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.     

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.     

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.     

Effects of Prosopis flexuosa on soil properties and the spatial pattern of understorey species in arid Argentina

Abstract. In arid zones dominant woody plants are capable of causing changes in microclimate and soil properties likely to affect species composition, as well as the establishment and spatial distribution of plant species. In North American and European deserts species richness appears to be higher under the canopy of shrubs and trees, in contrast with Chilean deserts where it seems to be lower. Since Prosopis flexuosa (Fabaceae, Mimosoideae) is the most conspicuous tree in the central Monte desert, Argentina, we analysed the effect of this species on the composition and abundance of the shrub and herbaceous layers and on soil properties. We considered two mesohabitats: ‘under P. flexuosa canopy’ and ‘intercanopy areas’. In addition, we analysed the differences between two microhabitats under canopies: ‘northern part of the canopy’ and ‘southern part of the canopy’. Results indicate that species composition and soil properties are affected by both mesohabitats and microhabitats. We found a higher number of shrubs under canopies, whereas that of grasses and perennial forbs increased in intercanopy areas. Concentrations of organic matter, nitrogen, potassium and phosphorus, factors limiting biological productivity in Monte desert soils, were significantly higher under than outside P. flexuosa canopies. Electrical conductivity and concentrations of Na⁺, Ca⁺⁺, Mg⁺⁺ were higher in the northern than in the southern microhabitats. No differences in species richness, evenness or diversity were found between mesohabitats or between microhabitats. We conclude that P. flexuosa modifies the spatial pattern of plant species in the shrub and herbaceous layers and the chemical conditions of the soil, generating spatial heterogeneity on different scales.     

Regional Estimate of Nitric Oxide Emissions Following Woody Encroachment: Linking Imaging Spectroscopy and Field Studies

Woody encroachment, a spatially explicit process of land-cover change, is known to affect the biophysical and biogeochemical properties of ecosystems. However, little information is available on the impacts of woody encroachment on N oxide emissions from savanna regions. We combined hyperspectral remote sensing and field measurements to quantify spatial patterns and estimate regional fluxes of soil N oxide emissions as they covary with vegetation cover and soil type across a semiarid rangeland in north Texas. Soil nitric oxide (NO) emissions were highly correlated with Prosopis canopy cover, allowing the extrapolation of NO fluxes from hyperspectral observations of woody cover. NO emissions were highly variable, ranging from 0 to 550 kg NO-N km^–2 y^–1 across the region, with the lowest emissions from shallow clay soils and highest from deeper upland clay loams. An estimate of annual NO emissions based on remotely derived Prosopis cover, temperature, and precipitation was 160 kg NO-N km^–2 y^–1, almost twice that of the value derived from traditional averaging of field measurements. We conclude that relationships between NO emissions and remotely sensed structure and composition are advantageous for quantifying NO emissions at the regional scale. This study also provides new insight into the role of woody encroachment on biogeochemical processes that are highly variable and otherwise difficult to measure at the regional scale.     

Positive interaction between invasive plants: The influence of Pyracantha angustifolia on the recruitment of native and exotic woody species

Abstract: Positive interactions between species are known to play an important role in the dynamics of plant communities, including the enhancement of invasions by exotics. We studied the influence of the invasive shrub Pyracantha angustifolia (Rosaceae) on the recruitment of native and exotic woody species in a secondary shrubland in central Argentina mountains. We recorded woody sapling recruitment and micro‐environmental conditions under the canopies of Pyracantha and the dominant native shrub Condalia montana (Rhamnaceae), and in the absence of shrub cover, considering these situations as three treatments. We found that native and exotic species richness were higher under Pyracantha than under the other treatments. Ligustrum lucidum (Oleaceae), an exotic bird‐dispersed shade‐tolerant tree, was the most abundant species recruiting in the area, and its density was four times higher under the canopy of Pyracantha. This positive interaction may be related to Pyracantha's denser shading, to the mechanical protection of its canopy against ungulates, and/or to the simultaneous fruit ripening of both woody invaders.     

Impacts of a mineral lick‐centred land use system on woody vegetation cover in an East African Savannah

Evaluation of woody vegetation changes with distance from a salt crater was conducted in the semi‐arid rangelands of southern Ethiopia. Data on live woody plants were collected over three seasons at 0, 1, 4, 6, 9 and 12 km from the salt crater. The density and diversity of woody plants differed significantly (p < .01) along the distance gradient. Six woody plant families were identified of which Fabaceae and Burseraceae were the dominant families. Acacia drepanolobium, Acacia nilotica, Commiphora africana and Acacia mellifera were among the severely encroaching woody species. There were high proportions of seedlings and saplings recorded closer to the salt crater showing a vigorous recruitment by woody plants. Woody plant encroachment along the 12‐km transect ranged from a low to severe encroachment, which could be translated into poor rangeland condition. Changes in soil characteristics increased grazing pressure and sedentary settlement around the salt crater, and the breakdown of traditional institutions seems to be major contributing factors to these vegetation changes. We suggest that severely encroached areas could be improved through a combination of methods such as bush clearing, prescribed fire, browsing animals and proper grazing management.     

Encroachment of woody plants and its impact on pastoral livestock production in the Borana lowlands, southern Oromia, Ethiopia

Encroachment of woody plants has been among the major threats to the livelihoods of Borana pastoralists and their ecosystem. An approach that integrated vegetation survey and pastoralists’ perception was followed to study the impacts of encroachment of woody plants in the Borana lowlands, Ethiopia. Density of woody species was determined in 192 plots of 500 m². Canopy cover of woody plants was estimated in 123 quadrates of 400 m². Pastoralists’ perception was assessed through group discussions and a semi‐structured questionnaire. Results showed that plant density was 3014 woody plants ha^?1. Cover of woody plants was 52%, indicating an increasing trend from ≤40% cover reported in the early 1990s. It was concluded that the increase of woody plants density and cover has crossed the critical threshold and has entered into the encroached condition. Principal components analysis (PCA) and redundancy analysis (RDA) also showed that woody plants were negatively correlated with herbaceous biomass. Commiphora africana, Acacia melliphera, A. drepanolobium, A. brevispica and Lannea rivae were among the dominant encroachers. RDA revealed that soil nutrients were positively correlated with woody plants density and cover. The pastoralists perceived that encroachment of woody plants had decreased the production of their grazingland. A ban on fire was perceived as the major factor that caused encroachment of woody plants. Re‐utilization of fire and strengthening of traditional rangeland management strategies are recommended.     

The Effect of Biological Soil Crusts on Throughput of Rainwater and N into Lake Michigan Sand Dune Soils

Biological soil crusts composed of cyanobacteria, green algae, bryophytes, and lichens colonize soils in arid and semiarid ecosystems worldwide and are responsible for significant N input to the soils of these ecosystems. Soil crusts also colonize active sand dunes in more humid regions, but studies of structure and function of such sand dune crusts are lacking. We identified the cyanobacterial, algal, and bryophytic constituents and N production and leachates of biological soil crusts that colonize beach dunes at the Indiana Dunes National Lakeshore along southern Lake Michigan in Indiana, USA. To determine the role of these crusts in this system, we conducted a greenhouse experiment in which intact soil cores with biological crusts were subjected to artificial rainfall over a full growing season. The volume and N content of leachate from the cores were quantified in relation to degree of crust development, taxonomic composition, rainfall volume and intensity, light intensity, and the presence of plant litter. Net N throughput significantly exceeded N inputs to cores in rainwater. Net N outputs from crusts to subsurface soil ranged from 0. 01 to 0.19 g NH ₄ ⁺ -N m⁻² yr⁻¹ and 0.01 to 0.61 g NO ₃ ^– N m⁻² yr⁻¹. Thus, total inorganic N inputs associated with biological soil crusts ranged from 0.02 g N m⁻² yr⁻¹ to 0.8 g N m⁻² yr⁻¹. High volume (≥2 cm) rainfall resulted in more N leaching than low volume events, and plant litter added over the surface of crusted soil cores significantly increased the amount of N in leachate. Exploratory path analysis revealed direct and indirect linkages among environmental factors, crust development, and crust composition in regulating the throughput of H₂O and N from these intact soil cores. Biological soil crusts at this site, combined with other properties of the soil surface, substantially increase N inputs to this water- and nutrient-limited sand dune ecosystem.     

Herbivore species and grazing intensity regulate community composition and an encroaching woody plant in semi-arid rangeland

Grazing by livestock can influence ecosystems in various ways, including altering plant communities, influencing woody plant encroachment, and determining livestock productivity. Evaluating long term effects of grazing on plant composition is valuable not only to understand herbivory on rangelands but to be able to address the primary factors that can threaten long term livestock productivity. We examined plant species composition and woody plant encroachment 45 years after the initiation of differing grazing treatments within a semiarid savanna of the southern Great Plains, USA. Grazing treatments varied in herbivore type (domestic cattle, sheep, and goats vs. goats only) and grazing intensity (heavy, moderate, and no-herbivory). All individual trees of Juniperus ashei Buchholz, the encroaching woody plant of the area, were removed prior to treatment initiation. Moderate and heavy grazing by a combination of species resulted in similar plant communities, while a history of heavy browsing by goats only and no-herbivory resulted in more distinct communities. Cover of J. ashei did not differ between mixed grazing and no-herbivory treatments, indicating that grazing was not responsible for woody plant encroachment. J. ashei cover within the browsed treatment was a third less compared to other treatments; compositional differences within this treatment are possibly due to reduced cover of woody vegetation. Declines in livestock productivity of the area are likely related to compositional changes resulting from increased woody plants. Livestock production within this semi-arid rangeland is likely unsustainable without management of woody plant encroachment, as communities tend to a closed canopy woodland.     

Interactive Effects of Three Ecosystem Engineers on Infiltration in a Semi-Arid Mediterranean Grassland

The redistribution of water in semi-arid environments is critical for the maintenance and survival of vegetation patches. We used a systems approach to examine the interactive effects of three engineers—Stipa tenacissima, biological soil crusts, and the European rabbit (Oryctolagus cuniculus)—on infiltration processes in a model gypseous semi-arid Mediterranean grassland. We measured the early (sorptivity) and later (steady-state infiltration) stages of infiltration at two supply potentials using disk permeameters, which allowed us to determine the relative effects of different engineers and soil micropores on water flow through large macropores. We detected few effects under tension when flow was restricted to matrix pores, but under ponding, sorptivity and steady-state infiltration adjacent to Stipa tussocks were 2–3 times higher than in intact or rabbit-disturbed biological soil crusts. Structural Equation Modeling (SEM) showed that both Stipa and biological soil crust cover exerted substantial and equal positive effects on infiltration under ponding, whereas indirectly, rabbit disturbance negatively affected infiltration by reducing crust cover. Under tension, when macropores were prevented from conducting water, Stipa had a direct negative effect and biological soil crust cover was relatively unimportant. More complex SEM models demonstrated that (1) Stipa primarily influenced biological soil crusts by reducing their richness, (2) rabbits exerted a small negative effect on crust richness, and (3) lichens were negatively, and mosses positively, correlated with a derived “infiltration” axis. Our results highlight the importance of biological soil crusts as key players in the maintenance of infiltration processes in Stipa grasslands, and demonstrate the modulating role played by rabbits through their surface disturbances.     

Shrub emergence and seedling growth in microenvironments created by Prosopis glandulosa

Abstract. Facilitation of the establishment of certain plant species by nurse plants is a common phenomenon in arid and semiarid ecosystems. The most commonly reported mechanisms of facilitation include cooler temperatures and increased soil nutrients beneath the nurse plant canopy, which favor establishment of other plant species. During conversion of upland grasslands to thorn woodland in southern Texas, Prosopis glandulosa appears to facilitate establishment of other woody plants, including Celtis pallida, whereas Acacia smallii occurs only in habitats between P. glandulosa canopies. We tested the hypothesis that light intensity and soils under P. glandulosa canopies facilitate seedling emergence and growth of C. pallida but inhibit seedling emergence and growth of A. smallii. In the field, C. pallida and A. smallii seeds were planted under P. glandulosa canopies and in adjacent interspaces. Percent emergence of C. pallida seedlings was greater under the canopy of P. glandulosa, whereas percent emergence of A. smallii seedlings was greater in interspaces. In a greenhouse experiment, seeds of each species were planted in pots filled with soil from under P. glandulosa canopies or from adjacent interspaces. Two treatments, shade and sunlight, were imposed and plants harvested seven weeks later. Seedling mass of both species was greater in canopy soil than in interspace soil in sunlight but mass of the two species did not differ between soil sources in shade. Canopy soils contained more total and available nitrogen than interspace soils. These results suggest that light is more limiting than nutrients under shaded conditions and so neither species can take advantage of the high nutrients beneath P. glandulosa. Shade and greater soil nutrients beneath P. glandulosa do not appear to be the major factors that facilitate C. pallida or inhibit A. smallii. Aggregation of C. pallida beneath P. glandulosa canopies appears to be a complex process that involves both passive facilitation (seed dispersal by birds) and active facilitation (reduction of seed dormancy by under-canopy temperatures) operating only during the seed germination stage with successional mechanisms other than facilitation operating during later stages of shrub establishment and growth.     

The influence of Acacia tortilis (Forssk.) Subsp. raddiana (Savi) and livestock grazing on grass species composition, yield and soil nutrients in arid environments of South Tunisia

The objectives of the study, conducted during the 2003/2004 growing season in the National Park of Bou Hedma (South Tunisia), were to quantify the effects of the single-woody species Acacia tortilis subsp. raddiana on grass species composition, on total plant cover, on density of perennial species, on dry matter (DM) yield and on soil nutrients at lightly and heavily grazed sites. In each study site, two subhabitats were distinguished, i.e. under tree canopies and open grasslands. In the lightly grazed site, the nutrient status of soil (organic matter, total N, extractable P, K⁺, Ca²⁺, Na⁺, Mg²⁺) under Acacia raddiana canopy, was found to be significantly higher (p<0.05) than under the open grassland. In the same way, total plant cover (p<0.05), density of perennial species (p<0.01) and DM yield (p<0.01) were significantly higher under tree canopies in the lightly grazed site. Heavy grazing proved to exert a strong overriding effect over the positive influences of the woody plants. For most studied parameters, a non-significant difference was recorded between canopied and uncanopied subhabitats. Some palatable species were frequently found under trees. In the heavily grazed site, these species are being replaced by less desirable species. This emphasizes the importance of conservation stocking rates and proper pasture management.