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.     

Seed dispersal by cattle may cause shrub encroachment of Grewia flava on southern Kalahari rangelands

Abstract. Shrub encroachment, i.e. the increase in woody plant cover, is a major concern for livestock farming in southern Kalahari savannas. We developed a grid‐based computer model simulating the population dynamics of Grewia flava, a common, fleshy‐fruited encroaching shrub. In the absence of large herbivores, seeds of Grewia are largely deposited in the sub‐canopy of Acacia erioloba. Cattle negate this dispersal limitation by browsing on the foliage of Grewia and dispersing seeds into the grassland matrix. In this study we first show that model predictions of Grewia cover dynamics are realistic by comparing model output with shrub cover estimates obtained from a time series of aerial photographs. Subsequently, we apply a realistic range of intensity of cattle‐induced seed dispersal combined with potential precipitation and fire scenarios. Based on the simulation results we suggest that cattle may facilitate shrub encroachment of Grewia. The results show that the severity of shrub encroachment is governed by the intensity of seed dispersal. For a high seed dispersal intensity without fire (equivalent to a high stocking rate) the model predicts 56% shrub cover and 85% cell cover after 100 yr. With fire both recruitment and shrub cover are reduced, which may, under moderate intensities, prevent shrub encroachment. Climate change scenarios with two‐fold higher frequencies of drought and wet years intensified shrub encroachment rates, although long‐term mean of precipitation remained constant. As a management recommendation we suggest that shrub encroachment on rangelands may be counteracted by frequent fires and controlling cattle movements to areas with a high proportion of fruiting Grewia shrubs.     

Shrub canopies influence soil temperatures but not nutrient dynamics: An experimental test of tundra snow–shrub interactions

Shrubs are the largest plant life form in tundra ecosystems; therefore, any changes in the abundance of shrubs will feedback to influence biodiversity, ecosystem function, and climate. The snow–shrub hypothesis asserts that shrub canopies trap snow and insulate soils in winter, increasing the rates of nutrient cycling to create a positive feedback to shrub expansion. However, previous work has not been able to separate the abiotic from the biotic influences of shrub canopies. We conducted a 3‐year factorial experiment to determine the influences of canopies on soil temperatures and nutrient cycling parameters by removing ~0.5 m high willow (Salix spp.) and birch (Betula glandulosa) shrubs, creating artificial shrub canopies and comparing these manipulations to nearby open tundra and shrub patches. Soil temperatures were 4–5°C warmer in January, and 2°C cooler in July under shrub cover. Natural shrub plots had 14–33 cm more snow in January than adjacent open tundra plots. Snow cover and soil temperatures were similar in the manipulated plots when compared with the respective unmanipulated treatments, indicating that shrub canopy cover was a dominant factor influencing the soil thermal regime. Conversely, we found no strong evidence of increased soil decomposition, CO₂ fluxes, or nitrate or ammonia adsorbtion under artificial shrub canopy treatments when compared with unmanipulated open tundra. Our results suggest that the abiotic influences of shrub canopy cover alone on nutrient dynamics are weaker than previously asserted.     

Changes in forest understory associated with Juniperus encroachment in Oklahoma,USA

Question: Does understory vegetation cover and richness decline along a gradient of increasing Juniperus virginiana midstory canopy cover and is that decline best correlated with litter accumulation? Location: Cross Timbers Forest in Payne County, OK, USA. Methods: We measured vegetation in forest gaps as well as forest areas without J. virginiana, at the inner and outer edge of J. virginiana canopies and near J. virginiana trunks (200 plots) and compared vegetation differences among location to light, litter, soil and microclimate variables. Results: Species richness (11 spp m^?2 to 6 spp m^?2) and summer vegetation cover (53.3% to 12.7%) declined with proximity to trunks. Regression indicated that richness declines (R²=0.08) and cover (R²=0.18) were best correlated with J. virginiana litter accumulation. Partial canonical correspondence analysis (pCCA) revealed two strong canonical axes, one related to litter/light and another to cover of Quercus spp. versus J. virginiana. Tree seedlings and woody vines dominated near J. virginiana. Forbs, graminoids and Quercus spp. seedlings were more common in areas without J. virginiana. Conclusions: Increasing J. virginiana and consequent litter additions alter understory biomass and composition and, through inhibiting Quercus spp. recruitment, may lead to changes in overstory composition. Decreases in herbaceous litter, which historically contributed to fuel accumulation, may have positive feedback effects on midstory encroachment by reducing the potential for prescribed burning.     

Variation in soil respiration under the tree canopy in a temperate mixed forest,central China,under different soil water conditions

The forest canopy cover can directly and indirectly affect soil conditions and hence soil carbon emission through soil respiration. Little is known, however, on the effects of canopy cover on soil respiration under the canopy of different tree species and soil water conditions. We have examined the variation in soil respiration at different soil water conditions (dry <10 %, wet >20 %, v/v) under different tree canopy covers in comparison with the canopy interspace in a temperate coniferous (Pinus armandii Franch) and broadleaved (Quercus aliena var. acuteserrata) mixed forest in central China. The results show that soil respiration measured under tree canopy cover varied with canopy size and soil water content. Soil respiration under small-sized canopies of P. armandii (PS) was higher than that under large-sized (PL) canopies, but the difference was only significant under the dry soil condition. However, soil respiration under large-sized canopies of Q. aliena (QL) was significantly greater than that under small-sized (QS) canopies under both dry and wet soil conditions. The difference in soil respiration between differently sized canopies of Q. aliena (33.5–35.8 %) was significantly greater than that between differently sized canopies of P. armandii (2.4–8.1 %). Differences in soil respiration between inter-plant gaps and under QS canopies in both the dry and wet soil conditions were significant. Significant increases in soil respiration (9.7–32.2 %) during the transition from dry to wet conditions were found regardless of canopy size, but the increase of soil respiration was significantly lower under P. armandii canopies (9.7–17.7 %) than under Q. aliena canopies (25.9–31.5 %). Our findings that the canopy cover of different tree species influences soil respiration under different soil moisture conditions could provide useful information for parameterizing and/or calibrating carbon flux models, especially for spatially explicit carbon models.     

Structure of biological soil crust communities in Callitris glaucophylla woodlands of New South Wales,Australia

Question: What is the nature of the relationships between cover, diversity and abundance of biological soil crusts, cover and diversity of vascular plants, and annual rainfall, soil texture and forestry practices in Callitris glaucophylla woodlands? Location: Arid and semi‐arid Callitris glaucophylla‐domi‐nated woodlands of eastern Australia. Methods: We documented soil crust‐forming mosses, lichens and liverworts at 83 woodland sites along a gradient of declining rainfall. Linear and non‐linear regression were used to examine relationships between soil crust species and attributes of vascular plant communities, and a similarity matrix (species abundance X sites) was subjected to Non‐metric Multi‐Dimensional Scaling (MDS), and Analysis of Similarities (ANOSIM) to show the degree of association between groups of taxa, and soil texture, rainfall classes and forestry practices. Results : We collected 86 taxa. Mosses were dominated by the family Pottiaceae, and lichens were dominated by squamulose forms. Average annual rainfall was highly correlated with soil crust community composition, and loamy soils supported a greater cover and diversity of taxa compared with sandy soils. Increases in tree cover were associated with significant, though weak, increases in abundance, but not diversity, of crusts. Crusts tended to be more diverse in areas that (1) had a sparse cover of ground‐storey plants; (2) were relatively stable ‐ as indicated by the proportion of perennial and/or native plants; (3) had more stable soil surfaces; and (4) were unlogged. Litter cover, overstorey thinning, and livestock grazing had no appreciable effect on crust diversity or cover. Conclusions : Callitris glaucophylla woodlands provide substantial habitat for soil crust organisms, and the dense tree cover and closed canopies of Callitris do not appear to have a major influence on the structure of biological crust communities. Unlike other woodland systems, relatively few patches would be required to reserve a high diversity of crust species.     

Spatial relations between shrubs and Prosopis glandulosa canopies

Abstract. Prosopis glandulosa, an arborescent legume, may act as a nurse plant that facilitates the establishment of other woody species. We hypothesized that attenuation of radiant energy and increased soil nutrients beneath P. glandulosa canopies facilitate establishment of subordinate shrubs and shrub cluster development. We determined the spatial distribution pattern of shrubs under P. glandulosa at three locations in southern Texas. Density of Celtis pallida, Zanthoxylum fagara, and total woody plants were comparable among the four cardinal directions at each location, which countered the prediction that shrub density would be greater on the north side of P. glandulosa canopies if attenuation of solar energy was a factor in cluster development. Total woody plant density increased with increasing P. glandulosa basal diameter, canopy radius, and height only at one location. Total woody plant density decreased with increasing total N in the upper 15 cm of soil at two of the three locations. Late in shrub cluster development, extraction of N from the soil and incorporation of N into plant tissue in dense shrub clusters may operate to inhibit further increases in subordinate shrub density.     

Woody plant encroachment reduces species richness of herb‐rich woodlands in southern Australia

Abstract Woody plants have been increasing in many woodland and savanna ecosystems owing to land use changes in recent decades. We examined the effects of encroachment by the indigenous shrub Leptospermum scoparium (Myrtaceae) on herb‐rich Eucalyptus camaldulensis woodlands in southern Australia. Species richness and compositional patterns were examined under the canopy of L. scoparium and in surrounding open areas to determine the species most susceptible to structural changes. Richness was significantly lower in areas of moderate to high L. scoparium cover (>15%), suggesting that a threshold shrub cover caused major change in this ecosystem. Shrubs were associated with a significant reduction in above‐ground biomass of the ground‐layer flora and a significant shift in community composition. The few species that were positively associated with high L. scoparium cover were also common in the woodland flora; no new species were recorded under the shrub canopy. Important environmental changes associated with L. scoparium cover were decreased light availability and increased litter cover, which were likely a consequence of encroachment. Leptospermum scoparium cover was also associated with greater surface soil moisture, which may be a consequence of increased shading under the shrub canopy or indicate favourable soil conditions for L. scoparium establishment. Reductions in species richness and abundance of the germinable seed bank were found in soil samples taken from under L. scoparium. With ongoing recruitment of L. scoparium and consequent increases in shrub cover, ground‐layer diversity in these species‐rich woodlands should continue to decline over time.     

Spatial and temporal changes in ecosystem carbon pools following juniper encroachment and removal

Proliferation of woody plants is a predominant global land cover change of the past century, particularly in dryland ecosystems. Woody encroachment and its potential impacts (e.g., livestock forage, wildlife habitat, hydrological cycling) have led to widespread brush management. Although woody plants may have substantial impacts on soils, uncertainty remains regarding woody encroachment and brush management influences on carbon (C) pools. Surface C pools (shallow soils and litter) may be particularly dynamic in response to encroachment and brush management. However, we have limited understanding of spatiotemporal patterns of surface C responses or how surface pools respond relative to aboveground C, litter, roots, and deep soil organic C. Spatial variability and lack of basic ecological data in woody-encroached dryland ecosystems present challenges to filling this data gap. We assessed the impact of western juniper (Juniperus occidentalis) encroachment and removal on C pools in a semi-arid sagebrush ecosystem. We used spatially-intensive sampling to create sub-canopy estimates of surface soil C (0–10 cm depth) and litter C pools that consider variation in tree size/age and sub-canopy location for live juniper and around stumps that were cut 7 years prior to sampling. We coupled the present size distribution of junipers with extensive existing allometric information about juniper in this region to estimate how landscape-level C pools would change through time under future management and land cover scenarios. Juniper encroachment and removal leads to substantial changes in C pools. Best-fit models for surface soil and litter C included positive responses to shrub basal diameter and negative responses to increasing relative distance from the bole to dripline. Juniper removal led to a net loss of surface C as a function of large decreases in litter C and small increases in surface soil C. At the landscape scale, deep soil C was the largest C pool (77 Mg C ha^?1), with an apparent lack of sensitivity to management. Overall, encroachment led to substantial increases in C storage over time as juniper size increased (excluding deep soil C, ecosystem C pools increased from 13.5 to 30.2 Mg C ha^?1 with transition from sagebrush-dominated to present encroachment levels). The largest pool of accumulation was juniper aboveground C, with important other pools including juniper roots, litter, and surface soil C. Woody encroachment and subsequent brush management can have substantive impacts on ecosystem C pools, although our data suggest the spatiotemporal patterns of surface C pools need to be properly accounted for to capture C pool responses. Our approach of coupling spatially-intensive surface C information with shrub distribution and allometric data is an effective method for characterizing ecosystem C pools, offering an opportunity for filling in knowledge gaps regarding woody encroachment and brush management impacts on local-to-regional ecosystem C pools.     

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.