Does desertification diminish biodiversity? Enhancement of ant diversity by shrub invasion in south‐western USA

The conversion of desert grasslands to shrublands is a long‐standing concern in the south‐western United States, but the effects of this change on native animals defy generalization. Here, I consider evidence that shrub invasion and encroachment, particularly that of honey mesquite (Prosopis glandulosa), has led to increasing ecological dominance and diversity of ants in general, as well as increases in specific native taxa. The effects of shrub invasion on ants were measured at two scales: (1) between Chihuahuan Desert landscapes that vary slightly in temperature and strongly in the dominance of mesquite, and (2) across variation in mesquite density occurring within a generally mesquite‐dominated landscape. Ant richness and numerical dominance was measured at pitfall traps over 2 years and baits were used to assess ecological dominance across different temperatures. The mesquite‐dominated Jornada site harboured four times the number of ant foragers found at the relatively ‘pristine’ Sevilleta site, with several ecologically dominant taxa driving this pattern, especially Dorymyrmex bicolor. Species richness and ecological dominance were also greatest at the Jornada. Within the Jornada landscape, turnover in species composition was related to mesquite density, but local richness and abundance was unrelated to mesquite density. Coupled with the results of previous manipulative experiments and comparative studies, there is support for the notion that ant diversity is not negatively affected by shrub invasion but that several taxa prosper from it. The Jornada is uniquely saturated by dominant ant taxa, perhaps as a consequence of an overall high level of shrub availability that provides a reliable source of carbohydrate‐rich plant exudates. This raises important questions about the relationship between environmental degradation, ecosystem productivity, and animal diversity.     

Effectiveness of sand-fixing measures on desert land restoration in Kerqin Sandy Land,northern China

In the semi-arid Kerqin Sandy Land of north China, land desertification and frequent sand storms in the spring strongly affect the growth of grassland vegetation and crops, and give rise to large reductions in yield as a result of wind erosion and sand dune movement. To bring desertification under control and reduce its influence on grassland and farmlands, many measures have been developed and implemented for stabilizing mobile sand dunes and restoring desertified grasslands. This study was conducted from 1996 to 2003 to evaluate the effectiveness of desert land restoration after implementation of sand-fixing measures. The results showed that construction of straw checkerboards and planting of shrub seedlings significantly enhances topsoil development on the dune surface, increasing silt and clay content and facilitating accumulation of soil carbon and total N, as well as accelerating an increase in plant diversity, vegetation cover and plant density. These findings suggest that using straw checkerboards and planting shrubs are successful methods for mobile sand dune stabilization and desertified grassland restoration in semiarid regions. The mechanism creating these changes is a reduction in wind erosion and improvement of the soil environment for plants. In addition, our results showed that construction of straw checkerboards was slightly more effective in vegetation and soil restoration in comparison with planting shrub seedlings, especially at the primary stage.     

Response of dominant grass and shrub species to water manipulation: an ecophysiological basis for shrub invasion in a Chihuahuan Desert Grassland

Increases in woody vegetation and declines in grasses in arid and semi-arid ecosystems have occurred globally since the 1800s, but the mechanisms driving this major land-cover change remain uncertain and controversial. Working in a shrub-encroached grassland in the northern Chihuahuan Desert where grasses and shrubs typically differ in leaf-level nitrogen allocation, photosynthetic pathway, and root distribution, we asked if differences in leaf-level ecophysiology could help explain shrub proliferation. We predicted that the relative performance of grasses and shrubs would vary with soil moisture due to the different morphological and physiological characteristics of the two life-forms. In a 2-year experiment with ambient, reduced, and enhanced precipitation during the monsoon season, respectively, the encroaching C(3) shrub (honey mesquite Prosopis glandulosa) consistently and substantially outperformed the historically dominant C(4) grass (black grama Bouteloua eriopoda) in terms of photosynthetic rates while also maintaining a more favorable leaf water status. These differences persisted across a wide range of soil moisture conditions, across which mesquite photosynthesis was decoupled from leaf water status and moisture in the upper 50 cm of the soil profile. Mesquite's ability to maintain physiologically active leaves for a greater fraction of the growing season than black grama potentially amplifies and extends the importance of physiological differences. These physiological and phenological differences may help account for grass displacement by shrubs in drylands. Furthermore, the greater sensitivity of the grass to low soil moisture suggests that grasslands may be increasingly susceptible to shrub encroachment in the face of the predicted increases in drought intensity and frequency in the desert of the southwestern USA.     

Restoration Practices Have Positive Effects on Breeding Bird Species of Concern in the Chihuahuan Desert

Woody plant encroachment into grasslands is a global concern. Efforts to restore grasslands often assume that removal of woody plants benefits biodiversity but assumptions are rarely tested. In the Chihuahuan Desert of the Southwestern United States, we tested whether abundances of grassland specialist bird species would be greater in plant communities resulting from treatment with herbicides to remove encroaching shrubs compared with untreated shrub‐dominated areas that represented pre‐treatment conditions. In 2010, we surveyed breeding birds and vegetation at 16 treated–untreated pairs. In 2011, we expanded the survey effort to 21 treated–untreated pairs, seven unpaired treatment areas, and five reference grassland areas. Vegetation in treatment areas had higher perennial grass foliar and basal cover and lower shrub foliar cover compared with untreated areas. Several regionally declining grassland specialists exhibited higher occurrence and relative abundance in treated areas. A shrubland specialist, however, was associated with untreated areas and may be negatively impacted by shrub removal. Bird community composition differed between treated and untreated areas in both years. Our results indicate that shrub removal can have positive effects on grassland specialist bird species, but that a mosaic of treated and untreated areas might be most beneficial for regional biodiversity .     

Directional climate change and potential reversal of desertification in arid and semiarid ecosystems

Our objective was to determine if long‐term increases in precipitation can maintain grasslands susceptible to desertification, and initiate a reversal of historic regime shifts on desertified shrublands. Perennial grass production and species richness in a multi‐year wet period were hypothesized to be greater than expected based on precipitation in a sequence of dry years. These responses were expected to differ for grasslands and shrublands with different dominant species and topo‐edaphic properties. Long‐term trends in desertification were documented using vegetation maps beginning in 1858, 1915, 1928, and 1998). These trends were compared with herbaceous and woody species responses to a sequence of dry (1994–2003) and wet years (2004–2008) for two grassland (uplands, playas) and three desertified shrubland types (honey mesquite, creosotebush, tarbush) in the Chihuahuan Desert. Analyses showed that both types of grasslands decreased in spatial extent since 1858 whereas areas dominated by mesquite or creosotebush increased. Production of upland grasslands in the wet period was greater than expected based on responses during the dry period whereas the relationships between species richness and precipitation was the same for both periods. Precipitation was not important to responses in playa grasslands in either period. For all ecosystem types, the production response in wet years primarily was an increase in herbaceous plants, and the most pronounced responses occurred on sandy sites (upland grasslands, mesquite shrubland). Results suggest that multiple wet years are needed to initiate a sequence of grass establishment and survival processes that can maintain upland grasslands without management inputs and lead to a state change reversal in desertified shrublands. Restoration strategies need to take advantage of opportunities provided by future climates while recognizing the importance of ecosystem type.     

Is the removal of aboveground shrub biomass an effective technique to restore a shrub‐encroached grassland?

Encroachment of woody plants into grasslands is a global phenomenon that has substantial impacts on pastoral productivity and ecosystem services. Over the past half century, pastoralists and land management agencies have explored various options to control woody plants in order to improve ecosystem services in shrub‐encroached grasslands. We examined the effectiveness of controlling the encroachment of the shrub Caragana microphylla into grassland in Inner Mongolia, China. We cut and removed all of the aboveground biomass from 450 shrubs, predicting that the effectiveness of this technique to control shrubs would depend on shrub morphology. Specifically, we expected that larger shrubs with more biomass would be more difficult to kill by cutting than smaller shrubs. A year after treatment, we found that cutting killed only 11% of the 450 treated shrubs, and of these, three‐quarters of the locations that they occupied reverted to grasses and one‐quarter to bare soil. Shrubs that survived the cutting treatment produced more stems and leaf biomass, and therefore had a greater leaf to stem ratio. Shrubs that died after cutting had a lower crown area and basal area, and less stem biomass than shrubs that resprouted within 12 months of cutting. There were no effects of shrub height on the fate of treated shrubs. Cutting had no effect on understory plant cover or richness, but reproductive plants were taller under shrubs that were not cut. Overall, our study showed that removing aboveground shrub biomass by cutting is an ineffective technique for “restoring” the original grassland community unless shrubs are very small. Strategic targeting of small shrubs would be a more effective technique for controlling the spread of C. microphylla in the long term.     

Winter survival of North American grassland birds is driven by weather and grassland condition in the Chihuahuan Desert

Populations of grassland birds that overwinter in the Chihuahuan Desert are declining more rapidly than other grassland birds, and survival during the non‐breeding season may have a strong influence on population trends of these species. Habitat loss and deterioration due to desertification may be contributing to these declines, and the winter ecology of grassland birds under these changing environmental conditions remains relatively unexplored. To fill this information gap, we estimated the survival of two grassland‐obligate sparrows, Baird's Sparrows (Ammodramus bairdii) and Grasshopper Sparrows (A. savannarum), on their wintering grounds in the Chihuahuan Desert, and investigated the role of habitat structure and weather on survival rates. We deployed radio‐transmitters on Baird's (N = 49) and Grasshopper (N = 126) sparrows near Janos, Chihuahua, and tracked birds from November to March during the winters of 2012–2013 and 2013–2014. Causes of mortality included avian predators, mammals, and possibly weather. We estimated an overall weekly winter survival probability of ? = 92.73% (95% CI[s] = 88.63–95.44%) for Baird's Sparrows in 2012–2013. We estimated a weekly winter survival probability of ? = 93.48% (95% CI[s] = 90.29–96.67%) and ? = 98.78% (95% CI[s] = 97.88–99.68%) for Grasshopper Sparrow in 2012–2013 and 2013–2014, respectively. Weekly winter survival was lower with colder daily minimum temperatures for both species and in areas with taller shrubs for Grasshopper Sparrows, with the shrubs potentially increasing predation risk by providing perches for Loggerhead Shrikes (Lanius ludovicianus). Our results highlight the need to maintain healthy grass structure in wintering areas to provide birds with food, protection from predators, and adequate cover from inclement weather. Our results also demonstrate that the presence of shrubs can lower winter survival, and suggest that shrub encroachment into the winter habitat of these sparrows may be an important driver of their population declines. Shrub removal could increase survival of wintering sparrows in the Chihuahuan Desert by reducing availability of perches for avian predators, thus reducing predation risk.     

Soil fungal composition changes with shrub encroachment in the northern Chihuahuan Desert

Woody species encroachment of grasslands globally causes many socioecological impacts, including loss of grazing pastures and decreased biodiversity. Soil microbial communities may partially regulate the pace of shrub encroachment, as plant-microbial interactions can strongly influence plant success. We measured fungal composition and activity under dominant plant species across a grassland to shrubland transition to determine if shrubs cultivate soil microbial communities as they invade. Specifically, soil microbial communities, abiotic soil properties, and extracellular enzyme activities were quantified for soils under four common Chihuahuan Desert plant species (three grasses, one shrub) in central New Mexico, U.S.A. Extracellular enzyme activity levels were fairly consistent under different plant species across the grassland to shrubland transition. Activity levels of two enzymes (alkaline phosphatase and beta-N-acetyl-glucosaminidase) were lower in the ecotone, presumably because soil organic matter content was also lower in ecotone soils. Community composition of soil fungi mirrored patterns in the plant community, with distinct plant and fungal communities in the shrubland and grassland, while grassland-shrubland ecotone soils hosted a mix of taxa from both habitats. We show that shrubs cultivate a distinct microbial community on the leading edge of the invasion, which may be necessary for shrub colonization, establishment, and persistence.     

The role of vegetation–microclimate feedback in promoting shrub encroachment in the northern Chihuahuan desert

Many arid and semi‐arid landscapes around the world are affected by a shift from grassland to shrubland vegetation, presumably induced by climate warming, increasing atmospheric CO₂ concentrations, and/or changing land use. This major change in vegetation cover is likely sustained by positive feedbacks with the physical environment. Recent research has focused on a feedback with microclimate, whereby cold intolerant shrubs increase the minimum nocturnal temperatures in their surroundings. Despite the rich literature on the impact of land cover change on local climate conditions, changes in microclimate resulting from shrub expansion into desert grasslands have remained poorly investigated. It is unclear to what extent such a feedback can affect the maximum extent of shrub expansion and the configuration of a stable encroachment front. Here, we focus on the case of the northern Chihuahuan desert, where creosotebush (Larrea tridentata) has been replacing grasslands over the past 100–150 years. We use a process‐based coupled atmosphere‐vegetation model to investigate the role of this feedback in sustaining shrub encroachment in the region. Simulations indicate that the feedback allows juvenile shrubs to establish in the grassland during average years and, once established, reduce their vulnerability to freeze‐induced mortality by creating a warmer microclimate. Such a feedback is crucial in extreme cold winters as it may reduce shrub mortality. We identify the existence of a critical zone in the surroundings of the encroachment front, in which vegetation dynamics are bistable: in this zone, vegetation can be stable both as grassland and as shrubland. The existence of these alternative stable states explains why in most cases the shift from grass to shrub cover is found to be abrupt and often difficult to revert.     

Desertification alters patterns of aboveground net primary production in Chihuahuan ecosystems

Abstract The Chihuahuan desert of New Mexico, USA, has changed in historical times from semiarid grassland to desert shrublands dominated by Larrea tridentata and Prosopis glandulosa. Similar displacement of perennial grasslands by shrubs typifies desertification in many regions. Such structural vegetation change could alter average values of net primary productivity, as well as spatial and temporal patterns of production. We investigated patterns of aboveground plant biomass and net primary production in five ecosystem types of the Jornada Basin Long‐Term Ecological Research (LTER) site. Comparisons of shrub‐dominated desertified systems and remnant grass‐dominated systems allowed us to test the prediction that shrublands are more heterogeneous spatially, but less variable over time, than grasslands. We measured aboveground plant biomass and aboveground net primary productivity (ANPP) by species, three times per year for 10 years, in 15 sites of five ecosystem types (three each in Larrea shrubland, Bouteloua eriopoda grassland, Prosopis dune systems, Flourensia cernua alluvial flats, and grass‐dominated dry lakes or playas). Spatial heterogeneity of biomass at the scale of our measurements was significantly greater in shrub‐dominated systems than in grass‐dominated vegetation. ANPP was homogeneous across space in grass‐dominated systems, and in most growing seasons was significantly more patchy in shrub vegetation. Substantial interannual variability in ANPP complicates comparison of mean values across ecosystem types, but grasslands tended to support higher ANPP values than did shrub‐dominated systems. There were significant interactions between ecosystem type and season. Grasslands demonstrated higher interannual variation than did shrub systems. Desertification has apparently altered the seasonality of productivity in these systems; grasslands were dominated by summer growth, while sites dominated by Larrea or Prosopis tended to have higher spring ANPP. Production was frequently uncorrelated across sites of an ecosystem type, suggesting that factors other than season, regional climate, or dominant vegetation may be significant determinants of actual NPP.     

Responses of different herb life-history groups to a dominant shrub species along a dune stabilization gradient

Shrubs play a pivotal role in reversing desertification and in promoting ecological rehabilitation in severe environments. However, how the interactions between shrubs and their understorey species change during restoration remains unclear. Here, we report the results of an observational study conducted in the Mu Us Desert of northern China. This study explored how dune stabilization and the size of individual shrubs affect shrub–herb interactions. In particular, we aimed to determine how different life-history groups and performance indicators (e.g., biomass and richness) of subordinate species respond to shrub–herb interactions during dune stabilization. The shrub Artemisia ordosica had positive effects on understorey species in this dune system. The ability of the shrub to promote the growth of understorey species increased with dune stabilization, but decreased from small to large shrubs. This effect was due to an increase in the relative abundance and biomass of perennials and their higher sensitivity to the positive effects of the shrubs. In contrast, the ability of shrubs to improve the richness of understorey species decreased with dune stabilization, but increased from small to large shrubs. This effect occurred because perennials suffered high strain during recruitment in disturbed open patches of the semi-fixed dunes, particularly below small shrubs. Our results support the theory claiming that communities are sets of hidden interaction groups that have contrasting responses (neutral for annuals, but facilitative for perennials) to dominant neighbors in a single community, depending on their functional strategies. Furthermore, our study highlights the high efficiency of A. ordosica in facilitating ecological restoration of dry and disturbed sandy communities.     

Rapid genetic restoration of a keystone species exhibiting delayed demographic response

Genetic founder effects are often expected when animals colonize restored habitat in fragmented landscapes, but empirical data on genetic responses to restoration are limited. We examined the genetic response of banner‐tailed kangaroo rats (Dipodomys spectabilis) to landscape‐scale grassland restoration in the Chihuahuan Desert of New Mexico, USA. Dipodomys spectabilis is a grassland specialist and keystone species. At sites treated with herbicide to remove shrubs, colonization by D. spectabilis is slow and populations persist at low density for ≥10 years (≥6 generations). Persistence at low density and low gene flow may cause strong founder effects. We compared genetic structure of D. spectabilis populations between treated sites and remnant grasslands, and we examined how the genetic response to restoration depended on treatment age, area, and connectivity to source populations. Allelic richness and heterozygosity were similar between treated sites and remnant grasslands. Allelic richness at treated sites was greatest early in the restoration trajectory, and genetic divergence did not differ between recently colonized and established populations. These results indicated that founder effects during colonization of treated sites were weak or absent. Moreover, our results suggested founder effects were not mitigated by treatment area or connectivity. Dispersal is negatively density‐dependent in D. spectabilis, and we hypothesize that high gene flow may occur early in the restoration trajectory when density is low. Our study shows genetic diversity can be recovered more rapidly than demographic components of populations after habitat restoration and that founder effects are not inevitable for animals colonizing restored habitat in fragmented landscapes.     

Nutrient losses in runoff from grassland and shrubland habitats in Southern New Mexico: I. rainfall simulation experiments

Rainfall simulation experiments were performed in areas of semiarid grassland (Bouteloua eriopoda) and arid shrubland (Larrea tridentata) in the Chihuahuan desert of New Mexico. The objective was to compare the runoff of nitrogen (N) and phosphorus (P) from these habitats to assess whether losses of soil nutrients are associated with the invasion of grasslands by shrubs. Runoff losses from grass- and shrub-dominated plots were similar, and much less than from bare plots located in the shrubland. Weighted average concentrations of total dissolved N compounds in runoff were greatest in the grassland (1.72 mg/1) and lowest in bare plots in the shrubland (0.55 mg/1). More than half of the N transported in runoff was carried in dissolved organic compounds. In grassland and shrub plots, the total N loss was highly correlated to the total volume of discharge. We estimate that the total annual loss of N in runoff is 0.25 kg/ha/yr in grasslands and 0.43 kg/ha/yr in shrublands — consistent with the depletion of soil N during desertification of these habitats. Losses of P from both habitats were very small.     

Plants, insects and birds in semi-natural pastures in relation to local habitat and landscape factors

The preservation of remaining semi-natural grasslands in Europe has a high conservation priority. Previously, the effects of artificial fertilisation and grazing intensity on grassland animal and plant taxa have been extensively investigated. In contrast, little is known of the effects of tree and shrub cover within semi-natural grasslands and composition of habitats in the surrounding landscape on grassland taxa. We evaluated the effect that each of these factors has on species richness and community structure of vascular plants, butterflies, bumble bees, ground beetles, dung beetles and birds surveyed simultaneously in 31 semi-natural pastures in a farmland landscape in south-central Sweden. Partial correlation analyses showed that increasing proportion of the pasture area covered by shrubs and trees had a positive effect on species richness on most taxa. Furthermore, species richness of nectar seeking butterflies and bumble bees were negatively associated with grazing intensity as reflected by grass height. At the landscape level, species richness of all taxa decreased (butterflies and birds significantly so) with increasing proportion of urban elements in a 1-km² landscape area centred on each pasture, while the number of plant and bird species were lower in landscapes with large proportion of arable fields. Our results differed markedly depending on whether the focus was on species richness or community structure. Canonical correspondence analyses (CCA) showed that the abundance of most taxa was ordered along a gradient describing tree cover within pastures and proportion of arable fields in the landscape. However, subsets of grassland birds and vascular plants, respectively, showed markedly different distribution patterns along axis one of the CCA. In contrast to current conservation policy of semi-natural pastures in Sweden, our results strongly advise against using a single-taxon approach (i.e., grassland vascular plants) to design management and conservation actions in semi-natural pastures. Careful consideration of conservation values linked to the tree and shrub layers in grasslands should always precede decisions to remove trees and shrubs on the grounds of promoting richness of vascular plants confined to semi-natural grasslands. Finally, the importance of landscape composition for mobile organisms such as birds entails that management activities should focus on the wider countryside and not exclusively on single pastures.     

Effects of experimental rainfall manipulations on Chihuahuan Desert grassland and shrubland plant communities

Aridland ecosystems are predicted to be responsive to both increases and decreases in precipitation. In addition, chronic droughts may contribute to encroachment of native C₃ shrubs into C₄-dominated grasslands. We conducted a long-term rainfall manipulation experiment in native grassland, shrubland and the grass–shrub ecotone in the northern Chihuahuan Desert, USA. We evaluated the effects of 5 years of experimental drought and 4 years of water addition on plant community structure and dynamics. We assessed the effects of altered rainfall regimes on the abundance of dominant species as well as on species richness and subdominant grasses, forbs and shrubs. Nonmetric multidimensional scaling and MANOVA were used to quantify changes in species composition in response to chronic addition or reduction of rainfall. We found that drought consistently and strongly decreased cover of Bouteloua eriopoda, the dominant C₄ grass in this system, whereas water addition slightly increased cover, with little variation between years. In contrast, neither chronic drought nor increased rainfall had consistent effects on the cover of Larrea tridentata, the dominant C₃ shrub. Species richness declined in shrub-dominated vegetation in response to drought whereas richness increased or was unaffected by water addition or drought in mixed- and grass-dominated vegetation. Cover of subdominant shrubs, grasses and forbs changed significantly over time, primarily in response to interannual rainfall variability more so than to our experimental rainfall treatments. Nevertheless, drought and water addition shifted the species composition of plant communities in all three vegetation types. Overall, we found that B. eriopoda responded strongly to drought and less so to irrigation, whereas L. tridentata showed limited response to either treatment. The strong decline in grass cover and the resistance of shrub cover to rainfall reduction suggest that chronic drought may be a key factor promoting shrub dominance during encroachment into desert grassland.     

Does shrub invasion indirectly limit grass establishment via seedling herbivory? A test at grassland‐shrubland ecotones

Question: Does shrub invasion at ecotones indirectly limit grass establishment by increasing mammalian seedling herbivory? Location: Chihuahuan Desert, New Mexico, USA. Methods: We tested the hypothesis that herbivore‐related mortality of seedlings of the dominant perennial grass Bouteloua eriopoda would be highest in shrub‐dominated portions of grassland‐shrubland ecotones. We tested the hypothesis in two Chihuahuan Desert sites featuring similar shrub encroachment patterns but different shrub species, grass cover, and different abundances of small mammals. Within each site we transplanted B. eriopoda seedlings to grass‐dominated, middle, and shrub‐dominated positions of replicate ecotones during the time of year (mid‐summer) when they would naturally appear and monitored seedling fates. We estimated population size/activity of putative small mammal herbivores. Results: Seedlings were killed by mammals in greater numbers in shrubland than in grassland or middle ecotone positions at the site with large herbivore numbers. At the site with low herbivore numbers, most seedlings were killed in middle ecotone positions. The abundance patterns of herbivores did not parallel patterns of seedling herbivory across the ecotones or between sites. Conclusions: Seedling herbivory is an important process and is related to vegetation composition, but the mechanisms underlying the relationship are not clear. We speculate that variation in small mammal foraging behavior may contribute to seedling herbivory patterns. Restoration strategies in the Chihuahuan Desert need to account for the abundance and/or behavior of native herbivores.     

Resource Islands Predict the Distribution of Heterotrophic Bacteria in Chihuahuan Desert Soils

The resource island hypothesis predicts that soil resources such as nitrogen, phosphorus, and water will be distributed evenly in grasslands but have a patchy distribution focused around plants in shrublands. This hypothesis predicts that microorganism numbers will follow resources and be (i) evenly distributed in grasslands, (ii) concentrated around individual plants in shrublands, and (iii) higher where resources are higher when comparing the same vegetation type. This study enumerated total heterotrophic bacteria and a subset of these, the nitrogen-efficient guild (NEG), in three shrublands (playa fringe mesquite, tar bush, and creosote) and two grasslands (playa and bajada). Both heterotrophs and NEG members followed the distribution pattern predicted by the resource island hypothesis. There were no significant differences in heterotroph or NEG numbers comparing at-plant and interplant samples for both the playa and bajada grasslands. Furthermore, populations were generally higher in nutrient-rich playa grasslands than nutrient-poor bajada grasslands. In contrast, both heterotroph and NEG numbers were higher at shrubs than between shrubs in all three shrub sites. These results suggest that resource abundance in resource islands predicts the distribution of heterotrophic bacterial numbers in desert soils.     

Grassland area determines beetle assemblage dissimilarity from surrounding floodplain forest

Patch size is known to affect biodiversity in fragmented landscapes, but is usually examined in systems where the surrounding matrix habitat is unfavourable. We examined beetle diversity in a floodplain ecosystem that is characterised by naturally occurring grassland patches within a dominant matrix of contrasting yet habitable forest. We asked whether differences in the beetle assemblage between grassland and forest vegetation depended on the area of the grassland patch, which is a function of its flooding frequency and duration: smaller grasslands tend to be higher on the floodplain and are flooded less often and for shorter periods than larger grasslands. We found a negative relationship between grassland area and beetle abundance and species richness, and a positive relationship between grassland area and compositional dissimilarity from the surrounding forest. As expected, we found an overall difference in composition between forest and grassland assemblages, with five beetle species more common in the grasslands. Our study indicates that floodplain grasslands not only support beetle assemblages that are distinct from the surrounding forest, but that assemblages from the larger grasslands are compositionally more distinct than those from smaller grasslands. A likely cause of this pattern is the reduced edge effects and greater environmental contrast between forest and large grasslands that may be exposed to greater variation in local climate. Ongoing changes to flood regimes and potential encroachment of forest plants may decrease grassland area in the future, which may reduce spatial heterogeneity in the insect community in this unique floodplain ecosystem.     

Post-Fire Resource Redistribution in Desert Grasslands: A Possible Negative Feedback on Land Degradation

Desert grasslands, which are very sensitive to external drivers like climate change, are areas affected by rapid land degradation processes. In many regions of the world the common form of land degradation involves the rapid encroachment of woody plants into desert grasslands. This process, thought to be irreversible and sustained by biophysical feedbacks of global desertification, results in the heterogeneous distribution of vegetation and soil resources. Most of these shrub-grass transition systems at the desert margins are prone to disturbances such as fires, which affect the interactions between ecological, hydrological, and land surface processes. Here we investigate the effect of prescribed fires on the landscape heterogeneity associated with shrub encroachment. Replicated field manipulation experiments were conducted at a shrub-grass transition zone in the northern Chihuahuan desert (New Mexico, USA) using a combination of erosion monitoring techniques, microtopography measurements, infiltration experiments, and isotopic studies. The results indicate that soil erosion is more intense in burned shrub patches compared to burned grass patches and bare interspaces. This enhancement of erosion processes, mainly aeolian, is attributed to the soil–water repellency induced by the burning shrubs, which alters the physical and chemical properties of the soil surface. Further, we show that by enhancing soil erodibility fires allow erosion processes to redistribute resources accumulated by the shrub clumps, thereby leading to a more homogeneous distribution of soil resources. Thus fires counteract or diminish the heterogeneity-forming dynamics of land degradation associated with shrub encroachment by enhancing local-scale soil erodibility. Author Contributions  SR—Conceived of or designed study, performed research, analyzed data, wrote the paper; PD—Conceived of or designed study, performed research, wrote the paper; LW—Performed research, analyzed data; GO—Contributed new methods, analyzed data; SC—Conceived of or designed study; CW—Performed research, contributed new methods or models; and SM—Contributed new methods or models.     

Carabid beetles as indicators for shrub encroachment in dry grasslands

Semi-natural grasslands are key habitats for biodiversity conservation in Central Europe. Shrub encroachment is one of the most threatening drivers of grassland degradation and affects soil properties, microclimate, and vegetation with possible impacts on higher trophic levels. We aimed to analyse the impact of shrub encroachment with broom (Cytisus scoparius) on carabid beetle diversity, species composition, and functional traits. In a field study on dry grasslands on the island of Hiddensee (Germany) we studied 15 sites along a gradient of increasing broom encroachment and classified them into three dry grassland types with low, medium, and high shrub cover. Our results provide evidence that shrub encroachment initially has positive effects on species richness and activity densities of dry grassland carabids. Carabid species composition differed among differently shrub-covered dry grassland types, and sites with low and high shrub cover were each characterised by unique carabid assemblages. The species composition of sites with a medium shrub biomass had a transitional character and contained species which are typical for open dry grassland, but also shared species with sites with a high shrub cover. Among functional trait parameters investigated, especially the body size of carabid beetles was related to environmental parameters associated with shrub encroachment. Body size was positively correlated to shrub biomass and soil humidity, but negatively to temperature. Eurytopy values of carabids were related to high litter cover, i.e. habitat generalist (eurytopic) species mainly occurred in densely shrub-encroached sites. In order to preserve unique carabid assemblages of open dry grasslands with stenotopic and smaller species, it is most important to prevent a shrub encroachment higher than about 60% cover. For management we suggest extensive grazing (by cattle, sheep or horses) to prevent shrub encroachment on dry grasslands. In areas with high shrub cover additionally the use of goats or mechanical removal of shrubs might be necessary.