After the hotspots are gone: Land use history and grassland plant species diversity in a strongly transformed agricultural landscape

Question: We asked how landscape configuration and present management influence plant species richness and abundance of habitat specialists in grasslands in a ‘modern’(much exploited and transformed) agricultural Swedish landscape. Location: Selaön, south‐eastern Sweden (59°24’ N, 17°10’ E). Methods: Present and past (150 and 50 years ago) landscape pattern was analysed in a 25 km² area. Species richness was investigated in 63 different grassland patches; grazed and abandoned semi‐natural grasslands, and grazed ex‐arable fields. Influence of landscape variables; area, past and present grassland connectivity, present management on total species richness, density and abundance of 25 grassland specialists was analysed. Results: Semi‐natural grasslands (permanent unfertilised pastures or meadows formed by traditional agricultural methods) had declined from 60% 150 years ago to 5% today. There was a significant decline in species richness and density in abandoned semi‐natural grasslands. Total species richness was influenced by present management, size and connectivity to present and past grassland pattern. Landscape variables did not influence species density in grazed semi‐natural grassland suggesting that maintained grazing management makes grassland patches independent of landscape context. The abundance of 16 grassland specialists was mainly influenced by management and to some extent also by landscape variables. Conclusion: Although species richness pattern reflect management and to some extent landscape variables, the response of individual species may be idiosyncratic. The historical signal from past landscapes is weak on present‐day species richness in highly transformed, agricultural landscapes. Generalizations of historical legacies on species diversity in grasslands should consider also highly transformed landscapes and not only landscapes with a high amount of diversity hotspots left.     

Recreating Grasslands in Swedish Rural Landscapes – Effects of Seed Sowing and Management History

Recent loss of plant species richness in Swedish semi-natural grasslands has led to an increase in grassland recreation and restoration. To increase the establishment of declining species favoured by grazing and to re-establish original species richness, seed sowing has been discussed as a conservation tool. In this study, I examined to what extent seed sowing in former arable fields increases species richness and generates a species composition typical of semi-natural grasslands. Six grassland species favoured by grazing (target species) and six generalist species favoured by ceased grazing, were studied in a seed-addition experiment. Four different seed densities were used on four different grassland categories, two grazed former arable fields, one continuously grazed grassland and one abandoned grassland. Target and generalist species emerged in all grassland categories, but seedling emergence was higher in the grazed than in the abandoned grassland. Target species had higher emergence in the two grasslands with the longest grazing continuity. Seedling emergence and frequency of established plants of each target species were positively associated. The largest fraction of seeds germinated at an intermediate sowing density, 20–50 seeds/dm², suggesting that aggregation of seeds positively affects emergence up to a certain threshold. In conclusion, artificial seed sowing may induce the recreation of typical grassland communities on former arable fields, which may be an important contribution to increase the total grassland area and species richness in the landscape.     

Effects of Restoration on Plant Species Richness and Composition in Scandinavian Semi-Natural Grasslands

Plant species richness in rural landscapes of northern Europe has been positively influenced by traditional management for millennia. Owing to abandonment of these practices, the number of species‐rich semi‐natural grasslands has decreased, and remaining habitats suffer from deterioration, fragmentation, and plant species decline. To prevent further extinctions, restoration efforts have increased during the last decades, by reintroducing grazing in former semi‐natural grasslands. To assess the ecological factors that might influence the outcome of such restorations, we made a survey of semi‐natural grasslands in Sweden that have been restored during the last decade. We investigated how plant species richness, species density, species composition, and abundance of 10 species that are indicators of grazing are affected by (1) the size of the restored site, (2) the time between abandonment of grazing and restoration, (3) the time elapsed since restoration, and (4) the abundance of trees and shrubs at the restored site. Only two factors, abundance of trees and shrubs and time since restoration, were positively associated with total species richness and species density per meter square at restored sites. Variation in species composition among restored sites was not related to any of the investigated factors. Species composition was relatively similar among sites, except in mesic/wet grasslands. The investigated factors had small effects on the abundance of the grazing‐indicator species. Only Campanula rotundifolia responded to restoration with increasing abundance and may thus be a suitable indicator of improved habitat quality. In conclusion, positive effects on species richness may appear relatively soon after restoration, but rare, short‐lived species are still absent. Therefore, remnant populations in surrounding areas may be important in fully recreating former species richness and composition.     

Does resource availability,resource heterogeneity or species turnover mediate changes in plant species richness in grazed grasslands?

Grazing by large ungulates often increases plant species richness in grasslands of moderate to high productivity. In a mesic North American grassland with and without the presence of bison (Bos bison), a native ungulate grazer, three non-exclusive hypotheses for increased plant species richness in grazed grasslands were evaluated: (1) bison grazing enhances levels of resource (light and N) availability, enabling species that depend on higher resource availability to co-occur; (2) spatial heterogeneity in resource availability is enhanced by bison, enabling coexistence of a greater number of plant species; (3) increased species turnover (i.e. increased species colonization and establishment) in grazed grassland is associated with enhanced plant species richness. We measured availability and spatial heterogeneity in light, water and N, and calculated species turnover from long-term data in grazed and ungrazed sites in a North American tallgrass prairie. Both regression and path analyses were performed to evaluate the potential of the three hypothesized mechanisms to explain observed patterns of plant species richness under field conditions. Experimental grazing by bison increased plant species richness by 25% over an 8-year period. Neither heterogeneity nor absolute levels of soil water or available N were related to patterns of species richness in grazed and ungrazed sites. However, high spatial heterogeneity in light and higher rates of species turnover were both strongly related to increases in plant species richness in grazed areas. This suggests that creation of a mosaic of patches with high and low biomass (the primary determinant of light availability in mesic grasslands) and promotion of a dynamic species pool are the most important mechanisms by which grazers affect species richness in high productivity grasslands.     

Size and heterogeneity rather than landscape context determine plant species richness in semi‐natural grasslands

Question: Is plant diversity in fragmented semi‐natural grasslands related to present and historical landscape context? Location: Southern Sweden. Methods: Plant diversity was described at 30 semi‐natural grassland sites in terms of total and specialist plant species richness at the site and species density at different scales (0.5–10 m²). These measures are commonly used to assess conservation value of semi‐natural grasslands. Landscape context was measured as contemporary connectivity to other semi‐natural grasslands, historical connectivity 50 years ago, amount of linear elements potentially suitable for dispersal (road verges, power line clearings), and amount of forest (inverse of the openness of the landscape). Results: The diversity measures were generally correlated with each other, implying that species richness in a subset of the grassland can predict the total richness. Plant species density at three scales (0.5 m², 10 m² and total) was related to the landscape context using an information theoretic approach. Results showed that total species richness increased with increased size of grasslands, contrary to earlier diversity studies in semi‐natural grasslands. Larger grasslands were more heterogeneous than smaller grasslands, and this is a likely reason for the species‐area relationship. Heterogeneity was also of high importance at the smaller scales (0.5 m², 10 m²). With increased amount of forest, total species richness increased but species density on 10 m² decreased. There was no influence of connectivity in either the contemporary or the historical landscape, contrary to previous studies. Conclusions: Grassland size and heterogeneity are of greater importance for plant diversity in semi‐natural grassland, than grassland connectivity in the landscape.     

Importance of grazing and soil acidity for plant community composition and trait characterisation in coastal dune grasslands

Question: Does grazing by large herbivores affect species composition or community‐wide variation in plant functional traits? Location: Dune grasslands at the Belgian coast. Methods: Plant cover and soil data were collected in 146 plots that were randomly selected at 26 grazed and ungrazed grassland sites. Plant community composition was assessed by Detrended Correspondence Analysis and mean values of plant trait categories were calculated across the plots. Results: Differentiation of plant composition and community‐wide plant trait characteristics was largely determined by grazing, soil acidity and their interaction. In ungrazed situations, a clear floristic distinction appears between acidic (non‐calcareous) and alkaline (calcareous) grasslands. In grazed situations, these floristic differences largely disappeared, indicating that grazing results in a decrease of natural variation in species composition. At higher soil pH, a larger difference in plant community composition and community‐wide plant traits was observed between grazed and ungrazed plots. In ungrazed situations, shifts in plant functional traits along the acidity gradient were observed. Conclusions: Grazing is responsible for shifts in plant community composition, and hence a decrease in plant diversity among grasslands at opposing acidity conditions in coastal dune grasslands. Therefore, care should be taken when introducing grazing as a system approach for nature conservation in dune grasslands as it may eliminate part of the natural variation in plant diversity along existing abiotic gradients.     

Effect of Hay Transfer on Long-Term Establishment of Vegetation and Grasshoppers on Former Arable Fields

The transfer of seed‐containing hay is a restoration measure for the introduction of plant species of local provenance. We investigated the effect of hay transfer on species richness and on long‐term establishment of target plant and grasshopper species on former arable fields with and without topsoil removal in comparison to reference sites in a nature reserve. Plant species richness, the number of target plant species, and Red List plant species were significantly positively affected by hay transfer, both on the scale of whole restoration fields and on permanent plots of 4 m². Eight years after the start of the restoration, only few of the transferred plant species had disappeared and some target species were newly found. Grasshoppers were affected not by hay transfer but by topsoil removal. The proportion of target grasshopper and plant species and Red List grasshopper species was higher on topsoil removal sites with low standing crop and high cover of bare soil than on sites without soil removal. On topsoil removal sites without hay, however, plant species richness was very low because of the slow natural dispersal of the target species. Vegetation and grasshopper communities still differed between restoration fields and the nature reserve. Nevertheless, our results indicate that the transfer of autochthonous seed‐containing hay is a successful method to establish species‐rich grasslands with a high proportion of target species.     

Problems, Approaches, and Results in Restoration of Dutch Calcareous Grassland During the Last 30 Years

This paper is based on research of the restoration of species‐rich calcareous grasslands in The Netherlands, over the last 30 years. Chalk grassland is a semi‐natural vegetation with a high density of species at a small scale. This type of vegetation was once widespread in Western Europe as common grazing land, mainly for flocks of sheep for which the main function was dung production. In some regions of Central Europe, these grasslands were also used for hay production. The dung was used to maintain arable field production at a reasonable level. In the chalk district in the southernmost part of The Netherlands some 25 sites of this vegetation, varying in area from 0.05–4.5 ha, are still present. Chalk grassland completely lost its significance for modern agricultural production after the wide application of artificial fertilizer following World War II. This grassland has a high conservation value both for plants and animal species, of which a large number of species are exclusively restricted to this biotope. When conservation activities started at a large scale in the early 1960s, three different types of restoration activities could be distinguished: (1) restoration of fertilized sites; (2) restoration of abandoned grasslands; and (3) recreation of chalk grassland on former arable fields. The main aim of the restoration attempt is to create and/or improve sustainable conditions for both plant and animal species characteristic of the chalk grassland ecosystem. In the process of restoration, several phases of different activities can be distinguished: (1) pre‐restoration phase, during which information of the land use history is collected and, based on these data, clear restoration goals are established; (2) initial restoration phase, during which effects of former, non‐conservational land use has to be undone in order to stimulate germination and establishment of target species originating from soil seed bank and species pool; (3) consolidation phase, including the introduction and continuation of a regular management system for sustainable conservation; and (4) long‐term conservation strategy, including measures to prevent disturbance from the outside and genetic erosion and extinction of locally endangered plant populations.     

From plant neighbourhood to landscape scales: how grazing modifies native and exotic plant species richness in grassland

The interactive effect of grazing and soil resources on plant species richness and coexistence has been predicted to vary across spatial scales. When resources are not limiting, grazing should reduce competitive effects and increase colonisation and richness at fine scales. However, at broad scales richness is predicted to decline due to loss of grazing intolerant species. We examined these hypotheses in grasslands of southern Australia that varied in resources and ungulate grazing intensity since farming commenced 170 years ago. Fine-scale species richness was slightly greater in more intensively grazed upper slope sites with high nutrients but low water supply compared to those that were moderately grazed, largely due to a greater abundance of exotic species. At broader scales, exotic species richness declined with increasing grazing intensity whether nutrients or water supply were low or high. Native species richness declined at all scales in response to increasing grazing intensity and greater resource supply. Grazing also reduced fine-scale heterogeneity in native species richness and although exotics were also characterised by greater heterogeneity at fine scales, grazing effects varied across scales. In these grasslands patterns of plant species richness did not match predictions at all scales and this is likely to be due to differing responses of native and exotic species and their relative abundance in the regional species pool. Over the past 170 years intolerant native species have been eliminated from areas that are continually and heavily grazed, whereas transient, light grazing increases richness of both exotics and natives. The results support the observation that the processes and scales at which they operate differ between coevolved ungulate—grassland systems and those in transition due to recent invasion of herbivores and associated plant species.     

Interacting effects of fertilization,mowing and grazing on plant species diversity of 1500 grasslands in Germany differ between regions

The relationship of different types of grassland use with plant species richness and composition (functional groups of herbs, legumes, and grasses) has so far been studied at small regional scales or comprising only few components of land use. We comprehensively studied the relationship between abandonment, fertilization, mowing intensity, and grazing by different livestock types on plant diversity and composition of 1514 grassland sites in three regions in North-East, Central and South-West Germany. We further considered environmental site conditions including soil type and topographical situation. Fertilized grasslands showed clearly reduced plant species diversity (?15% plant species richness, ?0.1 Shannon diversity on fertilized grasslands plots of 16 m²) and changed composition (?3% proportion of herb species), grazing had the second largest effects and mowing the smallest ones. Among the grazed sites, the ones grazed by sheep had higher than average species richness (+27%), and the cattle grazed ones lower (?42%). Further, these general results were strongly modulated by interactions between the different components of land use and by regional context: land-use effects differed largely in size and sometimes even in direction between regions. This highlights the importance of comparing different regions and to involve a large number of plots when studying relationships between land use and plant diversity. Overall, our results show that great caution is necessary when extrapolating results and management recommendations to other regions.     

Floristic diversity of meadow steppes in the Western Siberian Plain: effects of abiotic site conditions,management and landscape structure

Temperate grasslands have suffered from severe habitat loss and degradation worldwide. In Russia, vast areas of forest-steppe grasslands have been converted to cropland during Soviet times, whilst remaining grasslands were often intensively grazed. Contrastingly, the collapse of the Soviet Union have resulted in a massive reduction in livestock numbers and cessation of management. Albeit relatively large natural grassland areas remained in the Western Siberian Plain, their present condition is poorly studied. We analysed plant species composition, functional structure and richness of grassland communities and tested for the effect of local factors (management, abiotic site conditions) and landscape factors (patch size, proportion of land cover types) on diversity patterns. Abiotic site conditions, mainly soil moisture and salinity, differentiated distinct community types. Overall, species richness was highest in meadow steppe communities with lower soil moisture and salinity. Grazing intensity and litter accumulation due to cessation of management were significant negative related to species richness and shaped the functional structure. At the landscape scale, diversity in meadow steppe grasslands was higher in forest-grassland mosaics and in small remnants isolated in a matrix of cropland. Our findings highlight that meadow steppes suffered massively under the historical habitat loss and high grazing pressure. Small species-rich remnants are evidence of the former extent of meadow steppe habitats in agricultural landscape, but are likely threatened by an extinction debt. Low intense, irregular mowing maintained species-rich meadow steppe in forest- grassland mosaics, but currently such practices are declining.     

Use of sheep grazing in the restoration of semi‐natural meadows in northern Finland

Abstract. The biodiversity of species‐rich semi‐natural meadows is declining across Europe due to ceased management. In this study we aimed to find out how successfully the local species richness of an overgrown semi‐natural mesic meadow could be restored by sheep grazing after a long period of abandonment. The cover of vascular plant species in grazed plots and ungrazed exclosures was studied for five years and the responses of different functional plant groups were followed (herbs vs grasses, tall vs short species, species differing in flowering time, species representing different Grime's CSR strategies and species indicative of rich vs poor soil). Grazing increased species number by nearly 30%. On grazed plots the litter cover practically disappeared, favouring small herbs such as Rhinanthus minor, Ranunculus acris, Trifolium pratense and the grass Agrostis capillaris. Grazing decreased the cover of the late flowering tall herb Epilobium angustifolium but had no effect on the abundance of the early flowering tall herbs Anthriscus sylvestris or Geranium sylvaticum. We suggest that to succeed in restoration it is useful to determine the responses of different functional plant groups to grazing. Grassland managers need this information to optimize the methods and timing of management used in restoration. Additional management practices, such as mowing, may be needed in mesic meadows to decrease the dominance of tall species. The availability of propagules seemed to restrict further increase of species richness in our study area.     

The conservation benefit of mowing vs grazing for management of species‐rich grasslands: a multi‐site,multi‐year field experiment

Species‐rich semi‐natural grasslands in Europe are becoming more fragmented and many species that depend on this habitat type are rare and threatened today. Management methods like mowing and grazing are needed to preserve remaining grasslands. Because management is costly it is important to use the most cost‐effective as well as the most beneficial management method, but few studies have compared mowing and grazing. We investigated the effect of mowing and grazing on grassland vegetation using data from 11 long‐term field trials situated in southern Sweden. We calculated the change in the odds of finding species belonging to three different groups of indicators at the start of the treatment and after 8 and 14 years. The used indicator groups were indicators of good management, excess nitrogen and poor management. The results revealed an increase in the odds of finding indicators of good management in mowed plots and an increase in finding indicators of excess nitrogen in grazed plots. The odds of finding indicators of poor management remained unchanged. Results from sub‐analysis of the grazing intensity showed a more negative effect from grazing with low grazing intensity than normal/high grazing intensity. Therefore, mowing is the best long‐term management method for semi‐natural grasslands in Sweden and grazing using a low grazing intensity should be avoided.     

How can we preserve and restore species richness of pollinating insects on agricultural land?

During recent decades, concern about the loss of biodiversity on agricultural land has increased, and semi‐natural grasslands have been highlighted as critical habitats. Temperate European agricultural landscapes require distinct and appropriate management to prevent further impoverishment of the flora and fauna. This is especially urgent for pollinating insects that provide important ecosystem services. Our aim was to examine how species richness of three important groups of pollinating insects; solitary bees, butterflies and burnet moths are related to different farm characteristics, and if there are any differences between these three groups. A further aim was to test if red‐listed species are related to any farm characteristics. Species richness of solitary bees, butterflies and burnets was measured on all semi‐natural grasslands at 16 farms in a forest‐dominated area of 50 km² in southern Sweden, using systematic transect walks in April to September 2003 (only butterflies and burnets) and 2005. Species richness of solitary bees and butterflies was intercorrelated, both before and after controlling for the area of semi‐natural grassland. Species richness of solitary bees increased with the area of semi‐natural grassland. After controlling for the effect of the area of semi‐natural grassland species richness was strongly positively related with the density of the plant Knautia arvensis and negatively related with the proportion of grazed grassland. The results were similar for solitary bees and butterflies. The number of red‐listed solitary bees was positively related to the proportion of meadows with late harvest (after mid‐July) and decreased with increased farm isolation. The number of burnet species (all red‐listed) was positively related to vegetation height, flower density and the proportion of meadows with late harvest on a farm. Areas with a high density of K. arvensis and with traditional hay‐meadow with late harvest present, harbour most species. Promoting traditional hay‐meadows, late extensive grazing and the herb K. arvensis, people managing agricultural biodiversity can encompass high species richness of pollinating insects and support red‐listed species. Further, we suggest that the density of K. arvensis at a farm can be used as a biodiversity indicator, at least for pollinating insects.     

An attempt to restore a central European species‐rich mountain grassland through grazing

Abstract. This paper describes the effects of re‐establishing seasonal cattle grazing by 0.7 animal.ha^‐1 on vegetation in a long‐term abandoned, and partly degraded, semi‐natural mountain pasture in the ?umava National Park, Czech Republic. There was very uneven grazing intensity inside the locality, and grazing preference changed during the season: cattle grazed most of the time in productive but species‐poor Deschampsia cespitosa swards, but changed to a species‐rich Violion caninae stand in the middle of the summer. A species‐rich Carex rostrata community was only grazed at the end of the season. Species‐poor swards dominated by Nardus stricta and Carex brizoides were mainly used as resting areas. Both grazing and excluding from grazing had a negative effect on species diversity of the Deschampsia cespitosa swards. The soil seed bank contained only few species that are characteristic of mountain grassland communities, and seed dispersal of the target species by cattle dung was also found to be very limited. Thus both grazing and exclusion from grazing are probably of limited value for the restoration of species‐rich grasslands from species‐poor Deschampsia cespitosa swards in this case.     

Resilience of a high‐conservation‐value,semi‐arid grassland on fertile clay soils to burning,mowing and ploughing

In grassland reserves, managed disturbance is often necessary to maintain plant species diversity. We carried out experiments to determine the impact of fire, kangaroo grazing, mowing and disc ploughing on grassland species richness and composition in a nature reserve in semi‐arid eastern Australia. Vegetation response was influenced by winter–spring drought after establishment of the experiments, but moderate rainfall followed in late summer–autumn. Species composition varied greatly between sampling times, and the variability due to rainfall differences between seasons and years was greater than the effects of fire, kangaroo grazing, mowing or disc ploughing. In the fire experiment, species richness and composition recovered more rapidly after spring than autumn burning. Species richness and composition were similar to control sites within 12 months of burning and mowing, suggesting that removal of the dominant grass canopy is unnecessary to enhance plant diversity. Two fires (separated by 3 years) and post‐fire kangaroo grazing had only minor influence on species richness and composition. Even disc ploughing caused only a small reduction in native richness. The minor impact of ploughing was explained by the small areas that were ploughed, the once‐off nature of the treatment, and the high degree of natural movement and cracking in these shrink‐swell soils. Recovery of the composition and richness of these grasslands was rapid because of the high proportion of perennial species that resprout vegetatively after fire and mowing. There appears to be little conservation benefit from fire, mowing or ploughing ungrazed areas, as we could identify no native plant species dependent on frequent disturbance for persistence in this grassland community. However, the ability of the Astrebla‐ and Dichanthium‐dominated grasslands to recover quickly after disturbance, given favourable seasonal conditions, suggests that they are well adapted to natural disturbances (e.g. droughts, fire, flooding and native grazing).     

Scale‐dependence of vegetation‐environment relationships in semi‐natural grasslands

Questions: Which environmental and management factors determine plant species composition in semi‐natural grasslands within a local study area? Are vegetation and explanatory factors scale‐dependent? Location: Semi‐natural grasslands in Lærdal, Sognog Fjordane County, western Norway. Methods: We recorded plant species composition and explanatory variables in six grassland sites using a hierarchically nested sampling design with three levels: plots randomly placed within blocks selected within sites. We evaluated vegetation‐environment relationships at all three levels by means of DCA ordination and split‐plot GLM analyses. Results: The most important complex gradient determining variation in grassland species composition showed a broad‐scale relationship with management. Soil moisture conditions were related to vegetation variation on block scale, whereas element concentrations in the soil were significantly related to variation in species composition on all spatial scales. Our results show that vegetation‐environment relationships are dependent on the scale of observation. We suggest that scale‐related (and therefore methodological) issues may explain the wide range of vegetation‐environment relationships reported in the literature, for semi‐natural grassland in particular but also for other ecosystems. Conclusions: Interpretation of the variation in species composition of semi‐natural grasslands requires consideration of the spatial scales on which important environmental variables vary.     

Remnant habitats for grassland species in an abandoned Swedish agricultural landscape

Questions: Which factors influence the persistence of vascular grassland plants in long‐abandoned (at least 50 yr) arable fields and meadows? What might be the implications of current levels of species richness on abandoned arable fields and meadows for future restoration? Location: Forested highlands of Kilsbergen, south central Sweden. Methods: The abundance of all vascular plant species was investigated in three habitat types: former arable fields, hay meadows and outlands (pastures) at 27 farms, abandoned for either approximately 50 yr or 90 yr. Time since abandonment, tree cover, soil depth, degree of soil podsol development, size of the infield area and two measures of connectivity were used as predictors for species richness and species composition. Results: Former outland had denser tree cover, fewer species and fewer grassland species than former arable fields and hay meadows, irrespective of time since abandonment. Former hay meadows and arable fields with a longer time since abandonment were less rich in species, more wooded and had greater podsolization than meadows and fields abandoned at a later stage. Species richness was higher in hay meadows and arable fields at farms with larger infield area and deeper soils compared with farms with smaller infield area and shallower soils. The greatest richness of species and most open habitat were former arable fields at larger farms abandoned 50 yr before the study. Former arable fields had the highest number of grassland species. Conclusion: After 50 yr of abandonment, former arable fields were the most important remnant habitats for grassland species and may be a more promising target for restoration than formerly managed grasslands.     

Biophysical and human influences on plant species richness in grasslands: Comparing variegated landscapes in subtropical and temperate regions

A survey of grassy woodlands in the Queensland subtropics was conducted, recording herbaceous species richness at 212 sites on three properties (2756 ha). A range of habitats typical of cattle grazing enterprises was sampled and site variables included lithology, slope position, tree density, soil disturbance, soil enrichment and grazing. Results were compared with a previously published survey of temperate grasslands. Lithology, slope position and tree density had relatively minor effects on plant species richness, although in both surveys there was some evidence of lower species richness on the more fertile substrates. Soil disturbance and soil enrichment significantly reduced the richness of native species in both surveys, while exotic species were insensitive (subtropics) or increased (temperate) with disturbance. Rare native species were highly sensitive to disturbances, including grazing, in the temperate study. Although some trends were similar for rare species in the subtropics, the results were not significant and there were complex interactions between grazing, lithology and slope position. Grazing did not have a negative effect on native species richness, except in the closely grazed patches within pastures, and then only on the most intensively developed property. At the scale recorded (30 m²), the native pastures, roadsides and stock routes sampled in the subtropics appear to be among the most species‐rich grasslands ever reported, both nationally and globally. Native species richness was approximately 50% higher than the temperate survey figures across all the comparable habitats. While there are no clear reasons for this result, potential explanations are proposed.