Shrub invasion decreases diversity and alters community stability in northern Chihuahuan Desert plant communities.
ABSTRACT: BACKGROUND:Global climate change is rapidly altering species range distributions and interactions within communities. As ranges expand, invading species change interactions in communities which may reduce stability, a mechanism known to affect biodiversity. In aridland ecosystems worldwide, the range of native shrubs is expanding as they invade and replace native grassland vegetation with significant consequences for biodiversity and ecosystem functioning. METHODOLOGY:We used two long-term data sets to determine the effects of shrub encroachment by Larrea tridentata on subdominant community composition and stability in formerly native perennial grassland dominated by Bouteloua eriopoda in New Mexico, USA. PRINCIPAL FINDINGS:Our results indicated that Larrea invasion decreased species richness during the last 100 years. We also found that over shorter temporal scales species-poor subdominant communities in areas invaded by Larrea were less stable (more variable in time) compared to species rich communities in grass-dominated vegetation. Compositional stability increased as cover of Bouteloua increased and decreased as cover of Larrea increased. SIGNIFICANCE:Changes in community stability due to altered interspecific interactions may be one mechanism by which biodiversity declines in grasslands following shrub invasion. As global warming increases, shrub encroachment into native grasslands worldwide will continue to alter species interactions and community stability both of which may lead to a decline in biodiversity.
Project description:Shrub canopies in semi-arid environments often produce positive effects on soil fertility, and on the richness and biomass of understorey plant communities. However, both positive and negative effects of shrub encroachment on plant and soil attributes have been reported at the landscape-level. The contrasting results between patch- and landscape-level effects in shrublands could be caused by differences in the degree of shrub encroachment or grazing pressure, both of which are likely to reduce the ability of individual shrubs to ameliorate their understorey environment.We examined how grazing and shrub encroachment (measured as landscape-level shrub cover) influence patch-level effects of shrubs on plant density, biomass and similarity in species composition between shrub understories and open areas, and on soil stability, nutrient cycling, and infiltration in two semi-arid Australian woodlands.Individual shrubs had consistently positive effects on all plant and soil variables (average increase of 23% for all variables). These positive patch-level effects persisted with increasing shrub cover up to our maximum of 50% cover. Heavy grazing negatively affected most of the variables studied (average decline of 11%). It also altered, for some variables, how individual shrubs affected their sub-canopy environment with increasing shrub cover. Thus for species density, biomass and soil infiltration, the positive effect of individual shrubs with increasing shrub cover diminished under heavy grazing. SYNTHESIS:Our study refines predictions of the effects of woody encroachment on ecosystem structure and functioning by showing that heavy grazing, rather than differences in shrub cover, explains the contrasting effects on ecosystem structure and function between individual shrubs and those in dense aggregations. We also discuss how species-specific traits of the encroaching species, such as their height or its ability to fix N, might influence the relationship between their patch-level effects and their cover within the landscape.
Project description:We used a functional trait-based approach to assess the impacts of aridity and shrub encroachment on the functional structure of Mediterranean dryland communities (functional diversity (FD) and community-weighted mean trait values (CWM)), and to evaluate how these functional attributes ultimately affect multifunctionality (i.e. the provision of several ecosystem functions simultaneously). Shrub encroachment (the increase in the abundance/cover of shrubs) is a major land cover change that is taking place in grasslands worldwide. Studies conducted on drylands have reported positive or negative impacts of shrub encroachment depending on the functions and the traits of the sprouting or nonsprouting shrub species considered. FD and CWM were equally important as drivers of multifunctionality responses to both aridity and shrub encroachment. Size traits (e.g. vegetative height or lateral spread) and leaf traits (e.g. specific leaf area and leaf dry matter content) captured the effect of shrub encroachment on multifunctionality with a relative high accuracy (r(2) = 0.63). FD also improved the resistance of multifunctionality along the aridity gradient studied. Maintaining and enhancing FD in plant communities may help to buffer negative effects of ongoing global environmental change on dryland multifunctionality.
Project description: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.
Project description:In many cultural landscapes, the abandonment of traditional grazing leads to encroachment of pastures by woody plants, which reduces habitat heterogeneity and impacts biodiversity typical of semi-open habitats. We developed a framework of mutually interacting spatial models to locate areas where shrub encroachment in Alpine treeline ecosystems deteriorates vulnerable species' habitat, using black grouse Tetrao tetrix (L.) in the Swiss Alps as a study model. Combining field observations and remote-sensing information we 1) identified and located the six predominant treeline vegetation types; 2) modelled current black grouse breeding habitat as a function thereof so as to derive optimal habitat profiles; 3) simulated from these profiles the theoretical spatial extension of breeding habitat when assuming optimal vegetation conditions throughout; and used the discrepancy between (2) and (3) to 4) locate major aggregations of homogeneous shrub vegetation in otherwise suitable breeding habitat as priority sites for habitat restoration. All six vegetation types (alpine pasture, coniferous forest, Alnus viridis (Chaix), Rhododendron-dominated, Juniperus-dominated and mixed heathland) were predicted with high accuracy (AUC >0.9). Breeding black grouse preferred a heterogeneous mosaic of vegetation types, with none exceeding 50% cover. While 15% of the timberline belt currently offered suitable breeding habitat, twice that fraction (29%) would potentially be suitable when assuming optimal shrub and ground vegetation conditions throughout the study area. Yet, only 10% of this difference was attributed to habitat deterioration by shrub-encroachment of dense heathland (all types 5.2%) and Alnus viridis (4.8%). The presented method provides both a general, large-scale assessment of areas covered by dense shrub vegetation as well as specific target values and priority areas for habitat restoration related to a selected target organism. This facilitates optimizing the spatial allocation of management resources in geographic regions where shrub encroachment represents a major biodiversity conservation issue.
Project description:Regime shift from grasslands to shrub-dominated landscapes occur worldwide driven by altered land-use and climate change, affecting landscape function, biodiversity, and productivity. Warming winter temperatures are a main driver of expansion of the native, evergreen shrub, Morella cerifera, in coastal landscapes. Shrub establishment in these habitats alters microclimate, but little is known about seasonal differences and microclimate variance. We assessed influence of shrubs on microclimate variance, community composition, and community physiological functioning across three vegetation zones: grass, transitional, and shrub in a coastal grassland. Using a novel application of a time-series analysis, we interpret microclimatic variance modification and elucidate mechanisms of shrub encroachment at the Virginia Coast Reserve, Long-Term Ecological Research site. As shrub thickets form, diversity is reduced with little grass/forb cover, while transpiration and annual productivity increase. Shrub thickets significantly reduced temperature variance with a positive influence of one day on the next in maximum air, minimum air, and maximum ground temperature. We also show that microclimatic temperature moderation reduces summer extreme temperatures in transition areas, even before coalescence into full thickets. Encroachment of Morella cerifera on the Virginia barrier islands is driven by reduced local exposure to cold temperatures and enhanced by abiotic microclimatic modification and biotic physiological functioning. This shift in plant community composition from grassland to shrub thicket alters the role of barrier islands in productivity and can have impacts on the natural resilience of the islands.
Project description:Background:Changes in aboveground community composition and diversity following shrub encroachment have been studied extensively. Recently, shrub encroachment was associated with differences in belowground bacterial communities relative to non-encroached grassland sites hundreds of meters away. This spatial distance between grassland and shrub sites left open the question of how soil bacterial communities associated with different vegetation types might differ within the same plot location. Methods:We examined soil bacterial communities between shrub-encroached and adjacent (one m apart) grassland soils in Chinese Inner Mongolian, using high-throughput sequencing method (Illumina, San Diego, CA, USA). Results:Shrub-encroached sites were associated with dramatic restructuring of soil bacterial community composition and predicted metabolic function, with significant increase in bacterial alpha-diversity. Moreover, bacterial phylogenic structures showed clustering in both shrub-encroached and grassland soils, suggesting that each vegetation type was associated with a unique and defined bacterial community by niche filtering. Finally, soil organic carbon (SOC) was the primary driver varied with shifts in soil bacterial community composition. The encroachment was associated with elevated SOC, suggesting that shrub-mediated shifts in SOC might be responsible for changes in belowground bacterial community. Discussion:This study demonstrated that shrub-encroached soils were associated with dramatic restructuring of bacterial communities, suggesting that belowground bacterial communities appear to be sensitive indicators of vegetation type. Our study indicates that the increased shrub-encroached intensity in Inner Mongolia will likely trigger large-scale disruptions in both aboveground plant and belowground bacterial communities across the region.
Project description:Shrub encroachment into arid grasslands has been associated with reduced grass abundance, increased soil erosion, and local declines in biodiversity. Livestock overgrazing and the associated reduction of fine fuels has been a primary driver of shrub encroachment in the southwestern United States, but shrublands continue to persist despite livestock removal and grassland restoration efforts. We hypothesized that an herbivory feedback from native mammals may contribute to continued suppression of grasses after the removal of livestock. Our herbivore exclusion experiment in southeastern Arizona included five treatment levels and allowed access to native mammals based on their relative body size, separating the effects of rodents, lagomorphs, and mule deer. We included two control treatments and replicated each treatment 10 times (n = 50). We introduced uniform divisions of lawn sod (Cynodon dactylon) into each exclosure for 24-hr periods prior to (n = 2) and following (n = 2) the monsoon rains and used motion-activated cameras to document herbivore visitations. In the pre-monsoon trials, treatments that allowed lagomorph access had less sod biomass relative to other treatments (p < 0.001), averaging 44% (SD 36%) and 29% (SD 45%) remaining biomass after the 24-hr trial periods. Following the onset of monsoons, differences in remaining biomass among treatments disappeared. Desert cottontails (Sylvilagus audubonii) were detected more frequently than any of the other 11 herbivore species present at the site, accounting for 83% of detections during the pre-monsoon trials. Significantly more (p < 0.001) desert cottontails were detected during the pre-monsoon trials (2,077) compared to the post-monsoon trials (174), which coincided with biomass removal from lagomorph accessible treatments. We conclude that desert cottontails are significant consumers of herbaceous vegetation in shrub-encroached arid grasslands and they, along with other native herbivores, may act as a biotic feedback contributing to the competitive advantage and persistence of shrubs.
Project description:In recent decades, pastoral abandonment has produced profound ecological changes in the Alps. In particular, the reduction in grazing has led to extensive shrub encroachment of semi-natural grasslands, which may represent a threat to open habitat biodiversity. To reverse shrub encroachment, we assessed short-term effects of two different pastoral practices on vegetation and dung beetles (Coleoptera, Scarabaeoidea). Strategic placement of mineral mix supplements (MMS) and arrangement of temporary night camp areas (TNCA) for cattle were carried out during summer 2011 in the Val Troncea Natural Park, north-western Italian Alps. In 2012, one year after treatment, a reduction in shrub cover and an increase in bare ground cover around MMS sites was detected. A more intense effect was detected within TNCA through increases in forage pastoral value, and in the cover and height of the herbaceous layer. Immediately after treatment, changes in dung beetle diversity (total abundance, species richness, Shannon diversity, taxonomic and functional diversity) showed a limited disturbance effect caused by high cattle density. In contrast, dung beetle diversity significantly increased one year later both at MMS and TNCA sites, with a stronger effect within TNCA. Multivariate Regression Trees and associated Indicator Value analyses showed that some ecologically relevant dung beetle species preferred areas deprived of shrub vegetation. Our main conclusions are: i) TNCA are more effective than MMS in terms of changes to vegetation and dung beetles, ii) dung beetles respond more quickly than vegetation to pastoral practices, and iii) the main driver of the rapid response by dung beetles is the removal of shrubs. The resulting increase in dung beetle abundance and diversity, which are largely responsible for grassland ecosystem functioning, may have a positive effect on meso-eutrophic grassland restoration. Shrub encroachment in the Alps may therefore be reversed, and restoration of grassland enhanced, by using appropriate pastoral practices.
Project description:Altered disturbance regimes, increasing atmospheric CO2, and other processes have increased woody cover and homogenized vegetation in savannas across the planet. African savannas with extensive versus minimal woody cover often have vastly different animal communities. However, we lack a clear mechanistic understanding of why animal communities are changing with vegetation structure. Our goal for this study was to understand how vegetation structure in an African savanna shaped the perceived predation risk of small mammals, hence affecting their activity. Using a reciprocal measure of standard giving-up-densities, amount of food eaten, we found sharp declines in rodents' perceived predation risk and increased rodent activity underneath shrub cover. This response was consistent across species; however, species showed subtle differences in their responses to grassy vegetation. Our findings suggest that areas of minimal or extensive shrub cover (shrub encroachment) may be homogenizing rodents' perceptions of predation risk and thus shaping their use of space.
Project description:One of the most important changes taking place in drylands worldwide is the increase of the cover and dominance of shrubs in areas formerly devoid of them (shrub encroachment). A large body of research has evaluated the causes and consequences of shrub encroachment for both ecosystem structure and functioning. However, there are virtually no studies evaluating how shrub encroachment affects the ability of ecosystems to maintain multiple functions and services simultaneously (multifunctionality). We aimed to do so by gathering data from ten ecosystem functions linked to the maintenance of primary production and nutrient cycling and storage (organic C, activity of ?-glucosidase, pentoses, hexoses, total N, total available N, amino acids, proteins, available inorganic P and phosphatase activity), and summarizing them in a multifunctionality index (M). We assessed how climate, species richness, anthropic factors (distance to the nearest town, sandy and asphalted road, and human population in the nearest town at several historical periods) and encroachment by sprouting shrubs impacted both the functions in isolation and M along a regional (ca. 350 km) gradient in Mediterranean grasslands and shrublands dominated by a non-sprouting shrub. Values of M were higher in those grasslands and shrublands containing sprouting shrubs (43% and 62%, respectively). A similar response was found when analyzing the different functions in isolation, as encroachment by sprouting shrubs increased functions by 2%-80% compared to unencroached areas. Encroachment was the main driver of changes in M along the regional gradient evaluated, followed by anthropic factors and species richness. Climate had little effects on M in comparison to the other factors studied. Similar responses were observed when evaluating the functions in isolation. Overall, our results showed that M was higher at sites with higher sprouting shrub cover, longer distance to roads and higher perennial plant species richness. Our study is the first documenting that ecosystem multifunctionality in shrublands is enhanced by encroaching shrubs differing in size and leaf attributes. Our findings reinforce the idea that encroachment effects on ecosystem functioning cannot be generalized, and that are largely dependent on the traits of the encroaching shrub relative to those of the species being replaced.