandonment of manuring and mowing caused a reduction in N availability and N transformation processes, which correlated with a shift in the plant community towards more-conservative functional strategies and greater dominance by grasses. We propose that positive feedback between the grassland management regime and dominant plant functional strategy maintained high N availability in these managed subalpine grasslands. When traditional practices of mowing and manuring are neglected, direct management effects combined with the spread of grass species with conservative strategies force down N availability in the soil, reduce microbial activity, change the pH, and lead to a long-term loss of characteristic herbaceous subalpine-meadow species.     

Optimizing grassland management for flower-visiting insects in roadside verges

The decline of flower-visiting insects is a threat to ecological processes and to the services these insects provide. Roadside verges in the Netherlands span approximately 80,000 km and are often covered with semi-natural grasslands. As such, they also provide a suitable habitat for many insects, but this has received little attention so far. We investigated the effects of different management treatments on flower-visiting insects. We studied flower visitation in a 3 years old experimental set-up with five mowing treatments each replicated five times. Management types were: no management and mowing once or twice per year with and without the removal of hay, representing common forms of management and neglect. During an entire growing season, both flowers (number of species and inflorescences) as well as insects (total abundance and actual flower visits) were investigated. Mowing twice per year with removal of hay showed highest values for all measured variables and this effect persisted throughout the growing season. The early summer cut proved to be very important for insect feeding opportunities, due to the re-flowering of plants later in the growing season. Flower abundance showed high correlations with both plant species richness and the number of insect visits. Although overall, mowing twice a year with hay removal was the most beneficial treatment for flower-visiting insects, these plots were entirely devoid of flowers for some period right after mowing, indicating that a rotational scheme might further promote insect diversity and abundance.     

Plant community functional shifts in response to livestock grazing in intermountain depressional wetlands in British Columbia, Canada

Wetlands are ecologically and economically important ecosystems with high conservation value. Although wetland vegetation is strongly determined by abiotic factors, grazing disturbance may also be an important influence on this community. We evaluated the effects of livestock grazing on wetland vegetation in marsh and wet meadow zones in intermountain depressional wetlands in the southern interior of British Columbia, Canada. We sampled marshes and wet meadows in 36 wetlands along a grazing intensity gradient ranging from fully fenced and ungrazed wetlands to unfenced wetlands heavily grazed by livestock. The amount of bare ground was used as a surrogate measure of the intensity of livestock grazing. Vegetation community structure and composition was strongly associated with grazing intensity. Increased livestock grazing favored shorter-lived and smaller plants; conversely, the frequency of tall and rhizomatous species, which constitute the dominant plant species in these systems, declined with higher livestock use. The effects of grazing were more pronounced in the marsh than in the wet meadow. Associations between species richness and grazing differed between zones: native and exotic species richness showed a unimodal response in the marsh while in the wet meadow exotic richness increased and native richness showed no response. The relationship between exotic frequency and grazing was also inconsistent between zones, with a negative association in the marsh and a positive but weak association in the wet meadow. Grazing-related changes in vegetation along the grazing intensity gradient were substantial and may affect the habitat value of these wetlands for dependent wildlife such as breeding waterfowl.