高寒草地植物群落地上-地下净初级生产力权衡

植物生产力分配和权衡是植物生态学研究中的热点,反映植物对环境的适应性,是了解植物响应全球气候变化的关键。青藏高原作为气候变化敏感区,研究其植物地上与地下部分权衡对了解高寒草地植被生存策略和生态系统可持续发展具有重要意义。目前,生物量分配调控机制已被广泛研究,但主要使用植物根冠比和地上-地下生物量比等方法来表征植物分配模式,缺乏考虑因植物生长周期导致的差异。使用青藏高原高寒草地103个样点的地上和地下净初级生产力数据,分析高寒草甸和高寒草原两种主要草地类型的地上-地下净初级生产力权衡关系。利用气候因素和土壤因素等相关数据,结合方差分析、相关分析、相对重要性分析和结构方程模型的方法,探究环境因素对两种草地类型地上-地下净初级生产力权衡的影响机制。研究发现：（1）高寒草甸的地上净初级生产力、地下净初级生产力和土壤养分含量显著高于高寒草原（P<0.05）;（2）高寒草地植被生产力均向地下权衡（0.0199）,且高寒草原（0.0354）的权衡值高于高寒草甸（0.0173）;（3）结构方程模型发现,年平均降水量、土壤容重和土壤速效氮含量是影响高寒草甸生产力权衡的主导因子,而年平均温度和年平均降水量是影响高寒草原生产力权衡的主导因子。研究表明高寒草甸的生产力权衡主要受气候和土壤因素共同影响,而高寒草原主要受气候因素调节。研究为理解植物地上-地下生物量分配调控机制提供了新的视角和方法,对系统了解高寒草地生物量分配模式和准确预测高寒草地植被动态过程具有指导意义。     

Functional diversity of leaf nitrogen concentrations drives grassland carbon fluxes

Little is known about the role of plant functional diversity for ecosystem‐level carbon (C) fluxes. To fill this knowledge gap, we translocated monoliths hosting communities with four and 16 sown species from a long‐term grassland biodiversity experiment (‘The Jena Experiment’) into a controlled environment facility for ecosystem research (Ecotron). This allowed quantifying the effects of plant diversity on ecosystem C fluxes as well as three parameters of C uptake efficiency (water and nitrogen use efficiencies and apparent quantum yield). By combining data on ecosystem C fluxes with vegetation structure and functional trait‐based predictors, we found that increasing plant species and functional diversity led to higher gross and net ecosystem C uptake rates. Path analyses and light response curves unravelled the diversity of leaf nitrogen concentration in the canopy as a key functional predictor of C fluxes, either directly or indirectly via LAI and aboveground biomass.     

放牧和模拟增温对藏北高寒草地植物群落特征及生产力的影响

气候变化和放牧活动对草地植物物种多样性和生产力具有重要影响。为探索藏北高寒草地植物物种多样性和生产力对增温、放牧及其交互作用的响应, 于2011年在藏北高原开始建立增温实验平台, 2016年起增设放牧、增温+放牧实验, 连续2年(2016-2017年)观测了植物群落特征、群落组成、生产力和物种多样性。结果表明, 增温和放牧对高寒草地植物高度和净初级生产力具有显著交互作用。在放牧条件下, 增温对植物高度无显著影响; 但在不放牧条件下, 增温却显著增加了植物高度。在放牧条件下, 增温对净初级生产力的影响存在年际差异, 2016年增温对生产力无显著影响, 2017年增温显著降低了植物净初级生产力; 但在不放牧条件下, 增温对植物净初级生产力无显著影响。增温和放牧对高寒草地植物物种丰富度、盖度、重要值及多样性均无显著交互作用。植物盖度在增温和放牧条件下显著降低, 杂类草物种比例显著增加, 但物种多样性均无显著变化。研究表明, 增温和放牧显著改变高寒草地群落结构。未来气候变化条件下, 放牧活动加剧有可能导致高寒草地生产力降低。     

Spatial variations in aboveground net primary productivity along a climate gradient in Eurasian temperate grassland: effects of mean annual precipitation and its seasonal distribution

Concomitant changes of annual precipitation and its seasonal distribution within the context of global climate change have dramatic impacts on aboveground net primary productivity (ANPP) of grassland ecosystems. In this study, combining remote sensing products with in situ measurements of ANPP, we quantified the effects of mean annual precipitation (MAP) and precipitation seasonal distribution (PSD) on the spatial variations in ANPP along a climate gradient in Eurasian temperate grassland. Our results indicated that ANPP increased exponentially with MAP for the entire temperate grassland, but linearly for a specific grassland type, i.e. the desert steppe, typical steppe, and meadow steppe from arid to humid regions. The slope of the linear relationship appeared to be steeper in the more humid meadow steppe than that in the drier typical and desert steppes. PSD also had significant effect on the spatial variations in ANPP. It explained 39.4% of the spatial ANPP for the entire grassland investigated, being comparable with the explanatory power of MAP (40.0%). On the other hand, the relative contribution of PSD and MAP is grassland type specific. MAP exhibited a much stronger explanatory power than PSD for the desert steppe and the meadow steppe at the dry and wet end, respectively. However, PSD was the dominant factor affecting the spatial variation in ANPP for the median typical steppe. Our results imply that altered pattern of PSD due to climate change may be as important as the total amount in terms of effects on ANPP in Eurasian temperate grassland.     

Potential effects of warming and drying on peatland plant community composition

Boreal peatlands may be particularly vulnerable to climate change, because temperature regimes that currently constrain biological activity in these regions are predicted to increase substantially within the next century. Changes in peatland plant community composition in response to climate change may alter nutrient availability, energy budgets, trace gas fluxes, and carbon storage. We investigated plant community response to warming and drying in a field mesocosm experiment in northern Minnesota, USA. Large intact soil monoliths removed from a bog and a fen received three infrared warming treatments crossed with three water‐table treatments (n = 3) for five years. Foliar cover of each species was estimated annually. In the bog, increases in soil temperature and decreases in water‐table elevation increased cover of shrubs by 50% and decreased cover of graminoids by 50%. The response of shrubs to warming was distinctly species‐specific, and ranged from increases (for Andromeda glaucophylla) to decreases (for Kalmia polifolia). In the fens, changes in plant cover were driven primarily by changes in water‐table elevation, and responses were species‐ and lifeform‐specific: increases in water‐table elevation increased cover of graminoids – in particular Carex lasiocarpa and Carex livida– as well as mosses. In contrast, decreases in water‐table elevation increased cover of shrubs, in particular A. glaucophylla and Chamaedaphne calyculata. The differential and sometimes opposite response of species and lifeforms to the treatments suggest that the structure and function of both bog and fen plant communities will change – in different directions or at different magnitudes – in response to warming and/or changes in water‐table elevation that may accompany regional or global climate change.     

凋落物输入变化和氮添加对半干旱草原群落生产力及功能群组成的影响

不同的草原利用方式(围封、放牧和割草等)随着大气氮沉降的不断加剧, 改变了凋落物输入量。凋落物作为连接地上-地下碳循环过程的关键环节, 对草原生态系统生产力和碳循环过程影响显著。氮是草原生产力的主要限制因子, 凋落物输入量的变化对草原生态系统结构和功能的影响仍缺乏长期实验证据支持。该研究在内蒙古半干旱典型草原建立一个凋落物输入变化和氮添加控制实验平台, 通过连续6年对群落生产力和功能群组成的监测, 研究了凋落物添加与去除和氮添加对半干旱草原群落生产力和功能群组成的影响。研究发现: 1)凋落物输入量增加和氮添加均显著提高了群落生产力, 在对照和氮添加处理下, 凋落物去除处理导致生产力分别降低了8.4%和7.6%, 而凋落物添加处理使生产力分别提高了10.7%和6.3%; 2)不同植物功能群对凋落物输入变化和氮添加的响应存在差异, 导致群落功能群结构发生变化。随着凋落物输入量增加和氮添加, 群落优势功能群多年生禾草(包括多年生丛生禾草和多年生根茎禾草)的生物量显著提高, 对群落生产力的贡献增加, 在群落中的优势地位增强; 而另一优势功能群多年生杂类草生物量对凋落物和氮添加处理均无显著响应, 进而导致在氮添加处理下其对群落生物量的贡献比例显著降低; 3)凋落物输入主要改善土壤水分状况, 而氮添加则主要通过提高土壤养分含量, 促进群落生产力, 并通过影响主要功能群生物量, 导致群落结构发生变化。以上结果表明, 适当的草原管理方式如围封禁牧和降低放牧强度等都能通过增加凋落物的输入来提高草原生产力, 维持生态系统稳定性。而适量的氮等养分添加管理也有助于提高草原生产力, 促进其恢复。     

Abrupt declines in marine phytoplankton production driven by warming and biodiversity loss in a microcosm experiment

Rising sea surface temperatures are expected to lead to the loss of phytoplankton biodiversity. However, we currently understand very little about the interactions between warming, loss of phytoplankton diversity and its impact on the oceans' primary production. We experimentally manipulated the species richness of marine phytoplankton communities under a range of warming scenarios, and found that ecosystem production declined more abruptly with species loss in communities exposed to higher temperatures. Species contributing positively to ecosystem production in the warmed treatments were those that had the highest optimal temperatures for photosynthesis, implying that the synergistic impacts of warming and biodiversity loss on ecosystem functioning were mediated by thermal trait variability. As species were lost from the communities, the probability of taxa remaining that could tolerate warming diminished, resulting in abrupt declines in ecosystem production. Our results highlight the potential for synergistic effects of warming and biodiversity loss on marine primary production.     

Effects of climate warming on net primary productivity in China during 1961–2010

The response of ecosystems to different magnitudes of climate warming and corresponding precipitation changes during the last few decades may provide an important reference for predicting the magnitude and trajectory of net primary productivity (NPP) in the future. In this study, a process‐based ecosystem model, Carbon Exchange between Vegetation, Soil and Atmosphere (CEVSA), was used to investigate the response of NPP to warming at both national and subregional scales during 1961–2010. The results suggest that a 1.3°C increase in temperature stimulated the positive changing trend in NPP at national scale during the past 50 years. Regardless of the magnitude of temperature increase, warming enhanced the increase in NPP; however, the positive trend of NPP decreased when warming exceeded 2°C. The largest increase in NPP was found in regions where temperature increased by 1–2°C, and this rate of increase also contributed the most to the total increase in NPP in China's terrestrial ecosystems. Decreasing precipitation depressed the positive trend in NPP that was stimulated by warming. In northern China, warming depressed the increasing trend of NPP and warming that was accompanied by decreasing precipitation led to negative changing trends in NPP in large parts of northern China, especially when warming exceeded 2°C. However, warming stimulated the increase in NPP until warming was greater than 2°C, and decreased precipitation helped to increase the NPP in southern China.     

Effect of altered litter input and nitrogen addition on ecosystem aboveground primary productivity and plant functional group composition in a semiarid grassland

不同的草原利用方式(围封、放牧和割草等)随着大气氮沉降的不断加剧, 改变了凋落物输入量。凋落物作为连接地上-地下碳循环过程的关键环节, 对草原生态系统生产力和碳循环过程影响显著。氮是草原生产力的主要限制因子, 凋落物输入量的变化对草原生态系统结构和功能的影响仍缺乏长期实验证据支持。该研究在内蒙古半干旱典型草原建立一个凋落物输入变化和氮添加控制实验平台, 通过连续6年对群落生产力和功能群组成的监测, 研究了凋落物添加与去除和氮添加对半干旱草原群落生产力和功能群组成的影响。研究发现: 1)凋落物输入量增加和氮添加均显著提高了群落生产力, 在对照和氮添加处理下, 凋落物去除处理导致生产力分别降低了8.4%和7.6%, 而凋落物添加处理使生产力分别提高了10.7%和6.3%; 2)不同植物功能群对凋落物输入变化和氮添加的响应存在差异, 导致群落功能群结构发生变化。随着凋落物输入量增加和氮添加, 群落优势功能群多年生禾草(包括多年生丛生禾草和多年生根茎禾草)的生物量显著提高, 对群落生产力的贡献增加, 在群落中的优势地位增强; 而另一优势功能群多年生杂类草生物量对凋落物和氮添加处理均无显著响应, 进而导致在氮添加处理下其对群落生物量的贡献比例显著降低; 3)凋落物输入主要改善土壤水分状况, 而氮添加则主要通过提高土壤养分含量, 促进群落生产力, 并通过影响主要功能群生物量, 导致群落结构发生变化。以上结果表明, 适当的草原管理方式如围封禁牧和降低放牧强度等都能通过增加凋落物的输入来提高草原生产力, 维持生态系统稳定性。而适量的氮等养分添加管理也有助于提高草原生产力, 促进其恢复。     

短期氮添加对祁连山亚高山草地生产力及植物多样性的影响

全球氮沉降速率的急剧增加已显著地改变了生态系统的生产力及稳定性,特别是在受N限制较严重的亚高山草地生态系统。虽然氮沉降增加对草地生产力和植物多样性影响的研究报道已经很多,但是氮素沉降的生态系效应因气候区、草地系统类型、加氮水平、氮肥类型和试验时间长短等不同而差别很大。为了评估氮沉降增加对亚高山草地植物物种多样性和生产力的影响,通过在祁连山中部亚高山草地设置不同氮添加水平（0、2、5、10、15、25 g N m^-2 a^-1和50 g N m^-2 a^-1）的短期氮沉降增加模拟试验,探讨了生产力和物种多样性对不同水平氮添加的响应。结果显示：氮添加增加了禾本科（垂穗披碱草、赖草和草地早熟禾）和莎草科（矮嵩草）的地上生产力及其在群落生产力中所占的比例,主要表现在氮添加增加了禾本科和莎草科的株高和株数,降低了其他科（鹅绒委陵菜和葛缕子）的株高和株数;与生产力相比,植物多样性对氮添加的响应较慢,总体随着氮添加量的增加呈下降趋势但未达到显著水平;植物多样性与生产力呈显著的负相关关系。研究结果表明氮添加有助于提高禾本科和莎草科的生产力,进而提高群落生产力,但其他科的植物会被逐渐替代,导致群落植物物种多样性降低。研究结果可为我国亚高山草地的持续性管理提供一定的理论基础。     

放牧对贝加尔针茅草原群落植物多样性和生产力的影响

研究了不同放牧强度对贝加尔针茅草原群落植物多样性和生产力的影响。结果表明,在牧压梯度上,不同的植物表现出不同的生态适应对策,贝加尔针茅种群随着牧压的增加,种群株丛破碎化、小型化,羊草耐牧性较强,在中牧阶段生产力最高。群落初级生产力随着放牧强度的增加逐渐下降。较大的放牧压力下,群落中适口性差、耐牧的杂类草植物渐趋增加。而在更大的放牧压力下,群落逐步被耐牧的小丛生禾草、旱生小苔草、小灌木和灌木所替代。草地群落的稳定性随着放牧强度增加逐渐降低。放牧干扰对群落植物多样性和生产力及其稳定性的影响是不同步的,不对称的。植物群落初级生产力对放牧干扰的响应更迅速,变化更剧烈。     

气温上升对草地土壤微生物群落结构的影响

在 2 0世纪内 ,全球气温已经上升了 0 .6℃ ,并预计到本世纪末仍将上升 1.4～ 5 .8℃。全球气候变暖对生态系统的潜在影响 ,生态系统对气温上升的反馈已成为国际生态学界的研究热点 ,而且所研究的系统也已经从过去简化的模拟系统到复杂的真实生态系统。但是 ,现有对真实生态系统的研究大部分集中在地上植物群落和土壤气体交换等领域 ,对在土壤有机碳分解和保护中起决定作用的土壤微生物研究较少。为此 ,在美国大平原地区进行人工提高气温 (上升 1.8℃ ) ,来研究土壤微生物对气温上升的反应。结果表明 :增温对土壤微生物的总生物量没有显著效应 ,但可以提高微生物的 C∶ N比。另外 ,磷脂肪酸分析发现 ,气温上升显著降低土壤微生物量中的细菌比重 ,提高真菌的份额 ,从而显著提高了群落中真菌与细菌的比值。而且 ,通过对土壤微生物底物利用方式和磷脂肪酸特征的主成份分析 ,发现增温导致了土壤微生物群落结构的转变。可见 ,气温上升可能是通过提高土壤微生物中真菌的优势 ,而导致群落结构的变化。该变化将可以提高微生物对土壤有机碳的利用效率 ,并利于土壤有机碳的保护     

21世纪上半叶内蒙古草地植被净初级生产力变化趋势

基于中国气象局国家气候中心新发布的中短期适应气候变化的新情景(RCP4.5)和极端情景(RCP8.5)下的气候预估数据,采用空间化后的CENTURY模型模拟探讨2011－2050年内蒙古草地植被净初级生产力(NPP)的时空变化特征.结果表明: 区域尺度上,未来气候变化情景下内蒙古草地NPP年下降速率分别为0.57 g C·m^-2·a^-1(RCP4.5)、0.89 g C·m^-2·a^-1(RCP8.5);相对于基准时段,RCP4.5情景下内蒙古草地NPP在2020s、2030s、2040s分别下降11.6%、12.0%、18.0%,而RCP8.5情景下降幅分别为23.8%、21.2%、30.1%.不同气候情景下内蒙古草地NPP时空变化特征差异较大,但即使在RCP4.5下未来40年绝大部分草地NPP也将呈现下降趋势,15.6%的草地减产超过20%.这表明未来气候变化情景下内蒙古草地降水略增的态势不足以补偿因温度升高对草地植被初级生产力所产生的负面作用,草地资源的可持续发展将面临更大挑战.     

A landscape‐scale assessment of the relationship between grassland functioning,community diversity,and functional traits

Livestock farmers rely on a high and stable grassland productivity for fodder production to sustain their livelihoods. Future drought events related to climate change, however, threaten grassland functionality in many regions across the globe. The introduction of sustainable grassland management could buffer these negative effects. According to the biodiversity–productivity hypothesis, productivity positively associates with local biodiversity. The biodiversity–insurance hypothesis states that higher biodiversity enhances the temporal stability of productivity. To date, these hypotheses have mostly been tested through experimental studies under restricted environmental conditions, hereby neglecting climatic variations at a landscape‐scale. Here, we provide a landscape‐scale assessment of the contribution of species richness, functional composition, temperature, and precipitation on grassland productivity. We found that the variation in grassland productivity during the growing season was best explained by functional trait composition. The community mean of plant preference for nutrients explained 24.8% of the variation in productivity and the community mean of specific leaf area explained 18.6%, while species richness explained only 2.4%. Temperature and precipitation explained an additional 22.1% of the variation in productivity. Our results indicate that functional trait composition is an important predictor of landscape‐scale grassland productivity.     

Response of plant species richness and primary productivity in shrublands along a north–south gradient in Europe to seven years of experimental warming and drought: reductions in primary productivity in the heat and drought year of 2003

We used a nonintrusive field experiment carried out at six sites – Wales (UK), Denmark (DK), the Netherlands (NL), Hungary (HU), Sardinia (Italy – IT), and Catalonia (Spain – SP) – along a climatic and latitudinal gradient to examine the response of plant species richness and primary productivity to warming and drought in shrubland ecosystems. The warming treatment raised the plot daily temperature by ca. 1 °C, while the drought treatment led to a reduction in soil moisture at the peak of the growing season that ranged from 26% at the SP site to 82% in the NL site. During the 7 years the experiment lasted (1999–2005), we used the pin‐point method to measure the species composition of plant communities and plant biomass, litterfall, and shoot growth of the dominant plant species at each site. A significantly lower increase in the number of species pin‐pointed per transect was found in the drought plots at the SP site, where the plant community was still in a process of recovering from a forest fire in 1994. No changes in species richness were found at the other sites, which were at a more mature and stable state of succession and, thus less liable to recruitment of new species. The relationship between annual biomass accumulation and temperature of the growing season was positive at the coldest site and negative at the warmest site. The warming treatment tended to increase the aboveground net primary productivity (ANPP) at the northern sites. The relationship between annual biomass accumulation and soil moisture during the growing season was not significant at the wettest sites, but was positive at the driest sites. The drought treatment tended to reduce the ANPP in the NL, HU, IT, and SP sites. The responses to warming were very strongly related to the Gaussen aridity index (stronger responses the lower the aridity), whereas the responses to drought were not. Changes in the annual aboveground biomass accumulation, litterfall, and, thus, the ANPP, mirrored the interannual variation in climate conditions: the most outstanding change was a decrease in biomass accumulation and an increase in litterfall at most sites during the abnormally hot year of 2003. Species richness also tended to decrease in 2003 at all sites except the cold and wet UK site. Species‐specific responses to warming were found in shoot growth: at the SP site, Globularia alypum was not affected, while the other dominant species, Erica multiflora, grew 30% more; at the UK site, Calluna vulgaris tended to grow more in the warming plots, while Empetrum nigrum tended to grow less. Drought treatment decreased plant growth in several studied species, although there were some species such as Pinus halepensis at the SP site or C. vulgaris at the UK site that were not affected. The magnitude of responses to warming and drought thus depended greatly on the differences between sites, years, and species and these multiple plant responses may be expected to have consequences at ecosystem and community level. Decreases in biodiversity and the increase in E. multiflora growth at the SP site as a response to warming challenge the assumption that sensitivity to warming may be less well developed at more southerly latitudes; likewise, the fact that one of the studied shrublands presented negative ANPP as a response to the 2003 heat wave also challenges the hypothesis that future climate warming will lead to an enhancement of plant growth and carbon sequestration in temperate ecosystems. Extreme events may thus change the general trend of increased productivity in response to warming in the colder sites.     

Food webs obscure the strength of plant diversity effects on primary productivity

Plant diversity experiments generally find that increased diversity causes increased productivity; however, primary productivity is typically measured in the presence of a diverse food web, including pathogens, mutualists and herbivores. If food web impacts on productivity vary with plant diversity, as predicted by both theoretical and empirical studies, estimates of the effect of plant diversity on productivity may be biased. We experimentally removed arthropods, foliar fungi and soil fungi from the longest‐running plant diversity experiment. We found that fungi and arthropods removed a constant, large proportion of biomass leading to a greater reduction of total biomass in high diversity plots. As a result, the effect of diversity on measured plant productivity was much higher in the absence of fungi and arthropods. Thus, diversity increases productivity more than reported in previous studies that did not control for the effects of heterotrophic consumption.     

Sensitivity of plant species to warming and altered precipitation dominates the community productivity in a semiarid grassland on the Loess Plateau

Global warming and changes in precipitation patterns can critically influence the structure and productivity of terrestrial ecosystems. However, the underlying mechanisms are not fully understood. We conducted two independent but complementary experiments (one with warming and precipitation manipulation (+ or – 30%) and another with selective plant removal) in a semiarid grassland on the Loess Plateau, northwestern China, to assess how warming and altered precipitation affect plant community. Our results showed that warming and altered precipitation affected community aboveground net primary productivity (ANPP) through impacting soil moisture. Results of the removal experiment showed competitive relationships among dominant grasses, the dominant subshrub and nondominant species, which played a more important role than soil moisture in the response of plant community to warming and altered precipitation. Precipitation addition intensified the competition but primarily benefited the dominant subshrub. Warming and precipitation reduction enhanced water stresses but increased ANPP of the dominant subshrub and grasses, indicating that plant tolerance to drought critically meditated the community responses. These findings suggest that specie competitivity for water resources as well as tolerance to environmental stresses may dominate the responses of plant communities on the Loess Plateaus to future climate change factors.     

Deepened winter snow cover enhances net ecosystem exchange and stabilizes plant community composition and productivity in a temperate grassland

Global warming has greatly altered winter snowfall patterns, and there is a trend towards increasing winter snow in semi‐arid regions in China. Winter snowfall is an important source of water during early spring in these water‐limited ecosystems, and it can also affect nutrient supply. However, we know little about how changes in winter snowfall will affect ecosystem productivity and plant community structure during the growing season. Here, we conducted a 5‐year winter snow manipulation experiment in a temperate grassland in Inner Mongolia. We measured ecosystem carbon flux from 2014 to 2018 and plant biomass and species composition from 2015 to 2018. We found that soil moisture increased under deepened winter snow in early growing season, particularly in deeper soil layers. Deepened snow increased the net ecosystem exchange of CO₂ (NEE) and reduced intra‐ and inter‐annual variation in NEE. Deepened snow did not affect aboveground plant biomass (AGB) but significantly increased root biomass. This suggested that the enhanced NEE was allocated to the belowground, which improved water acquisition and thus contributed to greater stability in NEE in deep‐snow plots. Interestingly, the AGB of grasses in the control plots declined over time, resulting in a shift towards a forb‐dominated system. Similar declines in grass AGB were also observed at three other locations in the region over the same time frame and are attributed to 4 years of below‐average precipitation during the growing season. By contrast, grass AGB was stabilized under deepened winter snow and plant community composition remained unchanged. Hence, our study demonstrates that increased winter snowfall may stabilize arid grassland systems by reducing resource competition, promoting coexistence between plant functional groups, which ultimately mitigates the impacts of chronic drought during the growing season.