Ecosystem responses to woody plant encroachment in a semiarid savanna rangeland

Woody plant encroachment alters the structure and function of rangeland ecosystems. The objective of this study was to explore the association between woody plant encroachment and various ecosystem properties (i.e. vascular plant species diversity, richness, evenness, soil organic matter, herbaceous biomass, leaf litter and bare ground cover) in a semiarid savanna rangeland, and also to test whether the relationships were influenced by woody species composition, elevation and site. We carried out a vegetation survey in four rangeland sites in the lower Omo region of southwestern Ethiopia, and regressed each one of the ecosystem properties, separately, against woody plant density, elevation and site using multiple linear regressions. We found that vascular plant species diversity, richness and evenness increased with woody plant density, most likely due to increased spatial heterogeneity and soil microclimate improvement. Bare ground cover increased significantly, whereas herbaceous biomass and soil organic matter did not respond to woody encroachment. In a subsequent investigation, we used a redundancy analysis to assess whether ecosystem properties were influenced by the identity of encroaching woody plant species. Species diversity and richness responded positively to Lannea triphylla, whereas leaf litter responded positively to Grewia tenax and G. villosa. Our findings suggest that woody plant encroachment in a semiarid rangeland does alter ecosystem properties. However, its impact is highly variable, influenced by a set of factors including the level of encroachment and identity of encroaching woody species.     

Elevated atmospheric CO2: a nurse plant substitute for oak seedlings establishing in old fields

It has been hypothesized that elevated levels of atmospheric CO₂ (eCO₂) may facilitate the encroachment of woody plants into grasslands by reducing water stress. In east‐central Minnesota, sandy soils frequently create drought conditions for plants, and water limitation inhibits the establishment of oaks into old fields situated on these soils. Some have argued that eCO₂ should slow secondary succession by favoring fast‐growing early successional species. However, if oak encroachment into old fields is being inhibited by water stress, then eCO₂ could accelerate old‐field succession in this region. The purpose of this study was to test the hypothesis that eCO₂ will increase the establishment success of oak seedlings in an old field environment. The study was conducted with CO₂ levels controlled by free air CO₂ enrichment (FACE). In May 1999, four oak (Quercus ellipsoidalis) acorns were planted in each of 24 plots in each of six experimental FACE rings (n= 576), three of which received elevated levels (550 ppm) of CO₂. Half the plots in each ring were weeded during the first three summers of the experiment. In summer 2000, water input was manipulated during a 3.5‐week period, during which half the plots received regular watering while the other half received no water. Summer 2001 was dry, receiving 35% less rainfall than the mean level. Under hot and dry conditions, eCO₂ increased soil water levels in unweeded plots and enhanced oak establishment (survival and growth) in weeded plots. In 2006, after the eighth growing season following planting, survival was five times greater under elevated than ambient CO₂. The results showed that under hot and dry conditions, eCO₂ can act like a nurse plant for tree seedlings growing in bare and unshaded areas, increasing seedling survival and growth, and thereby expanding the establishment window for trees encroaching into a grassland environment.     

Trade‐offs between savanna woody plant diversity and carbon storage in the Brazilian Cerrado

Incentivizing carbon storage can be a win‐win pathway to conserving biodiversity and mitigating climate change. In savannas, however, the situation is more complex. Promoting carbon storage through woody encroachment may reduce plant diversity of savanna endemics, even as the diversity of encroaching forest species increases. This trade‐off has important implications for the management of biodiversity and carbon in savanna habitats, but has rarely been evaluated empirically. We quantified the nature of carbon‐diversity relationships in the Brazilian Cerrado by analyzing how woody plant species richness changed with carbon storage in 206 sites across the 2.2 million km² region at two spatial scales. We show that total woody plant species diversity increases with carbon storage, as expected, but that the richness of endemic savanna woody plant species declines with carbon storage both at the local scale, as woody biomass accumulates within plots, and at the landscape scale, as forest replaces savanna. The sharpest trade‐offs between carbon storage and savanna diversity occurred at the early stages of carbon accumulation at the local scale but the final stages of forest encroachment at the landscape scale. Furthermore, the loss of savanna species quickens in the final stages of forest encroachment, and beyond a point, savanna species losses outpace forest species gains with increasing carbon accumulation. Our results suggest that although woody encroachment in savanna ecosystems may provide substantial carbon benefits, it comes at the rapidly accruing cost of woody plant species adapted to the open savanna environment. Moreover, the dependence of carbon‐diversity trade‐offs on the amount of savanna area remaining requires land managers to carefully consider local conditions. Widespread woody encroachment in both Australian and African savannas and grasslands may present similar threats to biodiversity.     

Shrub encroachment alters composition and diversity of ant communities in abandoned grasslands of western Carpathians

Invasions of woody species into grasslands abandoned by agriculture are a global phenomenon, but their effects on diversity of other taxa have been rarely investigated across both regional and local scales. We quantified how shrub encroachment affected the activity, composition, and diversity of ant communities in managed and abandoned grasslands in western Carpathians of Central Europe across four regions and four shrub encroachment stages in each region. We surveyed ant communities on 48 sites in total, with each encroachment stage replicated three times in each region and twelve times overall. We used pitfall traps to sample ants over three years (2008, 2009, 2011) and identified 9,254 ant workers belonging to 33 species in total. Although the epigaeic activity and composition of ant communities varied with region, abandoned grasslands supported a greater species richness of ants than managed grasslands regardless of the region, and especially so in more advanced shrub encroachment stages. Since the woody colonization within grasslands was moderate even in the advanced encroachment stages (on average ~40 % of grassland colonized by woody species), it allowed coexistence of forest specialists (e.g. Temnothorax crassispinus) with species typical of open grasslands, thus increasing overall ant diversity. Managed grasslands were not only less species rich compared to abandoned grasslands, but they were characterized by different species (e.g. Lasius niger, Myrmica rugulosa). The differences in ant communities between managed and abandoned grasslands are likely to cause differences in ecological functions mediated by ants (e.g. predation of arthropods or plant seed dispersal).     

Influence of conspecific and heterospecific adults on riparian tree species establishment during encroachment of a humid palm savanna

Woody plant encroachment of savanna ecosystems has been related to altered disturbance regimes, mainly fire suppression and herbivore exclusion. In contrast, neighbourhood interactions among resident and colonising woody species have received little attention, despite their likely influence on the pattern and rate of tree establishment. We examined how resident palm trees (Butia yatay) and established adults of two riparian forest tree species (Allophylus edulis and Sebastiania commersoniana) influenced seed arrival and seedling performance of the latter two species in a humid savanna of east-central Argentina. Seed traps and seedlings of both riparian species were placed in herbaceous openings, and beneath palm, conspecific and heterospecific adult trees in two unburned savanna patches, and were monitored for 2 years. Only seeds of the bird-dispersed Allophylus arrived in palm microsites, yet survival of Allophylus seedlings near adult palms was limited by animal damage through trampling and burrowing, a non-trophic mechanism of apparent competition. Seeds of both riparian species dispersed into conspecific microsites, although adult trees selectively reduced growth of conspecific seedlings, a pattern consistent with the “escape hypothesis”. Further, survival of Sebastiania increased in the moister Allophylus microsites, suggesting a one-way facilitative interaction between woody colonisers. Our results indicate that dispersal facilitation by resident savanna trees may be critical to riparian species invasion after fire suppression. Distance-dependent effects of conspecific and heterospecific adult trees could contribute to shape the subsequent dynamics of woody seedling establishment. Overall, we show that indirect interactions can play a prominent role in savanna encroachment by non-resident woody species.     

Effects of termite mounds on composition,functional types and traits of plant communities in Pendjari Biosphere Reserve (Benin,West Africa)

Understanding the role of termite mounds in biodiversity and ecosystem functioning is a priority for the management of tropical terrestrial protected areas dominated by savannahs. This study aimed to assess the effects of termite mounds on the diversity of plant functional types (PFTs) and herbaceous’ net aboveground primary productivity (NAPP) in plant communities (PCs) of the Pendjari Biosphere Reserve. PCs were identified through canonical correspondence analysis performed on 96 phytosociological ‘relevés’ realized in plots of 900 m². PFTs’ diversity was compared between savannahs and mounds’ plots using generalized linear models. In each plot, 7 m² subplots were harvested and NAPP was determined. Linear mixed models were performed to assess change in herbaceous NAPP regarding species richness, graminoids’ richness, specific leaf area and termite mounds. There is no specific plant community related to mounds. However, the occurrence of termite mounds induced an increase of woody and forbs diversity while the diversity of legumes and graminoids decreased. These diversity patterns led to decreasing of PCs’ NAPP. This study confirms that termite‐induced resource heterogeneity supports niche differentiation theory and increased savannah encroachment by woody species.     

Forest encroachment into a Californian grassland: examining the simultaneous effects of facilitation and competition on tree seedling recruitment

Competition and facilitation are both considered major factors affecting the structure of plant assemblages, yet few studies have quantified positive, negative, and net effects simultaneously. In this study, we investigated the positive, negative, and net effects of tree saplings on the encroachment of two tree species, Douglas fir (Pseudotsuga menziesii) and tanoak (Lithocarpus densiflora), into a coastal California grassland. The study involved three components: sampling the spatial distributions of P. menziesii and L. densiflora in the grasslands, a field experiment examining seedling survival in different grassland environments, and a greenhouse experiment examining the effects of soil moisture on early seedling performance. The field experiment was conducted over a 2-year period, using Pseudotsuga in 2002 and both species in 2003. Seedlings were separated into four treatment groups: those planted in open grassland, in shaded grassland, under artificial (plastic) conifer saplings, and under natural Pseudotsuga saplings. Air temperature, relative humidity, soil moisture, incident radiation levels and fog water inputs were measured for each treatment group in 2003. In the greenhouse experiment, Pseudotsuga and Lithocarpus seedlings were grown for 13 weeks in watering treatments simulating the summer soil moisture conditions of the open grasslands and under Pseudotsuga saplings. Surveys of naturally established seedlings found that Lithocarpus occurred only under Pseudotsuga saplings, while most Pseudotsuga seedlings were located near but not directly under conspecific saplings. In the field experiment, positive effects of tree saplings were much larger than negative effects, resulting in strong net facilitation of seedling establishment. Survival for both species was always higher under the plastic and live trees than in the open or shade plots. The primary mechanism facilitating seedling survival appeared to be increased soil moisture caused by input of fog precipitation coupled with reduced microsite evaporation. The greenhouse experiment further showed that soil moisture strongly affected seedling performance, with both species having much higher photosynthetic rates in the higher moisture treatment. In the lower moisture treatment, Pseudotsuga seedlings had higher photosynthetic rates and stomatal conductance than Lithocarpus, suggesting they may be able to better tolerate the environmental conditions found in the open grasslands. Our combined results suggest that rate and patterning of woody plant encroachment can be strongly influenced by facilitation and that fog precipitation may play a key role in plant interactions.     

Positive versus negative environmental impacts of tree encroachment in South Africa

Woody plant encroachment in grasslands is a worldwide phenomenon. Despite many studies, the consequences of woody plant encroachment on sub-canopy vegetation and soil properties are still unclear. To better understand the impacts of trees on grassland properties we examined the following questions using a mountainous sub-tropical grassland of South Africa encroached by an indigenous tree, Acacia sieberiana as a case study: (1) Do trees increase sub-canopy herbaceous diversity, quality and biomass and soil nitrogen content? (2) Do large trees have a stronger effect than medium-sized trees on grass and soil properties? (3) Does the impact of trees change with the presence of livestock and position of trees in a catena? We studied grass and non-graminoid species diversity and biomass, grass quality and soil properties during the wet season of 2009. Nitrogen in grass leaves, soil cation exchange capacity and calcium and magnesium ion concentrations in the soil increased under tall Acacia versus open areas. Medium-sized Acacia decreased the gross energy content, digestibility and neutral detergent fibre of grasses but increased the species richness of non-graminoids. Tall and medium Acacia trees were associated with the presence of Senecio inaequidens, an indigenous species that is toxic to horses and cattle. The presence of livestock resulted in a decrease in herbaceous root biomass and an increase in soil carbon and leaf biomass of grass under Acacia. Tree position in the catena did not modify the impact of trees on the herbaceous layer and soil properties. For management of livestock we recommend retaining tall Acacia trees and partially removing medium-sized Acacia trees because the latter had negative effects on grass quality.     

Contrasting effects of grazing on the early stages of woody encroachment in a Neotropical savanna

Woody encroachment in savannas represents an ecological process of current global interest given its negative impact on ecosystem functioning, particularly on forage production. Traditional savanna models propose competition and niche differentiation as the main mechanisms allowing tree-grass coexistence. Demographic models, instead, propose abiotic and biotic factors as bottlenecks controlling vital rates and transitions from seeds to adult trees. The role played by domestic grazing on woody encroachment is yet controversial. Here, using a multistage tree life approach, we combine both models and evaluate the role of grazing and herbaceous vegetation on woody recruitment in a Neotropical savanna dominated by Vachellia caven, a successful and widely spread encroacher tree species. We performed three experiments to evaluate seed predation, seedling emergence and survival of V. caven by manipulating cattle grazing (grazed and ungrazed areas) and herbaceous vegetation presence (vegetated and unvegetated). Finally, we combined the results of the three experiments to estimate the probability of plant recruitment across these experimental factors. Grazing decreased seed predation by half, did not modify seedling emergence and decreased seedling survival. Herbaceous vegetation did not affect seed predation nor seedling emergence rate, but increased seedling survival. Overall, the net effect of grazing on V. caven recruitment was neutral since the increase in seed availability due to the reduction in seed predation rate was compensated by the negative effect of grazing on seedling survival. Our analysis revealed that cattle grazing and herbaceous vegetation had contrasting effects on the seed and seedling life stages. We propose that in order to restrain the early stages of encroachment, cattle grazing pressure could be managed following the seasonality of demographic tree transitions. Through rotational grazing amongst paddocks, stocking rates could be relaxed during the primary dispersal stage to maximize granivory, and then increased to enhance the chance of seedling consumption and trampling.     

Local native plant diversity responds to habitat loss and fragmentation over different time spans and spatial scales

Separating the threats that habitat loss and habitat fragmentation pose to biodiversity is challenging because these processes usually occur simultaneously. Additionally, their importance may be underestimated due to time-delayed extinction. In central Texas savannas, woody plant encroachment reduces the amount of habitat available to herbaceous species while fragmenting remaining habitat. We examined the relationships between present species richness and present and past habitat amount and fragmentation (measured as fractal dimension) using a series of aerial photographs taken over nearly 60 years. We show that woody plant encroachment, a common phenomenon in savannas worldwide, reduces the diversity of herbaceous vegetation through both habitat loss and fragmentation. Habitat loss has the strongest impact on species richness over short time spans and small spatial scales in these savannas. Habitat fragmentation, however, has the strongest impact over longer time spans and larger spatial scales and generates long-term extinction debts. We also demonstrate that examining habitat loss and habitat fragmentation across different time periods and at different spatial scales is essential for understanding their joint and individual effects on plant community composition.     

Shrub expansion stimulates soil C and N storage along a coastal soil chronosequence

Expansion of woody vegetation in grasslands is a worldwide phenomenon with implications for C and N cycling at local, regional and global scales. Although woody encroachment is often accompanied by increased annual net primary production (ANPP) and increased inputs of litter, mesic ecosystems may become sources for C after woody encroachment because stimulation of soil CO₂ efflux releases stored soil carbon. Our objective was to determine if young, sandy soils on a barrier island became a sink for C after encroachment of the nitrogen‐fixing shrub Morella cerifera, or if associated stimulation of soil CO₂ efflux mitigated increased litterfall. We monitored variations in litterfall in shrub thickets across a chronosequence of shrub expansion and compared those data to previous measurements of ANPP in adjacent grasslands. In the final year, we quantified standing litter C and N pools in shrub thickets and soil organic matter (SOM), soil organic carbon (SOC), soil total nitrogen (TN) and soil CO₂ efflux in shrub thickets and adjacent grasslands. Heavy litterfall resulted in a dense litter layer storing an average of 809 g C m^?2 and 36 g N m^?2. Although soil CO₂ efflux was stimulated by shrub encroachment in younger soils, soil CO₂ efflux did not vary between shrub thickets and grasslands in the oldest soils and increases in CO₂ efflux in shrub thickets did not offset contributions of increased litterfall to SOC. SOC was 3.6–9.8 times higher beneath shrub thickets than in grassland soils and soil TN was 2.5–7.7 times higher under shrub thickets. Accumulation rates of soil and litter C were highest in the youngest thicket at 101 g m^?2 yr^?1 and declined with increasing thicket age. Expansion of shrubs on barrier islands, which have low levels of soil carbon and high potential for ANPP, has the potential to significantly increase ecosystem C sequestration.     

Mechanical Land Clearing to Promote Establishment of Coastal Sandplain Grassland and Shrubland Communities

The decline in grasslands and other species‐rich early successional habitats on the coastal sandplains of the northeastern United States has spurred management to increase the area of these declining plant communities. We mechanically removed overstory oak and applied seed from a nearby sandplain grassland on the island of Martha’s Vineyard, Massachusetts, to evaluate this technique for creating an open oak community able to support sandplain herbaceous species. We compared vegetation structure and composition before and after clearing in an area of total tree removal (clearcutting), an area where 85% of tree basal area was removed (savanna cutting), and in adjacent coastal oak forest. Plant responses to clearcutting and savanna cutting were similar. Sandplain herbs colonized at high frequencies after seeding and increased herbaceous cover from less than 7% before clearing to 22–38% three growing seasons later. Pennsylvania sedge (Carex pensylvanica) increased in cover approximately 6‐fold, accounting for 84–90% of the increased herbaceous cover. Other native ruderals and exotic herbs reached 2 and less than or equal to 1%, cover, respectively, after three years. Species richness across cleared treatments increased from 30 to 79 species. All forest species were retained. Forest shrubs and trees initially declined from their dominant cover but rebounded after three years. Tree clearing plus seeding appeared to be a viable management practice for increasing cover of herbaceous sandplain species while causing minimal increases in exotic herbaceous cover. The long‐term persistence of sandplain herbs may require periodic disturbances that limit woody regrowth.     

The resprouting response of co‐occurring temperate woody plant and grass species to elevated [CO2]: An insight into woody plant encroachment of grasslands

The phenomenon of woody plant thickening in grasslands has been observed globally and is likely to have widespread ecological consequences. It has been proposed that woody plant thickening is driven in part by rising atmospheric [CO₂] enhancing the resprouting ability of woody plants relative to grasses so they respond more strongly after disturbances such as herbivory and fire. The aim of this study was to examine the CO₂ effect on the resprouting ability of 16 co‐occurring temperate woody plant and grass species (eight species from each growth form). Plants were grown in a controlled glasshouse experiment under ambient (400 ppm) and elevated [CO₂] (600 ppm) for 14 weeks after which their resprouting ability was measured. Root non‐structural carbohydrate (NSC_mass) and nitrogen (N_mass) storage was used as proxies to measure the resprouting ability of woody plants while for the grasses it was measured directly. We found that both the woody plants (22% on average; P = 0.003) and grasses (20% on average; P = 0.003) produced more biomass under elevated [CO₂]. Despite the woody plants not allocating additional carbon to belowground storage under elevated [CO₂], they had significantly greater root NSC_mass (23% on average; P = 0.007) due to increased root biomass production (8% on average; P = 0.007). In contrast, root N_mass of the woody plants did not differ between CO₂ treatments (P = 0.373). Surprisingly, the resprouting ability of the grasses did not significantly differ between the CO₂ treatments (P = 0.067). These results provide evidence that the differing resprouting response of woody plants and grasses under elevated [CO₂] may be contributing to woody plant encroachment of grasslands worldwide.     

Response of plant species and life form diversity to variable fire histories and biomass in the jarrah forest of south‐west Australia

Frequent fires reduce the abundance of woody plant species and favour herbaceous species. Plant species richness also tends to increase with decreasing vegetation biomass and cover due to reduced competition for light. We assessed the influence of variable fire histories and site biomass on the following diversity measures: woody and herbaceous species richness, overall species richness and evenness, and life form evenness (i.e. the relative abundance or dominance among six herbaceous and six woody plant life forms), across 16 mixed jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) forest stands in south‐west Australia. Fire frequency was defined as the total number of fires over a 30‐year period. Overall species richness and species evenness did not vary with fire frequency or biomass. However, there were more herbaceous species (particularly rushes, geophytes and herbs) where there were fewer shrubs and low biomass, suggesting that more herbaceous species coexist where dominance by shrubs is low. Frequently burnt plots also had lower number and abundance of shrub species. Life form evenness was also higher at both high fire frequency and low biomass sites. These results suggest that the impact of fire frequency and biomass on vegetation composition is mediated by local interactions among different life forms rather than among individual species. Our results demonstrate that measuring the variation in the relative diversity of different woody and herbaceous life forms is crucial to understanding the compositional response of forests and other structurally complex vegetation communities to changes in disturbance regime such as increased fire frequency.     

Survival,growth, and photosynthesis of tree seedlings competing with herbaceous vegetation along a water-light-nitrogen gradient

In herbaceous dominated patches and ecosystems, tree establishment is influenced partly by the ability of woody seedlings to survive and grow in direct competition with herbaceous vegetation. We studied the importance of season long wet and dry spells on the competitive interactions between herbaceous vegetation and oak seedlings along a light and nitrogen gradient in an infertile secondary successional grassland in central North America. We conducted a field experiment in which seedlings of bur oak (Quercus macrocarpa) and northern pin oak (Q. ellipsoidalis) were exposed to two levels of light (full sun and 80% shade), three levels of nitrogen input (0, 5, 15 g m^–1 yr^–1), and three levels of water input (low, medium and high). In addition, seedlings were grown with and without the presence of surrounding herbaceous vegetation under both light and all three water levels. Seedling survival, growth, and rate of photosynthesis were significantly affected by competition with herbaceous vegetation and these effects varied along the multiple resource gradient. Overall, seedling survival of both species was significantly greater in wetter and shaded plots and when surrounding herbaceous vegetation was removed and was lower in nitrogen enriched plots. We found that soil water was significantly affected by varying inputs of water, light, and the presence or absence of herbaceous vegetation, and that seedling survival and rate of photosynthesis were highly correlated with available soil water. Our findings show that the impact of season long wet and dry spells on tree seedling success in grasslands can be affected by light and soil nitrogen availability.     

Stochastic losses of fire‐dependent endemic herbs revealed by a 65‐year chronosequence of dispersal‐limited woody plant encroachment

The factors responsible for maintaining diverse groundcover plant communities of high conservation value in frequently burned wet pine savannas are poorly understood. While most management involves manipulating extrinsic factors important in maintaining species diversity (e.g., fire regimes), most ecological theory (e.g., niche theory and neutral theory) examines how traits exhibited by the species promote species coexistence. Furthermore, although many ecologists focus on processes that maintain local species diversity, conservation biologists have argued that other indices (e.g., phylogenetic diversity) are better for evaluating assemblages in terms of their conservation value. I used a null model that employed beta‐diversity calculations based on Raup–Crick distances to test for deterministic herbaceous species losses associated with a 65‐year chronosequence of woody species encroachment within each of three localities. I quantified conservation value of assemblages by measuring taxonomic distinctness, endemism, and floristic quality of plots with and without woody encroachment. Reductions in herb species richness per plot attributable to woody encroachment were largely stochastic, as indicated by a lack of change in the mean or variance in beta‐diversity caused by woody encroachment in the savannas studied here. Taxonomic distinctness, endemism, and floristic quality (when summed across all species) were all greater in areas that had not experienced woody encroachment. However, when corrected for local species richness, only average endemism and floristic quality of assemblages inclusive of herbs and woody plants were greater in areas that had not experienced woody encroachment, due to the more restricted ranges and habitat requirements of herbs. Results suggest that frequent fires maintain diverse assemblages of fire‐dependent herb species endemic to the region. The stochastic loss of plant species, irrespective of their taxonomic distinctness, to woody encroachment suggests that the relevance of niche partitioning or phylogenetic diversity to the management of biodiversity in wet pine savannas is minimal.     

Soil C and N responses to woody plant expansion in a mesic grassland

Changes in land management and reductions in fire frequency have contributed to increased cover of woody species in grasslands worldwide. These shifts in plant community composition have the potential to alter ecosystem function, particularly through changes in soil processes and properties. In semi-arid grasslands, the invasion of shrubs and trees is often accompanied by increases in soil resources and more rapid N and C cycling. We assessed the effects of shrub encroachment in a mesic grassland in Kansas (USA) on soil CO₂ flux, extractable inorganic N, and N mineralization beneath shrub communities (Cornus drummondii) and surrounding undisturbed grassland sites. In this study, a shift in plant community composition from grassland to shrubland resulted in a 16% decrease in annual soil CO₂ flux(4.78 kg CO₂ m^–2 year^–1 for shrub dominated sites versus 5.84 kg CO₂ m^–2 year^–1 for grassland sites) with no differences in total soil C or N or inorganic N. There was considerable variability in N mineralization rates within sites, which resulted in no overall difference in cumulative N mineralized during this study (4.09 g N m^–2 for grassland sites and 3.03 g N m^–2 for shrub islands). These results indicate that shrub encroachment into mesic grasslands does not significantly alter N availability (at least initially), but does alter C cycling by decreasing soil CO₂ flux.     

Increased tree densities in South African savannas: >50 years of data suggests CO2 as a driver

For the past century, woody plants have increased in grasslands and savannas worldwide. Woody encroachment may significantly alter ecosystem functioning including fire regimes, herbivore carrying capacity, biodiversity and carbon storage capacity. Traditionally, increases in woody cover and density have been ascribed to changes in the disturbance regime (fire and herbivores) or rainfall. Increased atmospheric CO₂ concentrations may also contribute, by increasing growth rates of trees relative to grasses. This hypothesis is still heavily debated because usually potential CO₂ effects are confounded by changes in land use (disturbance regime). Here we analyse changes in woody density in fire experiments at three sites in South African savannas where the disturbance regime (fire and herbivores) was kept constant for 30 and 50 years. If global drivers had significant effects on woody plants, we would expect significant increases in tree densities and biomass over time under the constant disturbance regime. Woody density remained constant in a semiarid savanna but tripled in a mesic savanna between the 1970s and 1990s. At the third site, a semiarid savanna near the southern limits of the biome, tree density doubled from the mid 1990s to 2010. Interpretation of the causes is confounded by population recovery after clearing, but aerial photograph analysis on adjacent non‐cleared areas showed an accompanying 48% increase in woody cover. Increased CO₂ concentrations are consistent with increased woody density while other global drivers (rainfall) remained constant over the duration of the experiments. The absence of a response in one semiarid savanna could be explained by a smaller carbon sink capacity of the dominant species, which would therefore benefit less from increased CO₂. Understanding how savannas and grasslands respond to increased CO₂ and identifying the causes of woody encroachment are essential for the successful management of these systems.     

Plant–community responses to shrub cover in a páramo grassland released from grazing and burning

Shrub encroachment can follow grazing or burning release in páramo grasslands. While encroachment decreases herbaceous species richness in some grassland systems, the effects of this process on the herbaceous community in páramo grasslands are currently unknown. We collected data on shrub cover, herbaceous‐species cover and species composition in a páramo grassland 12 years after release from burning and cattle grazing near Zuleta, Ecuador. Topographic and soil measures were also included as predictor variables of differences in community composition. Contrary to studies in other systems, shrub cover did not have a significant effect on herbaceous‐species richness, whereas shrub‐species richness significantly increased with shrub cover. However, shrub cover was associated with significant shifts in herbaceous–community composition. Most notably, there was an increase in some shade‐tolerant forbs and tall‐statured wetland grasses with increasing shrub cover, and a corresponding decrease in some short‐statured grasses and early successional forbs. These results could indicate that the ameliorative effects of shrubs (e.g. frost and wind protection) in harsh alpine environments may partially compensate for the expected competitive effect of shrubs due to shading.     

Effects of contemporary environment and Quaternary climate change on drylands plant diversity differ between growth forms

Previous studies on large‐scale patterns in plant richness and underlying mechanisms have mostly focused on forests and mountains, while drylands covering most of the world's grasslands and deserts are more poorly investigated for lack of data. Here, we aim to 1) evaluate the plant richness patterns in Inner Asian drylands; 2) compare the relative importance of contemporary environment, historical climate, vegetation changes, and mid‐domain effect (MDE); and 3) explore whether the dominant drivers of species richness differ across growth forms (woody vs herbaceous) and range sizes (common vs rare). Distribution data and growth forms of 13 248 seed plants were compiled from literature and species range sizes were estimated. Generalized linear models and hierarchical partitioning were used to evaluate the relative contribution of different factors. We found that habitat heterogeneity strongly affected both woody and herbaceous species. Precipitation, climate change since the mid‐Holocene and climate seasonality dominated herbaceous richness patterns, while climate change since the Last Glacial Maximum dominated woody richness patterns. Rare species richness was strongly correlated with precipitation, habitat heterogeneity and historical climatic changes, while common species richness was strongly correlated with MDE (woody) or climate seasonality (herbaceous). Temperature had little effects on the species richness patterns of all groups. This study represents the first evaluation of the large‐scale patterns of plant species richness in the Inner Asian drylands. Our results suggest that increasing water deficit due to anthropogenic activities combined with future global warming may increase the extinction risk of many grassland species. Rare species (both herbaceous and woody) may face severe challenges in the future due to increased habitat destruction caused by urbanization and resource exploitation. Overall, our findings indicate that the hypotheses on species richness patterns based on woody plants alone can be insufficient to explain the richness patterns of herbaceous species.