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.     

人工灌丛化对荒漠草原生态系统碳储量的影响

宁夏盐池县从20世纪70年代开始在荒漠草原上人工种植柠条灌木用以防风固沙和生态恢复,这一人为措施极大地改变了区域生态系统的植被结构和碳循环,而定量评估人工灌丛化对荒漠草原生态系统碳储量的影响,不仅能够揭示人类活动的碳循环反馈机制,而且可为地方政府生态治理提供理论指导。结合Biome-BGC模型和Logistics生长模型模拟了1958—2017年间荒漠草原人工灌丛化前后的碳储量变化,定量分析了人工灌丛化对生态系统碳储量和组分的影响。结果表明:(1)结合Biome-BGC模型和Logistics生长模型可以较准确地模拟出荒漠草原人工灌丛化过程中生态系统碳储量的变化。(2)人工灌丛化会快速改变荒漠草原的碳储量累积特征,柠条灌木种植后的快速生长阶段极大增强了生态系统的总碳储量,导致生态系统的碳储量特征由草地型向灌木型转变。(3)人工灌丛化改变了生态系统各类型碳储量的组分结构,其对地上植被和枯落物碳储量的影响非常明显,灌丛化后生态系统的植被和枯落物碳分别增加了6倍和1.76倍;因植被碳向土壤碳转化过程较慢,故人工灌丛化对地下土壤碳储量的影响在短期内较为微弱。以上结果显示,荒漠草原人工灌丛化能显...     

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     

Ecohydrological impacts of woody-plant encroachment: seasonal patterns of water and carbon dioxide exchange within a semiarid riparian environment

Across many dryland regions, historically grass‐dominated ecosystems have been encroached upon by woody‐plant species. In this paper, we compare ecosystem water and carbon dioxide (CO₂) fluxes over a grassland, a grassland–shrubland mosaic, and a fully developed woodland to evaluate potential consequences of woody‐plant encroachment on important ecosystem processes. All three sites were located in the riparian corridor of a river in the southwest US. As such, plants in these ecosystems may have access to moisture at the capillary fringe of the near‐surface water table. Using fluxes measured by eddy covariance in 2003 we found that ecosystem evapotranspiration (ET) and net ecosystem exchange of carbon dioxide (NEE) increased with increasing woody‐plant dominance. Growing season ET totals were 407, 450, and 639 mm in the grassland, shrubland, and woodland, respectively, and in excess of precipitation by 227, 265, and 473 mm. This excess was derived from groundwater, especially during the extremely dry premonsoon period when this was the only source of moisture available to plants. Access to groundwater by the deep‐rooted woody plants apparently decouples ecosystem ET from gross ecosystem production (GEP) with respect to precipitation. Compared with grasses, the woody plants were better able to use the stable groundwater source and had an increased net CO₂ gain during the dry periods. This enhanced plant activity resulted in substantial accumulation of leaf litter on the soil surface that, during rainy periods, may lead to high microbial respiration rates that offset these photosynthetic fluxes. March–December (primary growing season) totals of NEE were ?63, ?212, and ?233 g C m^?2 in the grassland, shrubland, and woodland, respectively. Thus, there was a greater disparity between ecosystem water use and the strength of the CO₂ sink as woody plants increased across the encroachment gradient. Despite a higher density of woody plants and a greater plant productivity in the woodland than in the shrubland, the woodland produced a larger respiration response to rainfall that largely offset its higher photosynthetic potential. These data suggest that the capacity for woody plants to exploit water resources in riparian areas results in enhanced carbon sequestration at the expense of increased groundwater use under current climate conditions, but the potential does not scale specifically as a function of woody‐plant abundance. These results highlight the important roles of water sources and ecosystem structure on the control of water and carbon balances in dryland areas.     

Temperature and precipitation controls over leaf‐ and ecosystem‐level CO2 flux along a woody plant encroachment gradient

Conversion of grasslands to woodlands may alter the sensitivity of CO₂ exchange of individual plants and entire ecosystems to air temperature and precipitation. We combined leaf‐level gas exchange and ecosystem‐level eddy covariance measurements to quantify the effects of plant temperature sensitivity and ecosystem temperature responses within a grassland and mesquite woodland across seasonal precipitation periods. In so doing, we were able to estimate the role of moisture availability on ecosystem temperature sensitivity under large‐scale vegetative shifts. Optimum temperatures (T_opt) for net photosynthetic assimilation (A) and net ecosystem productivity (NEP) were estimated from a function fitted to A and NEP plotted against air temperature. The convexities of these temperature responses were quantified by the range of temperatures over which a leaf or an ecosystem assimilated 50% of maximum NEP (Ω₅₀). Under dry pre‐ and postmonsoon conditions, leaf‐level Ω₅₀ in C₃ shrubs were two‐to‐three times that of C₄ grasses, but under moist monsoon conditions, leaf‐level Ω₅₀ was similar between growth forms. At the ecosystems‐scale, grassland NEP was more sensitive to precipitation, as evidenced by a 104% increase in maximum NEP at monsoon onset, compared to a 57% increase in the woodland. Also, woodland NEP was greater across all temperatures experienced by both ecosystems in all seasons. By maintaining physiological function across a wider temperature range during water‐limited periods, woody plants assimilated larger amounts of carbon. This higher carbon‐assimilation capacity may have significant implications for ecosystem responses to projected climate change scenarios of higher temperatures and more variable precipitation, particularly as semiarid regions experience conversions from C₄ grasses to C₃ shrubs. As regional carbon models, CLM 4.0, are now able to incorporate functional type and photosynthetic pathway differences, this work highlights the need for a better integration of the interactive effects of growth form/functional type and photosynthetic pathway on water resource acquisition and temperature sensitivity.     

青藏高原木本植物扩张对生长季地表昼夜温度的影响

木本植物扩张或灌丛化是全球性的生态环境问题。近年来青藏高原发生了大规模的木本植物扩张。然而木本植物在青藏高原扩张的时空分布特征及其对局地地表温度(LST)如何影响尚不清楚。基于MODIS土地覆盖产品识别出青藏高原木本植物扩张的空间分布,并利用移动窗口搜索法,探究其对生长季昼夜LST的影响规律及成因。结果表明,2001至2018年木本植物扩张的范围和程度均整体呈增加的趋势。在2018年,木本植物扩张使生长季白天LST降低(2.60±0.34)℃,夜间LST增加(0.94±0.22)℃,净效应使日均LST降低(0.83±0.24)℃。产生这种现象的原因是蒸散发增加((+13.46±6.65)mm/a)等引发的降温效应超过了以反照率减少(-0.031±0.003)为代表的增温效应。气候背景对该影响的空间分布具有相当的控制作用,即降水主导着白天LST的改变,但气温在夜间LST变化中占据更重要的地位。总体上,在气温越低、降水率越高、高程越低的地方发生的木本植物扩张更倾向于降低局地LST。与同一年中越湿润的地方越倾向于降温“相悖”的是,在不同的水文年,更干旱的年份对白天LST具有更强的降温作用,这...     

Impact of woody encroachment on soil organic carbon storage in the Lopé National Park,Gabon

This study quantifies changes in soil organic carbon (SOC) stock as a result of woody encroachment on savannas. Changes in SOC stocks occur below 30 cm depth, indicating the subsoil as the principal compartment contributing to SOC sequestration, and suggesting the need to consider the entire profile (0–100 cm) to thoroughly assess the effect of woody encroachment on SOC stocks.     

The ecological impact of bush encroachment on the yield of grasses in Borana rangeland ecosystem

The ecological impact of woody encroachment and the responses of herbage yield to encroachment were assessed at three locations in Borana rangeland at the end of the growing season. The study was carried out in two communal grazing areas (Medhecho and Dubluk) and one Government ranch (Dida‐Tuyura) in bush and/or shrub encroached and non‐encroached sites. In each area, three altitude ranges were distinguished and in each altitude range one transect, covering both encroached and non‐encroached rangeland, was selected. The assessment was based on the yield and botanical composition of the herbaceous layer. The grasses Cenchrus ciliaris, Chrysopogon aucheri and Panicum coloratum were common or dominant in both encroached and non‐encroached sites. Pennisetum mezianum and Pennise‐tum stramineum were typically found in encroached vegetation. The relative yield increased with non‐encroached sites and varied at different altitude ranges from about 106% to about 150%, thus increases ranged from 75% in Medhecho to 350% in Dubluk as determined from the lower values of the ranges. The encroached vegetation had a significantly lower score for herbage yield than the non‐encroached vegetation for most of the sites, although the differences were small. Differences based on altitude range were also significant for Eragrostis papposa and Pennisetum stramineum, while the three areas showed a significant difference for the mean yield of Aristida adscensionis, Cenchrus ciliaris and Eragrostis papposa.     

Soil phosphorus does not keep pace with soil carbon and nitrogen accumulation following woody encroachment

Soil carbon, nitrogen, and phosphorus cycles are strongly interlinked and controlled through biological processes, and the phosphorus cycle is further controlled through geochemical processes. In dryland ecosystems, woody encroachment often modifies soil carbon, nitrogen, and phosphorus stores, although it remains unknown if these three elements change proportionally in response to this vegetation change. We evaluated proportional changes and spatial patterns of soil organic carbon (SOC), total nitrogen (TN), and total phosphorus (TP) concentrations following woody encroachment by taking spatially explicit soil cores to a depth of 1.2 m across a subtropical savanna landscape which has undergone encroachment by Prosopis glandulosa (an N₂ fixer) and other woody species during the past century in southern Texas, USA. SOC and TN were coupled with respect to increasing magnitudes and spatial patterns throughout the soil profile following woody encroachment, while TP increased slower than SOC and TN in topmost surface soils (0–5 cm) but faster in subsurface soils (15–120 cm). Spatial patterns of TP strongly resembled those of vegetation cover throughout the soil profile, but differed from those of SOC and TN, especially in subsurface soils. The encroachment of woody species dominated by N₂‐fixing trees into this P‐limited ecosystem resulted in the accumulation of proportionally less soil P compared to C and N in surface soils; however, proportionally more P accrued in deeper portions of the soil profile beneath woody patches where alkaline soil pH and high carbonate concentrations would favor precipitation of P as relatively insoluble calcium phosphates. This imbalanced relationship highlights that the relative importance of biotic vs. abiotic mechanisms controlling C and N vs. P accumulation following vegetation change may vary with depth. Our findings suggest that efforts to incorporate effects of land cover changes into coupled climate–biogeochemical models should attempt to represent C‐N‐P imbalances that may arise following vegetation change.     

The effects of warming-shifted plant phenology on ecosystem carbon exchange are regulated by precipitation in a semi-arid grassland

Background

The longer growing season under climate warming has served as a crucial mechanism for the enhancement of terrestrial carbon (C) sink over the past decades. A better understanding of this mechanism is critical for projection of changes in C cycling of terrestrial ecosystems.

Methodology/Principal Findings

A 4-year field experiment with day and night warming was conducted to examine the responses of plant phenology and their influences on plant coverage and ecosystem C cycling in a temperate steppe in northern China. Greater phenological responses were observed under night than day warming. Both day and night warming prolonged the growing season by advancing phenology of early-blooming species but without changing that of late-blooming species. However, no warming response of vegetation coverage was found for any of the eight species. The variances in species-level coverage and ecosystem C fluxes under different treatments were positively dependent upon the accumulated precipitation within phenological duration but not the length of phenological duration.

Conclusions/Significance

These plants'' phenology is more sensitive to night than day warming, and the warming effects on ecosystem C exchange via shifting plant phenology could be mediated by precipitation patterns in semi-arid grasslands.     

草原灌丛化对生态系统多功能性的影响

草原灌丛化现象在干旱半干旱区广泛发生,影响了生态系统的结构、过程和功能。生态系统具有同时提供多种功能的能力,即生态系统多功能性。灌丛化是否会引起草原生态系统多功能性的减少,其内在的作用机制又是什么？这些问题仍有待明晰。理解草原灌丛化对生态系统多功能性的影响,对于促进草原地区"草-畜-人"平衡和实现区域可持续发展至关重要。从响应规律、影响路径和控制因素三个方面总结评述了草原灌丛化对生态系统多功能性影响的研究进展,主要包括:(1)阐明了单一生态系统功能和多种生态系统功能对草原灌丛化的响应特征;(2)从生物路径、非生物路径以及气候变化和人类活动的影响方面探讨了灌丛化对生态系统多功能性的影响路径;(3)从灌丛化物种、灌丛化阶段和草原类型三个方面明晰了草原灌丛化对生态系统多功能性影响的控制因素。在此基础上,针对灌丛化对生态系统多功能性的影响机制,对生产-生态功能权衡的影响等方面对未来研究进行了展望,并面向可持续发展目标探讨了灌丛化生态系统的可持续管理路径。研究可为我国灌丛化草原的恢复和管理提供支撑。     

Multiple trade‐offs regulate the effects of woody plant removal on biodiversity and ecosystem functions in global rangelands

Woody plant encroachment is a major land management issue. Woody removal often aims to restore the original grassy ecosystem, but few studies have assessed the role of woody removal on ecosystem functions and biodiversity at global scales. We collected data from 140 global studies and evaluated how different woody plant removal methods affected biodiversity (plant and animal diversity) and ecosystem functions (plant production, hydrological function, soil carbon) across global rangelands. Our results indicate that the impact of removal is strongly context dependent, varying with the specific response variable, removal method, and traits of the target species. Over all treatments, woody plant removal increased grass biomass and total groundstorey diversity. Physical and chemical removal methods increased grass biomass and total groundstorey biomass (i.e., non‐woody plants, including grass biomass), but burning reduced animal diversity. The impact of different treatment methods declined with time since removal, particularly for total groundstorey biomass. Removing pyramid‐shaped woody plants increased total groundstorey biomass and hydrological function but reduced total groundstorey diversity. Environmental context (e.g., aridity and soil texture) indirectly controlled the effect of removal on biomass and biodiversity by influencing plant traits such as plant shape, allelopathic, or roots types. Our study demonstrates that a one‐size‐fits‐all approach to woody plant removal is not appropriate, and that consideration of woody plant identity, removal method, and environmental context is critical for optimizing removal outcomes. Applying this knowledge is fundamental for maintaining diverse and functional rangeland ecosystems as we move toward a drier and more variable climate.     

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.     

Leaf gas exchange and water status responses of a native and non-native grass to precipitation across contrasting soil surfaces in the Sonoran Desert

Arid and semi-arid ecosystems of the southwestern US are undergoing changes in vegetation composition and are predicted to experience shifts in climate. To understand implications of these current and predicted changes, we conducted a precipitation manipulation experiment on the Santa Rita Experimental Range in southeastern Arizona. The objectives of our study were to determine how soil surface and seasonal timing of rainfall events mediate the dynamics of leaf-level photosynthesis and plant water status of a native and non-native grass species in response to precipitation pulse events. We followed a simulated precipitation event (pulse) that occurred prior to the onset of the North American monsoon (in June) and at the peak of the monsoon (in August) for 2002 and 2003. We measured responses of pre-dawn water potential, photosynthetic rate, and stomatal conductance of native (Heteropogon contortus) and non-native (Eragrostis lehmanniana) C₄ bunchgrasses on sandy and clay-rich soil surfaces. Soil surface did not always amplify differences in plant response to a pulse event. A June pulse event lead to an increase in plant water status and photosynthesis. Whereas the August pulse did not lead to an increase in plant water status and photosynthesis, due to favorable soil moisture conditions facilitating high plant performance during this period. E. lehmanniana did not demonstrate heightened photosynthetic performance over the native species in response to pulses across both soil surfaces. Overall accumulated leaf-level CO₂ response to a pulse event was dependent on antecedent soil moisture during the August pulse event, but not during the June pulse event. This work highlights the need to understand how desert species respond to pulse events across contrasting soil surfaces in water-limited systems that are predicted to experience changes in climate.     

Direct and indirect effects of grazing constrain shrub encroachment in semi‐arid Patagonian steppes

Question: What are the long‐term effects of grazing exclusion on the population structure and dynamics of, and interactions among, three dominant shrub species? Location: Grass‐shrub Patagonian steppe, Chubut, Argentina. Methods: Permanent plots were established in grazed paddocks and paddocks excluded from grazing in representative Patagonian rangelands. Shrub abundance, population size‐structure, short‐term (two 3‐yr periods) and long‐term (matrix models) population dynamics, and neighborhood interactions of three native and codominant shrub species (Mulinum spinosum, Senecio filaginoides and Adesmia volckmanni) were measured and analysed using different statistical approaches. Results: The total density of shrubs was 74% higher in paddocks excluded from grazing, owing mainly to increases in Mulinum (80%) and Senecio (68%) species. However, differences in size structure between ungrazed and grazed paddocks were only detected in Mulinum. Demographic rates differed between shrub species, time‐periods and grazing conditions. In particular, recruitment in the short term (especially in wet years) and population growth rate in the long term (λ) were higher in paddocks excluded from grazing only in Mulinum populations. Senecio populations showed a marginal increase in recruitment and mortality independent of the grazing condition in the wet and dry period. Grazing exclusion modified the balance of neighborhood interactions among the three shrub species. In grazing‐exclusion paddocks, there was a balance between positive and negative interspecific interactions, while in grazed paddocks there were more negative intraspecific and interspecific interactions, resulting in a net negative balance of neighborhood interactions. Conclusions: Our understanding of woody encroachment in arid rangelands can be informed through evaluation of direct and indirect effects of grazing exclusion on the abundance and demography of dominant woody species. In Patagonian arid steppes, the occurrence of woody encroachment in rangelands excluded from grazing can be explained by altered responses in plant‐animal and plant‐plant interactions among shrub species.     

Rates of woody encroachment in African savannas reflect water constraints and fire disturbance

Aim

The aims of this study were to (1) estimate current rates of woody encroachment across African savannas; (2) identify relationships between change in woody cover and potential drivers, including water constraints, fire frequency and livestock density. The found relationships led us to pursue a third goal: (3) use temporal dynamics in woody cover to estimate potential woody cover.

Location

Sub‐Saharan African savannas.

Methods

The study used very high spatial resolution satellite imagery at sites with overlapping older (2002–2006) and newer (2011–2016) imagery to estimate change in woody cover. We sampled 596 sites in 38 separate areas across African savannas. Areas with high anthropogenic impact were avoided in order to more clearly identify the influence of environmental factors. Relationships between woody cover change and potential drivers were identified using linear regression and simultaneous autoregression, where the latter accounts for spatial autocorrelation.

Results

The mean annual change in woody cover across our study areas was 0.25% per year. Although we cannot explain the general trend of encroachment based on our data, we found that change rates were positively correlated with the difference between potential woody cover and actual woody cover (a proxy for water availability; p < .001), and negatively correlated with fire frequency (p < .01). Using the relationship between rates of encroachment and initial cover, we estimated potential woody cover at different rainfall levels.

Main conclusions

The results indicate that woody encroachment is ongoing and widespread across African savannas. The fact that the difference between potential and actual cover was the most significant predictor highlights the central role of water availability and tree–tree competition in controlling change in woody populations, both in water‐limited and mesic savannas. Our approach to derive potential woody cover from the woody cover change trajectories demonstrates that temporal dynamics in woody populations can be used to infer resource limitations.