Contrasting needs of grassland dwellers: habitat preferences of endangered steppe beetles (Coleoptera)

Temperate grasslands are local biodiversity hotspots. In Europe, their extent was mostly reduced to isolated habitat patches, whose biota is subject to extinction debt. Knowledge on requirements of dry-grassland inhabitants is thus vital to slow down decline of grassland biodiversity. We studied habitat requirements of eight flightless steppe beetles, including some of the most endangered dry-grassland specialists of the continent. The beetles were sampled using 167 pitfall traps at a pannonian dry-grassland fragment, the Pouzdrany steppe, SE Czech Republic, from March to November 2006. The number of each species captures in each trap was related to vegetation and abiotic habitat characteristics; captures of all beetles were related to each other. Two of the studied species required relatively humid microhabitats, including tall-grass steppe with litter (Carabus hungaricus, Carabidae) and grassland of high herb cover (Meloe proscarabaeus, Meloidae). Others were associated with xeric habitats (e.g. Meloe scabriusculus) and their early-successional stages, including short-turf vegetation (Dorcadion fulvum, D. pedestre, Cerambycidae) and/or bare-ground patches (Blaps lethifera, Tenebrionidae; Meloe decorus, M. uralensis). Our findings point to key importance of early-successional vegetation for grassland biodiversity, and to the fact that locally co-occurring and closely related grassland specialists may exhibit contrasting habitat needs. Spatially and temporarily highly diversified patch management creating a fine scale mosaic of various seral stages from bare soil to tall-grass steppe is therefore the most appropriate approach for managing isolated grasslands. Prescribed burning and support of burrowing herbivores are recommended and discussed together with other measures for restoration of habitat diversity in dry-grassland fragments.     

Habitat and landscape effects on abundance of Missouri's grassland birds

Of 6 million ha of prairie that once covered northern and western Missouri, <36,500 ha remain, with planted, managed, and restored grasslands comprising most contemporary grasslands. Most grasslands are used as pasture or hayfields. Native grasses largely have been replaced by fescue (Festuca spp.) on most private lands (almost 7 million ha). Previously cropped fields set aside under the Conservation Reserve Program (CRP) varied from a mix of cool-season grasses and forbs, or mix of native warm-season grasses and forbs, to simple tall-grass monocultures. We used generalized linear mixed models and distance sampling to assess abundance of 8 species of breeding grassland birds on 6 grassland types commonly associated with farm practices in Missouri and located in landscapes managed for grassland-bird conservation. We selected Bird Conservation Areas (BCAs) for their high percentage of grasslands and grassland-bird species, and for <5% forest cover. We used an information-theoretic approach to assess the relationship between bird abundance and 6 grassland types, 3 measures of vegetative structure, and 2 landscape variables (% grassland and edge density within a 1-km radius). We found support for all 3 levels of model parameters, although there was less support for landscape than vegetation structure effects likely because we studied high-percentage-grassland landscapes (BCAs). Henslow's sparrow (Ammodramus henslowii) counts increased with greater percentage of grassland, vegetation height-density, litter depth, and shrub cover and lower edge density. Henslow's sparrow counts were greatest in hayed native prairie. Dickcissel (Spiza americana) counts increased with greater vegetation height-density and were greatest in planted CRP grasslands. Grasshopper sparrow (A. savannarum) counts increased with lower vegetation height, litter depth, and shrub cover. Based on distance modeling, breeding densities of Henslow's sparrow, dickcissel, and grasshopper sparrow in the 6 grassland types ranged 0.9–2.6, 1.4–3.2, and 0.1–1.5 birds/ha, respectively. We suggest different grassland types and structures (vegetation height, litter depth, shrub cover) are needed to support priority grassland-bird species in Missouri. © 2011 The Wildlife Society.     

Influence of revegetation on predation rates by introduced red foxes (Vulpes vulpes) in south‐eastern Australian farmland

Interactions between wild carnivore abundance and landscape composition in agricultural landscapes are poorly understood despite their importance to both production and conservation. In Australia, introduced red foxes (Vulpes vulpes) prey on both native species and lambs in the temperate agricultural regions. Historically these areas were extensively cleared of native vegetation. Recently revegetation programmes have been implemented, but there is concern that this may benefit foxes and hence increase their impacts. We used an artificial prey placed in eight revegetated (6–12% cover) and 10 cleared (0–1.5% cover) landscapes of ～700 ha to assess how these landscapes influenced fox predation rates. In June and August 2006 (winter) when we expect fox populations to be relatively stable following juvenile dispersal, predation rates were 1.5–2 times higher in revegetated landscapes than in cleared landscapes. We found no evidence of microhabitat effects on predation rates suggesting these landscape‐level differences were probably due to differences in fox population density. In April 2007 (autumn) the results were more variable, possibly indicating more transient populations including dispersing juveniles at this time. Our results suggest that the impact of foxes on highly vulnerable prey could increase with revegetation. However, the benefits of revegetation to prey may offset negative impacts of foxes and future work is required to assess the likely net effects.     

Cultural monuments and nature conservation: a review of the role of kurgans in the conservation and restoration of steppe vegetation

Steppe is among the most endangered biomes of the world, especially in Eastern Europe, where more than 90 % of original steppes have been destroyed due to conversion into croplands, afforestation and other human activities. Currently, steppe vegetation is often restricted to places unsuitable for ploughing, such as ancient burial mounds called kurgans. The aim of our study was to collect and synthesise existing knowledge on kurgans by a review of research papers and grey literature. The proportion of kurgans covered by steppe vegetation increases from west to east and from lowlands to uplands. Despite their small size, kurgans act as biodiversity hotspots and harbour many red-listed species. High overall species richness and a high proportion of grassland specialists are maintained by a pronounced fine-scale environmental heterogeneity. The main factors threatening the biodiversity of kurgans are intensified agriculture and construction works. We conclude that kurgans can play a crucial role in preserving steppe vegetation, especially in intensively used agricultural landscapes in the western part of the steppe zone. Despite the vital role of kurgans in sustaining steppe vegetation, we identified serious knowledge gaps on their distribution, vegetation, flora and fauna and their potential role in steppe restoration.     

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.     

Burrowing activities of kangaroo rats and patterns in plant species dominance at a shortgrass steppe-desert grassland ecotone

Abstract. Our objective was to evaluate the effects of burrowing activities by banner-tail kangaroo rats (Dipodomys spectabilis Merriam) on plant community structure and species dominance for two patch types at the ecotone between shortgrass steppe and desert grassland in New Mexico, USA. 10 mounds produced by kangaroo rats were selected in patches dominated by Bouteloua gracilis (the dominant in shortgrass steppe communities) and 10 mounds were selected in patches dominated by B. eriopoda (the dominant in Chihuahuan desert grasslands). Plant cover and density by species were sampled from three locations associated with each mound: the mound proper, the edge of the mound in the transition area, and the off-mound vegetation. Similar cover of B. eriopoda for the edges of mounds in both patch types indicates the ability of this species to respond to animal disturbances regardless of the amount of cover in the surrounding undisturbed vegetation. By contrast, cover of B. gracilis was low for all mounds and mound edges in patches dominated by this species. Much higher cover of B. eriopoda on mound edges compared to the undisturbed vegetation in B. gracilis-dominated patches indicates that kangaroo rats have important positive effects on this species. Lower cover of perennial grasses and higher cover of forbs, shrubs, and succulents on the edges of mounds in B. eriopoda-dominated patches compared to patches dominated by B. gracilis indicate the importance of surrounding vegetation to plant responses on disturbed areas. Our results show that kangaroo rats have important effects on both species dominance and composition for different patch types, and may provide a mechanism for small-scale dominance patterns at an ecotone; thus providing further support for their role as keystone species in desert grasslands.     

Cape ground squirrels as ecosystem engineers: modifying habitat for plants,small mammals and beetles in Namib Desert grasslands

Burrowing and foraging of semi‐fossorial rodents can affect species distribution and composition. Ground squirrels dig large burrow systems for refuge from predators and temperature extremes. Burrowing and foraging around burrows by squirrels may affect habitat and resource distributions for other organisms. We examined the impact of Cape ground squirrels (Xerus inauris) on vegetation, small mammals and beetles during winter and summer in grasslands on the edge of the Namib Desert. At each burrow system and paired control site without burrows, we estimated plant cover and height using quadrats (N = 8 paired sites), small mammal abundance and species richness using mark‐recapture techniques (N = 8 paired sites) and beetle abundance and species richness using pitfall traps (N = 6 paired sites, winter only). Squirrel burrowing and foraging activities resulted in lower plant cover and height, higher small mammal abundance and lower beetle abundance and species richness. Squirrels also reduced more plant cover in winter compared to summer, but had no effect on small mammal species richness. Furthermore, plant cover and height were higher in summer, whereas small mammal abundance and species richness were higher in winter. Our results suggest that Cape ground squirrels are important ecosystem engineers that influence plant and animal communities in the Namib Desert grasslands.     

Species diversity of remnant calcareous grasslands in south eastern Germany depends on litter cover and landscape structure

Species diversity depends on, often interfering, multiple ecological drivers. Comprehensive approaches are hence needed to understand the mechanisms determining species diversity. In this study, we analysed the impact of vegetation structure, soil properties and fragmentation on the plant species diversity of remnant calcareous grasslands, therefore, in a comparative approach.We determined plant species diversity of 18 calcareous grasslands in south eastern Germany including all species and grassland specialists separately. Furthermore, we analysed the spatial structure of the grasslands as a result of fragmentation during the last 150 years (habitat area, distance to the nearest calcareous grassland and connectivity in 1830 and 2013). We also collected data concerning the vegetation structure (height of the vegetation, cover of bare soil, grass and litter) and the soil properties (content of phosphorous and potassium, ratio of carbon and nitrogen) of the grassland patches. Data were analysed using Bayesian multiple regressions.We observed a habitat loss of nearly 80% and increasing isolation between grasslands since 1830. In the Bayesian multiple regressions the species diversity of the studied grasslands depended negatively on cover of litter and to a lower degree on the distance to the nearest calcareous grassland in 2013, whereas soil properties had no significant impact.Our study supports the observation that vegetation structure, which strongly depends on land use, is often more important for the species richness of calcareous grasslands than fragmentation or soil properties. Even small and isolated grasslands may, therefore, contribute significantly to the conservation of species diversity, when they are still grazed.     

Distribution and effects of tree leaf litter on vegetation composition and biomass in a forest-grassland ecotone

Aims After abandonment of grasslands, secondary succession leads to the invasion by woody species. This process begins with the accumulation of tree litter in the forest–grassland ecotone. Our objectives were to determine the relationships between litter amounts and vegetation composition and cover along natural forest–grassland ecotones and to experimentally study the initial effects of tree litter accumulation on grassland vegetation and on microsite conditions.Methods We established 11 transects varying from 12 to 15 m in length in different forest–grassland ecotones in the Lahn-Dill highlands, Germany, and measured the mass and cover of tree litter and the cover and composition of vegetation at five sequential positions along each transect by using 1 m 2 plots with five replications. In a field experiment, we established plots subjected to different litter amounts (0, 200 and 600g m ?2) and evaluated changes in grassland vegetation, soil temperature and soil nutrient availability below the litter layer.Important findings Tree litter amounts decrease from 650 to 65g m ?2 across the forest–grassland ecotone. Vegetation changed from shrubs and annual species (adapted to more stressful conditions) in the forests edge to grasses, rosettes and hemirosette species (with higher competitive abilities) in the grassland. These anthropogenic forest–grassland ecotones showed abrupt edges, and the two adjacent ecosystems were characterized by different species pools and functional groups. In the field experiment, the presence of a litter layer reduced vegetation biomass and cover; the species richness was only reduced in the treatment with high litter (600g m ?2). Additionally, adding litter on top of vegetation also reduced thermal amplitude and the number of frost days, while increasing the availability of some nutrients, such as nitrogen and aluminium, the latter being an indicator of soil acidification. Adding a tree litter layer of 600g m ?2 in grassland areas had strong effects on the composition and diversity of grassland vegetation by reducing the cover of several key grassland species. In, or near, forest edges, litter accumulation rapidly changes established vegetation, microsite conditions and soil nutrients.     

The influence of wild boar <Emphasis Type="Italic">(Sus scrofa)</Emphasis> on microhabitat quality for the endangered butterfly <Emphasis Type="Italic">Pyrgus malvae</Emphasis> in the Netherlands

The decline of open habitats in Europe, such as semi-natural grasslands and heathlands, has caused a general decline in biodiversity, which has been well documented for butterflies. Current conservation practices often involve grazing by domestic livestock to maintain suitable butterfly habitats. The extent to which wild ungulates may play a similar role remains largely unknown. Through their rooting activity, wild boar could be effective to reduce grass encroachment and restore pioneer microhabitats that are vital to many grassland insects in temperate climates. Here, we assessed the microhabitat requirements of Pyrgus malvae, an endangered butterfly of heathland and grassland habitats in the Netherlands, with special attention for the influence of wild boar rooting. To date, oviposition site selection of this species has concentrated on calcareous grasslands, whereas we also include heathlands. Overall, larval occupancy was higher in warm, open and sparsely vegetated microhabitats, which supports earlier findings. In heathland, microhabitat occupancy was positively affected by bryophyte and litter cover. In heath-grassland mosaic, microhabitat occupancy was also influenced by bryophyte and litter cover, but in addition low grass cover increased occupancy by favouring host plants. In grassland, only low grass cover and host plant cover determined microhabitat quality. Across all habitats, occupied microhabitats were characterized by lower vegetation as well as higher average daytime temperatures than unoccupied microhabitats. We discovered that wild boar play an important role in reducing grass cover by shallow rooting in grass patches, thereby increasing host plant availability. Hence, wild boar may have an added value in maintaining and restoring P. malvae microhabitats in grassland habitats in addition to grazing by domestic livestock.     

Multi-scale impacts of crested wheatgrass invasion in mixed-grass prairie

Most of North America’s northern Great Plains have been cultivated for crop production, leaving remnants of natural mixed-grass prairie fragmented and threatened by alien plant invasions. The region’s most widespread alien perennial forage crop, crested wheatgrass (Agropyron cristatum sensu amplo), has invaded native grassland and raised concerns regarding its ecological impact. To evaluate impacts at multiple scales of organization, adjacent invaded and uninvaded mixed-grass prairie were sampled at eight widely separated locations. At the population level, native C₃ mid-grasses and forbs were less abundant in invaded grasslands, while native C₃ and C₄ short-grass abundance was not different. At community and landscape levels, diversity was lower in invaded grasslands largely because of lower forb species richness and cover, and crested wheatgrass dominance of both cover (14% basal cover) and seedbank (404 seeds m⁻²). At the ecosystem level, both vegetation and litter biomass were greater in invaded grasslands, however, below ground organic matter (roots and litter), soil organic carbon, total nitrogen and phosphorus were not different. Crested wheatgrass invasion of mixed-grass prairie was associated with lower diversity within and among plant communities, and appears to simplify the composition of mixed-grass prairie landscapes. Hypotheses for crested wheatgrass dominance and persistence following invasion are suggested.     

Disproportionate effects of non‐colonial small herbivores on structure and diversity of grassland dominated by large herbivores

The response of semiarid grasslands to small, non‐colonial herbivores has received little attention, focusing primarily on the effects of granivore assemblages on annual plant communities. We studied the long‐term effects of both small and large herbivores on vegetation structure and species diversity of shortgrass steppe, a perennial semiarid grassland considered marginal habitat for small mammalian herbivores. We hypothesized that 1) large generalist herbivores would affect more abundant species and proportions of litter‐bare ground‐vegetation cover through non‐selective herbivory, 2) small herbivores would affect less common species through selective but limited consumption, and 3) herbivore effects on plant richness would increase with increasing aboveground net primary production (ANPP). Plant community composition was assessed over a 14‐year period in pastures grazed at moderate intensities by cattle and in exclosures for large (cattle) and large‐plus‐small herbivores (additional exclusion of rabbits and rodents). Exclusion of large herbivores affected litter and bare ground and basal cover of abundant, common and uncommon species. Additional exclusion of small herbivores did not affect uncommon components of the plant community, but had indirect effects on abundant species, decreased the cover of the dominant grass Bouteloua gracilis and total vegetation, and increased litter and species diversity. There was no relationship between ANPP and the intensity of effects of either herbivore body size on richness. Exclusion of herbivores of both body sizes had complementary and additive effects which promoted changes in vegetation composition and physiognomy that were linked to increased abundance of tall and decreased abundance of short species. Our findings show that small mammalian herbivores had disproportionately large effects on plant communities relative to their small consumption of biomass. Even in small‐seeded perennial grasslands with a long history of intensive grazing by large herbivores, non‐colonial small mammalian herbivores should be recognized as an important driver of grassland structure and diversity.     

Effects of Habitat on Competition Between Kit Foxes and Coyotes

Abstract: San Joaquin kit foxes (Vulpes macrotis mutica) are an endangered species with a narrow geographic range whose natural populations are limited by predation by coyotes (Canis latrans). In the warm, arid grassland and shrubland habitats where kit foxes occur, coyotes are more cover dependent than kit foxes, creating the possibility of habitat segregation. Effects of habitat variation on coyote and kit fox competition are unknown. We assessed exploitation and interference competition between coyotes and kit foxes in grassland and shrubland habitats to determine if such competition varies among habitats. With respect to exploitation competition, we evaluated habitat and spatial partitioning, diet, prey abundance, and survival for kit foxes and coyotes at the Lokern Natural Area in central California, USA, from January 2003 through June 2004. Kit foxes partitioned habitat, space, and diet with coyotes. Coyotes primarily used shrubland habitats whereas kit foxes selectively used burned grasslands. Kit foxes and coyotes had high dietary overlap with regards to items used, but proportional use of items differed between the 2 species. Kit foxes selected for Heermann's kangaroo rats (Dipodomys heermanni), which were closely tied to shrub habitats. With respect to interference competition, predation was the primary source of mortality for kit foxes, and survival of individual kit foxes was inversely related to proportion of shrub habitat within their home ranges. Our results suggest that a heterogeneous landscape may benefit kit foxes by providing habitat patches where predation risk may be lower.     

Clustered animal burrows yield higher spatial heterogeneity

An understanding of the relationships between spatial heterogeneity and disturbance regime is important for establishing the mechanisms necessary to maintain biodiversity. Our objective was to examine how the configuration of disturbance by burrowing rodents (Siberian marmot) affected the spatial heterogeneity of vegetation and soil nutrient properties. We established three 2500-m² (50 m × 50 m) isolated-burrows plots and three 2500-m² clustered-burrows plots in a Mongolian grassland. Each plot was subdivided into 4-m² quadrats, and the plant species richness, percent coverage, and soil nutrient properties in the quadrats were surveyed. Spatial heterogeneity was calculated for vegetation using the mean dissimilarity of species composition among sample quadrats, and geostatistical analysis was used to calculate soil properties. Heterogeneous patches of plants such as Achnatherum splendens and higher nutrient concentrations were found only near the clustered burrows. As a result, spatial heterogeneities of vegetation and soil nutrient properties were higher in the clustered colony than those in the isolated colony. The configuration of disturbance patches affected the spatial heterogeneity at the landscape level through the spatial pattern of disturbance frequency.     

Biodiversity of semi‐arid Mediterranean grasslands: Impact of grazing and afforestation

Question: What is the impact of grazing and/or afforestation on grassland diversity, species composition and cover parameters? Location: Semi‐arid Mediterranean grasslands of Jordan. Methods: Vegetation, litter, bare soil and rock cover were compared among four management types – free grazing and protected from grazing with three levels of tree cover. Species composition, plant cover, species richness and evenness were used to evaluate differences in vegetation among management types. Species composition differences among management types were also investigated. Results: Semi‐arid Mediterranean grasslands harbour appreciable levels of plant biodiversity. Grazing did not affect plant diversity, indicating the high resilience against and adaptation to grazing; however,grazing affected species composition and cover parameters. Afforestation seems to protect soil through higher litter cover but its impact on plant biodiversity was negative and markedly affected species composition. Conclusions: Neither protection from grazing or massive afforestation alone are sufficient for conserving biodiversity in this system. A management model is suggested where the landscape should be maintained as a mosaic of four management types: complete protection from grazing, grazing rotation, planting sparse trees in eroded areas and revegetating degraded areas using native, herbaceous and grazing tolerant species.     

Spontaneous steppe restoration on abandoned cropland in Kazakhstan: grazing affects successional pathways

After the collapse of the Soviet Union in 1991, 12 million ha of cropland were abandoned in the steppe zone of Kazakhstan. At the same time livestock numbers crashed, leaving large areas of steppe without any grazing. We aimed to investigate, to which degree spontaneous succession on former cropland leads to the restoration of native steppe vegetation, and how this process is influenced by changing grazing patterns. We recorded biomass and vegetation characteristics as well as local soil and landscape variables in 151 quadrats of 100 m², covering 89 plots on former cropland abandoned in the early 1990s and 62 reference plots of near-natural steppe grassland that was never ploughed. About half of the plots of each category were located in a remote region where grazing has been absent for ca. 20 years, whereas the other half was located in a region with moderate livestock grazing. While there were no differences in the diversity, structure and plant life form composition of currently grazed and un-grazed near-natural steppe grasslands, corresponding successional plots on abandoned arable land exhibited significant differences. Grazed plots on former fields showed higher species richness and a higher cover of dwarf shrubs (mostly Artemisia spec.), ruderals and perennial herbs. At the same time, immigration of typical steppe species was much more successful. Contrary, in the absence of any grazing we found species-poor swards dominated by Stipa lessingiana and Leymus ramosus exhibiting an increasing frequency of wildfires due to litter accumulation. After 15–20 years, secondary steppe grasslands still differed substantially from their near-natural references. Our results suggest that grazing is mandatory to fully restore the original near-natural steppe vegetation and the underlying processes of pyric herbivory.     

Ecosystem engineering varies spatially: a test of the vegetation modification paradigm for prairie dogs

Colonial, burrowing herbivores can be engineers of grassland and shrubland ecosystems worldwide. Spatial variation in landscapes suggests caution when extrapolating single‐place studies of single species, but lack of data and the need to generalize often leads to ‘model system’ thinking and application of results beyond appropriate statistical inference. Generalizations about the engineering effects of prairie dogs (Cynomys sp.) developed largely from intensive study at a single complex of black‐tailed prairie dogs C. ludovicianus in northern mixed prairie, but have been extrapolated to other ecoregions and prairie dog species in North America, and other colonial, burrowing herbivores. We tested the paradigm that prairie dogs decrease vegetation volume and the cover of grasses and tall shrubs, and increase bare ground and forb cover. We sampled vegetation on and off 279 colonies at 13 complexes of 3 prairie dog species widely distributed across 5 ecoregions in North America. The paradigm was generally supported at 7 black‐tailed prairie dog complexes in northern mixed prairie, where vegetation volume, grass cover, and tall shrub cover were lower, and bare ground and forb cover were higher, on colonies than at paired off‐colony sites. Outside the northern mixed prairie, all 3 prairie dog species consistently reduced vegetation volume, but their effects on cover of plant functional groups varied with prairie dog species and the grazing tolerance of dominant perennial grasses. White‐tailed prairie dogs C. leucurus in sagebrush steppe did not reduce shrub cover, whereas black‐tailed prairie dogs suppressed shrub cover at all complexes with tall shrubs in the surrounding habitat matrix. Black‐tailed prairie dogs in shortgrass steppe and Gunnison's prairie dogs C. gunnisoni in Colorado Plateau grassland both had relatively minor effects on grass cover, which may reflect the dominance of grazing‐tolerant shortgrasses at both complexes. Variation in modification of vegetation structure may be understood in terms of the responses of different dominant perennial grasses to intense defoliation and differences in foraging behavior among prairie dog species. Spatial variation in the engineering role of prairie dogs suggests spatial variation in their keystone role, and spatial variation in the roles of other ecosystem engineers. Thus, ecosystem engineering can have a spatial component not evident from single‐place studies.     

Landscape-scale connectivity and fragment size determine species composition of grassland fragments

As a consequence of agricultural intensification and habitat fragmentation since the mid-20th century, biological diversity has declined considerably throughout the world, particularly in Europe. We assessed how habitat and landscape-scale heterogeneity, such as variation in fragment size (small vs. large) and landscape configuration (measured as connectivity index), affect plant and arthropod diversity. We focused on arthropods with different feeding behaviour and mobility, spiders (predators, moderate dispersal), true bugs (mainly herbivores and omnivores with moderate dispersal), wild bees (pollinators with good dispersal abilities), and wasps (pollinators, omnivores with good dispersal abilities). We studied 60 dry grassland fragments in the same region (Hungarian Great Plain); 30 fragments were represented by the grassland component of forest-steppe stands, and 30 were situated on burial mounds (kurgans). Forest-steppes are mosaics of dry grasslands with small forests in a matrix of plantation forests. Kurgans are ancient burial mounds with moderately disturbed grasslands surrounded by agricultural fields. The size of fragments ranged between 0.16–6.88 ha (small: 0.16–0.48 ha, large: 0.93–6.88 ha) for forest-steppes and 0.01–0.44 ha (small: 0.01–0.10 ha and large: 0.20–0.44 ha) for kurgans. Fragments also represented an isolation gradient from almost cleared and homogenous landscapes, to landscapes with relatively high compositional heterogeneity. Fragment size, connectivity, and their interaction affected specialist and generalist species abundances of forest-steppes and kurgans. Large fragments had higher species richness of ground-dwelling spiders, and the effect of connectivity was more strongly positive for specialist arthropods and more strongly negative for generalists in large than in small fragments. However, we also found a strong positive impact of connectivity for generalist plants in small kurgans in contrast to larger ones. We conclude that besides the well-known effect of enhancing habitat quality, increasing connectivity between fragments by restoring natural and semi-natural habitat patches would help to maintain grassland biodiversity.     

Long‐term effects of the abandonment of grazing on steppe‐like grasslands

Question: What are the consequences of grazing abandonment on the Stipa lessingiana dominated steppe‐like grasslands? What is the relative importance of management and environmental factors in causing variation in species composition and abundance in the continuously grazed and abandoned grassland stands? Location: Transylvanian Lowland, Romania. Methods: Repeated vegetation mapping of a grassland stand, where grazing was abandoned 35 years ago; re‐sampling six grassland stands surveyed 29–57 years ago. For revealing long‐term changes in species composition and rank abundance PCoA ordination was applied. The relative importance of management and environmental factors in structuring vegetation were explored by CCA ordination. Diversity, evenness and the relative number and abundance of red‐listed species were compared between managed and abandoned stands. Results: Our results pointed out that grasslands which were formerly grazed and dominated by S. lessingiana, in the long‐term absence of grazing, have been transformed into a S. pulcherrima dominated type. Management, probably by creating bare surfaces and preventing litter accumulation, had the strongest effect on the species composition and abundance in the grasslands. Abandoned grassland stands had lower diversity and evenness compared to continuously grazed stands. While at the same time, the relative number of threatened, rare species did not differ between managed and abandoned sites. Conclusion: Maintaining extensively grazed, as well as un‐managed, Stipa dominated grasslands would be important in order to create various habitat conditions for plant species, especially threatened and rare species, and promote diversity on the landscape scale.     

Location of long‐term communal burrows of a threatened arid‐zone lizard in relation to soil and vegetation

The great desert skink (Liopholis kintorei) of the Egerniinae subfamily (Reptilia: Scincidae) is a communal burrowing lizard that inhabits arid spinifex grasslands in central Australia. Great desert skink activity is centred in and around the burrows which are inhabited for many years. However, it is not known whether skinks select burrow sites with specific attributes or how continuing occupancy of burrows is influenced by the surrounding habitat; especially post‐fire, when plant cover is reduced. Here, we test whether great desert skink burrows in areas burnt 2 years previously and in longer unburnt areas are associated with particular habitat attributes, and whether there are differences between occupied and recently abandoned burrow sites. Vegetation composition, cover and soil surface characteristics at 56 established great desert skink burrows, including occupied and recently unoccupied burrows, were compared with 56 random nearby non‐burrow control sites. Burrow sites had higher plant cover compared with the surrounding landscape in both recently burnt and longer unburnt areas and were more likely to be associated with the presence of shrubs. Soil stability and infiltration were also higher at burrow sites. However, we found no evidence that burrows with lower cover were more likely to be abandoned. Our results suggest that great desert skinks may actively select high cover areas for burrow construction, although differences between burrow and control sites may also partly reflect local changes to plant cover and composition and soil properties resulting from burrow construction and long‐term habitation of a site. Further research should determine if burrows with shrubs or higher plant cover provide greater protection from predators, more structural stability for burrow construction, increased prey abundance or other benefits. We recommend that maintenance of areas with relatively higher plant cover be prioritized when managing great desert skink habitat.