Long‐term livestock exclusion facilitates native woody plant encroachment in a sandy semiarid rangeland

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A patch-dynamics approach to savanna dynamics and woody plant encroachment – Insights from an arid savanna

The coexistence of woody and grassy plants in savannas has often been attributed to a rooting-niche separation (two-layer hypothesis). Water was assumed to be the limiting resource for both growth forms and grasses were assumed to extract water from the upper soil layer and trees and bushes from the lower layers. Woody plant encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and is believed to be the result of overgrazing in ‘two-layer systems’. Recent research has questioned the universality of both the two-layer hypothesis and the hypothesis that overgrazing is the cause of woody plant encroachment.

We present an alternative hypothesis explaining both tree–grass coexistence and woody plant encroachment in arid savannas. We propose that woody plant encroachment is part of a cyclical succession between open savanna and woody dominance and is driven by two factors: rainfall that is highly variable in space and time, and inter-tree competition. In this case, savanna landscapes are composed of many patches (a few hectares in size) in different states of transition between grassy and woody dominance, i.e. we hypothesize that arid savannas are patch-dynamic systems. We summarize patterns of tree distribution observed in an arid savanna in Namibia and show that these patterns are in agreement with the patch-dynamic savanna hypothesis. We discuss the applicability of this hypothesis to fire-dominated savannas, in which rainfall variability is low and fire drives spatial heterogeneity.

We conclude that field studies are more likely to contribute to a general understanding of tree–grass coexistence and woody plant encroachment if they consider both primary (rain and nutrients) and secondary (fire and grazing) determinants of patch properties across different savannas.     

Fire-tolerance mechanisms of common woody plant species in a semiarid savanna in south-western Zimbabwe

The frequency of fire has increased in savannas yet few studies have assessed how plants persist when subjected to long‐term disturbance by fire. We investigated the contributions of bark thickness and resprouting to the persistence of woody plants in two fire trials that were started in 1948 and 1949. The number of resprouts per individual, bark thickness, basal diameter and height of woody plants were measured in unburnt plots and those burnt annually, triennially and quinquennially during the late dry season. Changes in tree density, number of resprouts and individuals in different height classes between 1963 and 2002 were assessed. Bark thickness varied among species and also increased with increases in basal diameter. Generally, plants with thick bark survived fire more than those with thin bark. Resprouting was the major fire survival strategy for most species. The number of resprouts produced per plant ranged from 4 ± 3 (Acacia rehmanniana) to 14 ± 9 (Pseudolachnostylis maprouneifolia). Fire reduced species richness in plots burnt annually and triennially by 47% and 6% respectively. Species richness increased in unburnt plots (5%) and those burnt quinquennially (16%). Most woody species survived fire through a combination of traits.     

Changes in aboveground primary production and carbon and nitrogen pools accompanying woody plant encroachment in a temperate savanna

When woody plant abundance increases in grasslands and savannas, a phenomenon widely observed worldwide, there is considerable uncertainty as to whether aboveground net primary productivity (ANPP) and ecosystem carbon (C) and nitrogen (N) pools increase, decrease, or remain the same. We estimated ANPP and C and N pools in aboveground vegetation and surface soils on shallow clay and clay loam soils undergoing encroachment by Prosopis glandulosa in the Southern Great Plains of the United States. Aboveground Prosopis C and N mass increased linearly, and ANPP increased logarithmically, with stand age on clay loam soils; on shallow clays, Prosopis C and N mass and ANPP all increased linearly with stand age. We found no evidence of an asymptote in trajectories of C and N accumulation or ANPP on either soil type even following 68 years of stand development. Production and accumulation rates were lower on shallow clay sites relative to clay loam sites, suggesting strong edaphic control of C and N accumulation associated with woody plant encroachment. Response of herbaceous C mass to Prosopis stand development also differed between soil types. Herbaceous C declined with increasing aboveground Prosopis C on clay loams, but increased with increasing Prosopis C on shallow clays. Total ANPP (Prosopis+herbaceous) of sites with the highest Prosopis basal area were 1.2 × and 4.0 × greater than those with the lowest Prosopis basal area on clay loam and shallow clay soils, respectively. Prosopis ANPP more than offset declines in herbaceous ANPP on clay loams and added to increased herbaceous ANPP on shallow clays. Although aboveground C and N pools increased substantially with Prosopis stand development, we found no corresponding change in surface soil C and N pools (0–10 cm). Overall, our findings indicate that Prosopis stand development significantly increases ecosystem C and N storage/cycling, and the magnitude of these impacts varied with stand age, soil type and functional plant traits     

Grass bud responses to fire in a semiarid savanna system

Increasingly, land managers have attempted to use extreme prescribed fire as a method to address woody plant encroachment in savanna ecosystems. The effect that these fires have on herbaceous vegetation is poorly understood. We experimentally examined immediate (<24 hr) bud response of two dominant graminoids, a C₃ caespitose grass, Nassella leucotricha, and a C₄ stoloniferous grass, Hilaria belangeri, following fires of varying energy (J/m²) in a semiarid savanna in the Edwards Plateau ecoregion of Texas. Treatments included high‐ and low‐energy fires determined by contrasting fuel loading and a no burn (control) treatment. Belowground axillary buds were counted and their activities classified to determine immediate effects of fire energy on bud activity, dormancy, and mortality. High‐energy burns resulted in immediate mortality of N. leucotricha and H. belangeri buds (p < .05). Active buds decreased following high‐energy and low‐energy burns for both species (p < .05). In contrast, bud activity, dormancy, and mortality remained constant in the control. In the high‐energy treatment, 100% (n = 24) of N. leucotricha individuals resprouted while only 25% (n = 24) of H. belangeri individuals resprouted (p < .0001) 3 weeks following treatment application. Bud depths differed between species and may account for this divergence, with average bud depths for N. leucotricha 1.3 cm deeper than H. belangeri (p < .0001). Synthesis and applications: Our results suggest that fire energy directly affects bud activity and mortality through soil heating for these two species. It is imperative to understand how fire energy impacts the bud banks of grasses to better predict grass response to increased use of extreme prescribed fire in land management.     

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.     

Responses of microbial components of the rhizosphere to plant management strategies in a semiarid rangeland

Summary Easily soluble heavy-metal fractions from different soils, a garbage-sewage sludge compost and peat were extracted by EUF. Blanks were determined by extracting distilled water. As the rubber seal of the extraction chamber contained Zn, the obtained Zn values were not reliable. The relative standard deviations of extracted micronutrients were 29.1% for Fe and 20.5% for Mn, Fe, Mn, Zn, Cu, Pb, Cd and Cr were not only found in the filters but also in the extracts.The extraction of CrIII and CrVI solutions showed that CrVI mainly migrated into the anode extract. CrIII was found mainly in the cathode filter and cathode extract, a smaller part however was obviously oxidized to CrVI and migrated into the anode extract. Consequently, CrIII and CrVI in soils could not be distinguished unequivocally by EUF.The amounts of Zn, Cu, Pb and Cd extracted by EUF from various substrates were small compared with the quantities extracted by 2N HCl. The heavy metal contents of the leaves were mostly in the order of those of the EUF extract.Several vineyard soils as well as peat were mixed with increasing quantities of Grünsalz (green salt), a fertilizer consisting mainly of iron sulphate. High amounts of Grünsalz (100–200 g/200 g soil) were necessary to raise soluble Fe in calcareous soils. In peat, however, small Grünsalz additions (1 g/50 g peat) were sufficient. Soluble Mn and Cu increased too when Grünsalz was added to soil or peat. These results give valuable information on how grapevine chlorosis can be reduced by the use of Grünsalz or mixtures of peat and Grünsalz.     

Temporal changes in the dominance of tree functional traits,but no changes in species diversity during woody plant encroachment in a Brazilian savanna

Questions : Woody encroachment in savannas has been associated with changing taxonomic composition and ecosystem function. Interestingly, there is little understanding of how encroachment impacts plant functional diversity and how those changes relate to plant demography, a crucial mediator between taxonomic composition and ecosystem function. Location : Southeastern Brazil. Methods: Using a landscape scale fire suppression experiment in a diverse Brazilian savanna, we quantify how change in species composition over seven years impacted vegetative and reproductive tree functional diversity as determined by new recruits, dead and surviving trees. Results: Over seven years, tree above-ground biomass increased by 15%, while total species richness did not change. Despite minor changes, species composition remained overall similar (82%), with few species contributing significantly to plot dissimilarity over time. There were small changes in vegetative traits, where the community-weighted mean increased in maximum tree height (↑ 2.1%) and specific leaf area (↑ 5.3%), and decreased in wood density (↓ 1.3%) and bark thickness (↓ 9.4%). Changes in reproductive traits were larger than in vegetative traits, with an increase in the prevalence of monoecy (↑ 32.6%), dioecy (↑ 44.2%), large seeds (↑ 20.3%), animal-mediated seed dispersal (↑ 4.9%) and pollination by very small insects (↑ 45.5%), and a decrease in the prevalence of hermaphroditism (↓ 9%), small seeds (6.8%) and pollination by small insects (12.5%). The overall decrease in bark thickness and increase in monoecy and dioecy were mainly driven by characters of the new recruits, while the overall increase in specific leaf area (SLA) and decrease in small seeds appeared largely determined by the loss of trees possessing those traits. Conclusions: Encroachment leads to changes that are likely increasing ecosystem vulnerability to fire and drought. Further, the compositional changes observed appear to drive marked change in reproductive traits, indicating increasing dependence on animals for dispersal and reproduction. Understanding post-hoc encroachment impacts in an era of widespread pervasive encroachment is fundamental to reconciling ecosystem functions such as nutrient cycling and pollination services as there is a loss of species with open ecosystem life-history strategies. Among savannas, there remains an urgent need to understand relationships between woody cover and ecosystem function to determine thresholds in woody cover promoting resilient savanna ecosystems.     

Changes to soil organic N dynamics with leguminous woody plant encroachment into grasslands

Encroachment of nitrogen-fixing trees and shrubs into grasslands and savannas is a well-documented land cover change that occurs worldwide. In the Rio Grande Plains region of southern Texas, previous studies have shown woody encroachment by leguminous Prosopis glandulosa (mesquite) trees increases soil C and N, decreases microbial biomass N relative to soil N, and accelerates N mineralization and nitrification. We examined responses of the dominant organic N components in soil (amino acids and amino sugars) and two soil-bound protein-N acquiring enzymes (arylamidase and β-N-acetylglucosaminidase) along a grassland-to-woodland successional chronosequence to determine changes to soil N chemistry and extractability. The proportion of total N held within amino compounds was significantly lower in the woodlands (47 %) relative to the grassland soils (62 %). This increase in non-hydrolysable N was accompanied by increases in plant cell wall derived amino acids (e.g. hydroxyproline, serine) and losses of microbial amino sugars, indicating the woodland organic N pool was altered in composition and potentially in quality, either because it was more structurally protected or difficult to degrade due to polymerization/condensation reactions. Soil carbon-normalized activities of both soil-bound N-acquiring enzymes were significantly higher in woodland soils, consistent with changes in the biochemical composition of organic N. Although soil total N increases following woody encroachment, this additional organic N appears to be less extractable by chemical hydrolysis and thus potentially in more refractory forms, which may limit microbial N accessibility, slow the cycling of soil organic carbon, and contribute to observed soil C and N accrual in these systems.     

The sensitivity of ecosystem carbon exchange to seasonal precipitation and woody plant encroachment

Ongoing, widespread increases in woody plant abundance in historical grasslands and savannas (woody encroachment) likely will interact with future precipitation variability to influence seasonal patterns of carbon cycling in water-limited regions. To characterize the effects of woody encroachment on the sensitivity of ecosystem carbon exchange to seasonal rainfall in a semi-arid riparian setting we used flux-duration analysis to compare 2003-growing season NEE data from a riparian grassland and shrubland. Though less seasonally variable than the grassland, shrubland NEE was more responsive to monsoon rains than anticipated. During the 2004-growing season we measured leaf gas exchange and collected leaf tissue for δ¹³C and nitrogen content analysis periodically among three size classes of the dominant woody-plant, Prosopis velutina and the dominant understory species, Sporobolus wrightii, a C₄ bunchgrass, present at the shrubland. We observed size-class and plant functional type independent patterns of seasonal plant performance consistent with greater-than-anticipated sensitivity of NEE in the shrubland. This research highlights the complex interaction between growing-season precipitation, plant-available alluvial groundwater and woody plant abundance governing ecosystem carbon balance in this semi-arid watershed.     

Leaf chemistry of woody plants in relation to season,canopy retention and goat browsing in a semiarid subtropical savanna

Abstract It is assumed that the phytochemistry of browse species protects their biomass and nutrients against herbivory. In this study we were primarily interested in (i) seasonal and phenology‐related variations in leaf chemistry, and (ii) chemistry‐related variations in the feeding behaviour of domestic goats. Such knowledge would guide management‐orientated modelling of browse–browser interactions in seasonal, subtropical zones where goats are abundant. The browse species studied were typical of semiarid savannas in southern Africa: Grewia occidentalis L. (Tiliaceae), Scutia myrtina (Burm. f) Kurz (Rhamnaceae), Diospyros lycioides Desf. ssp. lycioides (Ebenaceae), Rhus longispina Eckl. and Zeyh. (Anacardiaceae), Ehretia rigida (Thunb.) Druce (Boraginaceae) and Acacia karroo Hayne (Mimosoideae). Nitrogen (N), phosphorus (P), condensed tannins (CT), protein‐precipitating tannins (PPT), total phenols (TP), cellulose and lignin concentrations were estimated for each species during the late dormant and early growing seasons. N, P, CT and TP were elevated during the growth season, while cellulose, lignin and PPT decreased. Unlike cytoplasm contents, which varied seasonally, cell wall and vacuole contents varied both seasonally and among species. Except that seasonal variation in N of deciduous species was greater than that of evergreen species, leaf phenology was not related to variations in forage quality. Short‐term intake rates were not related to primary metabolite concentrations, but were positively related to secondary metabolites. Elevated intake rates of putative defences were concluded to be side‐effects of attempts by goats to increase nutrient intake rate, indicating tolerance of chemical defences. Without support for the hypothesis that chemical defences are correlated to canopy retention, we propose an alternative hypothesis, that defences are distributed among woody plants in semiarid, subtropical savannas according to shoot morphology because it affects the vulnerability of plant parts to browsers.     

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.     

Linking woody species diversity with plant available water at a landscape scale in a Brazilian savanna

Question: How is the diversity of woody species in a seasonally dry savanna related to plant available water (PAW)? Location: Savannas in central Brazil. Methods: Two‐dimensional soil resistivity profiles to 10‐m depth previously measured along three 10 m × 275 m replicate transects revealed differences in belowground water resources among and within transects: (1) driest/most heterogeneous; (2) wettest/least heterogeneous; and (3) PAW‐intermediate. All woody plants along these transects were identified to species, and height and basal circumference measured. Species diversity was evaluated for the whole transect (total diversity), 100‐m² plots (alpha‐diversity) and dissimilarity among 100‐m² plots within transects (beta‐diversity). Correlation analyses were conducted between PAW and vegetation variables at the 100‐m² scale. Results: The driest/most heterogeneous transect had the lowest total species diversity, while the wettest/least heterogeneous transect showed the lowest beta‐diversity. Floristic variation was correlated with PAW in all transects. In the most heterogeneous transect, species density was positively correlated with PAW in the 0‐400 cm soil layer. Evenness and Simpson's diversity were negatively correlated with PAW in the 700‐1000 cm soil layer. Conclusion: Woody species diversity was related to PAW at a fine spatial scale. Abundant PAW in the top 4 m of soil may favour many species and increase species total diversity. Conversely, abundant PAW at depth may result in lower evenness and total diversity, probably because the few species adapted to obtaining deep soil water can become dominant. Environmental changes altering soil water availability and partitioning in soil layers could affect the diversity of woody plants in this savanna.     

Explaining grass-nutrient patterns in a savanna rangeland of southern Africa

Aim The search for possible factors influencing the spatial variation of grass quality is an important step towards understanding the distribution of herbivores, as well as a step towards identifying crucial areas for conservation and restoration. A number of studies have shown that grass quality at a regional scale is influenced by climatic variables. At a local scale, site factors and their interaction are considered important. In this study, we aimed at examining environmental correlates of grass quality at a local scale. The study also sought to establish if biotic factors interact significantly with abiotic factors in influencing a variation in grass quality. Location The study area is located in the Kruger National Park of South Africa. The study area stretches from west (22°49′ S and 31°01′ E) to east, (22°44′ S and 31°22′ E) covering an area of about 25 × 6 km in the far northern region of the Kruger National Park. Methods We collected environmental data such as soil texture, percentage grass cover and biomass as well as grass samples for chemical analysis from specific locations in the study area. In addition, a digital elevation model (DEM) with a resolution of 5 m was used to derive elevation, slope and aspect using a geographic information system (GIS), which were related to grass quality. We used correlation analysis and anova to relate environmental variables to grass quality. Multivariate analysis techniques were used to simultaneously analyse and explore the complex interactions between variables. Results and conclusions Our results indicate that there is a significant relationship between grass quality parameters and site‐specific factors such as slope, altitude, percentage grass cover, aspect and soil texture. Relatively, percentage grass cover and soil texture were more critical in explaining a variation in grass quality. Plant characteristics such as species type interact significantly with slope, altitude and geology in influencing nutrient distribution. The results of this study may give a better insight on foliar nutrient distribution patterns at a landscape scale in savanna rangelands. Furthermore, the results of this study may help in the selection of ancillary information, which could be used in conjunction with other data such as remotely sensed data to map grass quality – an important step towards understanding the distribution and feeding patterns of wildlife. However, we acknowledge that this study is based on one seasonal snapshot, therefore some slightly different findings may be obtained during other times of the year. Nevertheless, the study has revealed that under the conditions experienced during the study period, nutrient distribution varies with varying biotic and abiotic factors.     

Ecosystem properties of semiarid savanna grassland in West Africa and its relationship with environmental variability

The Dahra field site in Senegal, West Africa, was established in 2002 to monitor ecosystem properties of semiarid savanna grassland and their responses to climatic and environmental change. This article describes the environment and the ecosystem properties of the site using a unique set of in situ data. The studied variables include hydroclimatic variables, species composition, albedo, normalized difference vegetation index (NDVI), hyperspectral characteristics (350–1800 nm), surface reflectance anisotropy, brightness temperature, fraction of absorbed photosynthetic active radiation (FAPAR), biomass, vegetation water content, and land‐atmosphere exchanges of carbon (NEE) and energy. The Dahra field site experiences a typical Sahelian climate and is covered by coexisting trees (~3% canopy cover) and grass species, characterizing large parts of the Sahel. This makes the site suitable for investigating relationships between ecosystem properties and hydroclimatic variables for semiarid savanna ecosystems of the region. There were strong interannual, seasonal and diurnal dynamics in NEE, with high values of ~?7.5 g C m^?2 day^?1 during the peak of the growing season. We found neither browning nor greening NDVI trends from 2002 to 2012. Interannual variation in species composition was strongly related to rainfall distribution. NDVI and FAPAR were strongly related to species composition, especially for years dominated by the species Zornia glochidiata. This influence was not observed in interannual variation in biomass and vegetation productivity, thus challenging dryland productivity models based on remote sensing. Surface reflectance anisotropy (350–1800 nm) at the peak of the growing season varied strongly depending on wavelength and viewing angle thereby having implications for the design of remotely sensed spectral vegetation indices covering different wavelength regions. The presented time series of in situ data have great potential for dryland dynamics studies, global climate change related research and evaluation and parameterization of remote sensing products and dynamic vegetation models.     

Neutral responses of plant community Ca concentration to nitrogen enrichment in a semiarid grassland

氮沉降对半干旱草原植物群落钙浓度的影响 钙(Ca)是植物生长所必需的营养物质,牧草中钙的含量对反刍动物的饲粮和健康具有重要意义。目前,关于氮沉降增加是否改变牧草的Ca浓度仍不是十分清楚。我们于2008–2015年在中国北方半干旱草原开展了氮沉降速率增加实验,测定了不同氮处理下每个样方内所有植物的Ca浓度,并且 根据各样方内每个物种的Ca浓度及其相对生物量,计算出功能群水平和群落水平的植物Ca浓度。研究结果表明,尽管物种水平和功能群水平的Ca浓度对氮沉降表现为负响应,但群落水平Ca浓度在整个氮添 加速率梯度上保持稳定。由于杂类草Ca浓度显著高于禾草,杂类草相对生物量的增加抵消了物种水平和功能群水平Ca浓度对氮沉降的负响应。另外,群落Ca库对氮添加速率表现为正饱和响应,且阈值为10 g N m−² yr−¹。本研究表明了氮沉降背景下植物相对生物量的变化对调控牧草Ca浓度和储量具有重要作用。