Above- and belowground responses to nitrogen addition in a Chihuahuan Desert grassland

Increased available soil nitrogen can increase biomass, lower species richness, alter soil chemistry and modify community structure in herbaceous ecosystems worldwide. Although increased nitrogen availability typically increases aboveground production and decreases species richness in mesic systems, the impacts of nitrogen additions on semiarid ecosystems remain unclear. To determine how a semiarid grassland responds to increased nitrogen availability, we examined plant community structure and above- and belowground net primary production in response to long-term nitrogen addition in a desert grassland in central New Mexico, USA. Plots were fertilized annually (10 g N m⁻²) since 1995 and NPP measured from 2004 to 2009. Differences in aboveground NPP between fertilized and control treatments occurred in 2004 following a prescribed fire and in 2006 when precipitation was double the long-term average during the summer monsoon. Presumably, nitrogen only became limiting once drought stress was alleviated. Belowground NPP was also related to precipitation, and greatest root growth occurred the year following the wettest summer, decreasing gradually thereafter. Belowground production was unrelated to aboveground production within years and unrelated to nitrogen enrichment. Species richness changed between years in response to seasonal precipitation variability, but was not altered by nitrogen addition. Community structure did respond to nitrogen fertilization primarily through increased abundance of two dominant perennial grasses. These results were contrary to most nitrogen addition studies that find increased biomass and decreased species richness with nitrogen fertilization. Therefore, factors other than nitrogen deposition, such as fire or drought, may play a stronger role in shaping semiarid grassland communities than soil fertility.     

Effects of plant species richness on stand structure and productivity

Aims Aboveground biomass production commonly increases with species richness in plant biodiversity experiments. Little is known about the direct mechanisms that cause this result. We tested if by occupying different heights and depths above and below ground, and by optimizing the vertical distribution of leaf nitrogen, species in mixtures can contribute to increased resource uptake and, thus, increased productivity of the community in comparison with monocultures.Methods We grew 24 grassland plant species, grouped into four nonoverlapping species pools, in monoculture and 3- and 6-species mixture in spatially heterogeneous and uniform soil nutrient conditions. Layered harvests of above- and belowground biomass, as well as leaf nitrogen and light measurements, were taken to assess vertical canopy and root space structure.Important findings The distribution of leaf mass was shifted toward greater heights and light absorption was correspondingly enhanced in mixtures. However, only some mixtures had leaf nitrogen concentration profiles predicted to optimize whole-community carbon gain, whereas in other mixtures species seemed to behave more 'selfish'. Nevertheless, even in these communities, biomass production increased with species richness. The distribution of root biomass below ground did not change from monocultures to three- and six-species mixtures and there was also no indication that mixtures were better than monocultures at extracting heterogeneously as compared to homogeneously distributed soil resources. We conclude that positive biodiversity effect on aboveground biomass production cannot easily be explained by a single or few common mechanisms of differential space use. Rather, it seems that mechanisms vary with the particular set of species combined in a community.     

Does aboveground vegetation composition resemble soil seed bank during succession in specialized vegetation on gypsum soil?

This paper evaluates the aboveground vegetation in relation to the soil seed bank throughout a 60-year succession process following agricultural abandonment in a semi-arid Mediterranean gypsum habitat. There is little information regarding the relationship between these two community components in the context of succession on semi-arid gypsum soils. Aboveground vegetation and the corresponding seed bank of gypsum plant communities were sampled through a chronosequence of 24 abandoned fields. Generalized linear models were used to model seed species richness and density, redundancy analyses to model the effect of time since abandonment and the effect of soil physicochemical parameters on seed bank species composition, and Mantel tests to analyze resemblance between above- and belowground species composition. In this last case, the effect of time since abandonment was controlled using a partial Mantel test. Mantel correlograms using time intervals instead of distances were used to describe the resemblance of above- to belowground species occurrence in different aged fields. No significant variability in seed species richness, seed density, or species composition due to time since abandonment was found. Differences in seed species composition were mainly due to small spatial scale predictors such as slope and soil calcium content. High correlations between species composition in the soil seed bank and the aboveground vegetation were detected during succession. The lack of a significant trend in aboveground species replacement over time was also reflected in seed bank composition. We concluded that the rapid establishment of strict gypsophyte species relied mainly on the long-term persistence of these species in the seed bank.     

放牧干扰下高寒草甸物种和生活型丰富度与地上及地下生物量的关系

明晰放牧干扰下高寒草甸植物丰富度与生物量的相关关系,为草地植物不同生长时期生物量的预测提供依据。设置6个放牧强度样地,连续3a放牧,2014年进行3个季节(6月、8月、10月)的植物丰富度和地上、地下生物量调查,对比分析放牧干扰下物种和生活型丰富度(生活型的种类)分别与地上、地下生物量的相关关系。结果表明:(1)物种和生活型丰富度与地上生物量均受放牧强度的显著影响,物种丰富度仅在8月与放牧强度显著负相关,生活型丰富度在10月随放牧强度单峰变化,地上生物量在不同季节均与放牧强度显著负相关,而地下生物量与放牧强度无关。(2)物种丰富度与地上和地下生物量均受季节的显著影响,物种丰富度和地上生物量仅在低强度放牧区随季节呈单峰变化,地下生物量在中等强度放牧区随季节呈单峰变化;生活型丰富度与季节无关。(3)放牧干扰前物种和生活型丰富度与地上和地下生物量均显著正相关。3a放牧后仅在8月,物种丰富度只与地上生物量显著正相关,生活型丰富度与地上和地下生物量均显著正相关。(4)对于不同放牧强度,物种丰富度仅在低强度放牧区与地上生物量显著正相关,而生活型丰富度在所有放牧强度区均与地上生物量显著正相关。综上所述,放牧干扰扰乱了高寒草甸丰富度与生物量之间的关系,尤其影响了物种丰富度与地下生物量之间的相关关系。生活型丰富度与地上生物量之间的显著关系不受放牧强度干扰,使生活型丰富度在预测生物量方面表现出优势。     

Density may alter diversity–productivity relationships in experimental plant communities

Contemporary biodiversity experiments, in which plant species richness is manipulated and aboveground productivity of the system measured, generally demonstrate that lowering plant species richness reduces productivity. However, we propose that community density may in part compensate for this reduction of productivity at low diversity. We conducted a factorial experiment in which plant functional group richness was held constant at three, while plant species richness increased from three to six to 12 species and community density from 440 to 1050 to 2525 seedlings m⁻². Response variables included density, evenness and above- and belowground biomass at harvest. The density gradient converged slightly during the course of the experiment due to about 10% mortality at the highest sowing density. Evenness measured in terms of aboveground biomass at harvest significantly declined with density, but the effect was weak. Overall, aboveground, belowground and total biomass increased significantly with species richness and community density. However, a significant interaction between species richness and community density occurred for both total and aboveground biomass, indicating that the diversity–productivity relationship was flatter at higher than at lower density. Thus, high species richness enabled low-density communities to reach productivity levels otherwise seen only at high density. The relative contributions of the three functional groups C₃, C₄ and nitrogen-fixers to aboveground biomass were less influenced by community density at high than at low species richness. We interpret the interaction effects between community density and species richness on community biomass by expanding findings about constant yield and size variation from monocultures to plant mixtures.     

Linking Above- and Belowground Responses to 16 Years of Fertilization,Mowing, and Removal of the Dominant Species in a Temperate Grassland

Species-rich oligotrophic meadows are affected by a wide range of management interventions that influence their functioning and capacity to deliver ecosystem services, but long-term studies on the above- and belowground adaptations to different management tools are still scarce. We focused on the interactive effects of NPK fertilization, mowing, and removal of the initially dominant species (Molinia caerulea) on plant, soil, and microbial responses in wet oligotrophic grassland in a 16-year full-factorial manipulative experiment. Changes in vegetation composition, soil pH, and nutrient availability were accompanied by altered microbial phospholipid fatty acid (PLFA) composition, whereas treatment effects on soil microbial biomass and carbon (C) mineralization were mainly related to changes in soil organic matter (SOM) content and nutrient availability. Fertilization decreased plant species richness aboveground and lowered SOM storage and microbial activity belowground. Mowing preserved high plant diversity and led to more efficient recycling of N within the grassland, whereas Molinia removal significantly affected only plant community composition. Mowing combined with fertilization maintained high species richness only in the short term. Belowground, mowing reduced N leaching from the fertilized system but did not prevent SOM depletion, soil acidification, and concomitant adverse effects on soil microbes. We conclude that annual mowing is the appropriate type of extensive management for oligotrophic species-rich meadows, but the concomitant nutrient depletion should not be compensated for by regular NPK fertilization due to its adverse effects on soil quality.     

Differential responses of soil bacteria,fungi, archaea and protists to plant species richness and plant functional group identity

Plants are known to influence belowground microbial community structure along their roots, but the impacts of plant species richness and plant functional group (FG) identity on microbial communities in the bulk soil are still not well understood. Here, we used 454‐pyrosequencing to analyse the soil microbial community composition in a long‐term biodiversity experiment at Jena, Germany. We examined responses of bacteria, fungi, archaea, and protists to plant species richness (communities varying from 1 to 60 sown species) and plant FG identity (grasses, legumes, small herbs, tall herbs) in bulk soil. We hypothesized that plant species richness and FG identity would alter microbial community composition and have a positive impact on microbial species richness. Plant species richness had a marginal positive effect on the richness of fungi, but we observed no such effect on bacteria, archaea and protists. Plant species richness also did not have a large impact on microbial community composition. Rather, abiotic soil properties partially explained the community composition of bacteria, fungi, arbuscular mycorrhizal fungi (AMF), archaea and protists. Plant FG richness did not impact microbial community composition; however, plant FG identity was more effective. Bacterial richness was highest in legume plots and lowest in small herb plots, and AMF and archaeal community composition in legume plant communities was distinct from that in communities composed of other plant FGs. We conclude that soil microbial community composition in bulk soil is influenced more by changes in plant FG composition and abiotic soil properties, than by changes in plant species richness per se.     

The relationship between species richness and productivity in four typical grasslands of northern China

The relationship between plant species richness and primary productivity has long been acentral topic in biodiversity research.In this paper,we examine the relationship between species richness and productivity in four typical grasslands of Northern China at different spatial scales.At the community scale,a positive correlation was found for six of seven communities.A unimodal pattern was found only for one community (Stipa glareosa community),while at a large scale (vegetation type or landscape/region),the relationship was also found significantly positive.Species richness ranged from 4 to 35 species,and community aboveground productiand aboveground productivity were found in alpine meadow,followed by meadow steppe,typical steppe and desert steppe.     

黄土高原典型草原群落结构和土壤水分对划破的响应

划破是草原改良的基础措施之一,划破强度是划破措施的关键环节,划破对草原健康持续管理有重要意义。目前的研究主要集中在划破对植物群落结构和生产力的影响上,然而草原植物群落与土壤水分对划破强度的响应尚不清楚。在黄土高原典型草原开展不同程度的草地划破试验,探究不同划破强度（27.4%、46.3%和61.9%）对草地植物群落物种多样性、生物量和土壤水分的影响。结果表明：3个划破强度下划破带物种丰富度显著低于未划破带1-3种/m²,划破带和未划破带群落相似性分别低于整区23.85%-119.23%和44.43%-84.55%。地上生物量随物种丰富度的增加而增大,且地上生物量与Simpson指数和Shannon Weiner指数显著负相关。3个划破强度下未划破带地下生物量和总生物量分别高于划破带88.2%-134.6%、52.4%-67.8%、2.5%-16.6%和103.9%-152.9%、59.3%-75.8%、9.1%-22.6%。植物群落物种丰富度和地上生物量随划破强度的增加呈"驼峰"型曲线变化,当划破强度分别为43.7%-55.3%和43.8%-45.7%时,植物群落物种丰富度和生物量均最高。本试验阐明了划破对典型草原植物群落特征和土壤水分的作用机制,研究结果为采用划破措施实现草地培育和草原修复提供了科学依据,对保护草地生物多样性和提高生产力具有重要意义。     

Additive effects of functionally dissimilar above- and belowground organisms on a grassland plant community

Aims In nature, plant communities are affected simultaneously by a variety of functionally dissimilar organisms both above and below the ground. However, there is a gap of knowledge on interactive effects of functionally dissimilar organisms on plant communities that is needed to be filled to better understand and predict the general impact of biotic factors on plant communities.Methods We conducted a full-factorial mesocosm study to investigate the individual and combined impacts of above- and belowground functionally dissimilar organisms on a grassland plant community. We studied the effects of aboveground herbivores (Helix aspersa, Gastropoda), arbuscular mycorrhizal fungi (AMF; Glomus spp., Glomeromycota) and endogeic earthworms (Aporrectodea spp., Lumbricidae) on the diversity, structure and productivity of an experimental grassland plant community and each other.Important findings Aboveground herbivory by snails decreased, AMF increased and earthworms had no effects on the diversity of the grassland plant community, while their combined effects were additive. The biomass of the plant community was negatively affected by snails and AMF, while no effects of earthworms or interaction effects were found. The plant species were differently affected by snails and AMF. No effects of the above- and belowground organisms on each other's performance were detected. Since the effects of the functionally dissimilar organisms on the grassland plant community were mainly independent, the results indicate that their combined effects may be predicted by knowing the individual effects, at least under the conditions used in the present mesocosm study.     

Responses of Plant Community Composition and Biomass Production to Warming and Nitrogen Deposition in a Temperate Meadow Ecosystem

Climate change has profound influences on plant community composition and ecosystem functions. However, its effects on plant community composition and biomass production are not well understood. A four-year field experiment was conducted to examine the effects of warming, nitrogen (N) addition, and their interactions on plant community composition and biomass production in a temperate meadow ecosystem in northeast China. Experimental warming had no significant effect on plant species richness, evenness, and diversity, while N addition highly reduced the species richness and diversity. Warming tended to reduce the importance value of graminoid species but increased the value of forbs, while N addition had the opposite effect. Warming tended to increase the belowground biomass, but had an opposite tendency to decrease the aboveground biomass. The influences of warming on aboveground production were dependent upon precipitation. Experimental warming had little effect on aboveground biomass in the years with higher precipitation, but significantly suppressed aboveground biomass in dry years. Our results suggest that warming had indirect effects on plant production via its effect on the water availability. Nitrogen addition significantly increased above- and below-ground production, suggesting that N is one of the most important limiting factors determining plant productivity in the studied meadow steppe. Significant interactive effects of warming plus N addition on belowground biomass were also detected. Our observations revealed that environmental changes (warming and N deposition) play significant roles in regulating plant community composition and biomass production in temperate meadow steppe ecosystem in northeast China.     

Larger aboveground neighbourhood scales maximise similarity but do not eliminate discrepancies with belowground plant diversity in a Mediterranean shrubland

Plant and Soil - An unresolved question in plant ecology is whether diversity of the aboveground and belowground compartments of a plant community is similar at different neighbourhood scales. We...     

Differentiation between responses of primary productivity and phosphorus exploitation to species richness

Several studies have shown that ecosystem functioning increases with increasing species richness. Most of these studies examined the effects of species richness on primary productivity. The underlying mechanism that explains this pattern is usually the selection effect. The higher the diversity in plant communities the higher the chance in including a very productive species that dominates the community, or a legume species that brings N into the soil. Less attention has been given so far to the effects of species richness on phosphorus exploitation. The aim of this work was to investigate the effect of species richness on aboveground primary productivity and P accumulation in a plant diversity experiment. For this reason, 14 grassland plant species were grown in containers as monocultures and in mixtures of 2-, 3-, 4-, 8-, 11- and 14-species combinations. Results show that the aboveground phytomass and total P increased with increasing species richness. Complementarity effects, probably through partitioning of resources, were most apparent in the highest levels of species richness, and were observed to be greater for total P in comparison to phytomass. Selection effects generally were greater for phytomass than for total P; they were significantly positive at the 2- to 8-species combinations but close to 0 or negative in the highest levels of species richness. The increases in phytomass and total P at the highest levels of species richness appeared to be caused by the increased performance of intermediate-productive species. Responsible Editor: Tibor Kalapos.     

黑河中游荒漠草地地上和地下生物量的分配格局

草地生态系统中地上和地下生物量的分配方式对于研究生态系统碳储量和碳循环有着重要的意义。为了解黑河中游荒漠草地的地上和地下生物量分配格局, 从群落和个体两个水平对黑河中游的地上和地下生物量进行了调查。结果表明: 群落水平上地上生物量介于3.2-559.2 g·m^-2之间, 地下生物量介于3.3-188.2 g·m^-2之间, 个体水平上地上生物量介于6.1-489.0 g·株^-1之间, 地下生物量介于2.4-244.2 g·株^-1之间, 群落水平上的根冠比(R/S)为0.10-2.49, 个体水平上为0.07-1.55, 地下生物量均小于地上生物量, 群落水平上R/S值大于个体水平。群落和个体水平地上和地下生物量的拟合斜率分别为1.1001和0.9913, 与1没有显著差异, 说明地上与地下生物量呈等速生长关系。群落和个体水平土壤表层0-20 cm和0-30 cm的根系生物量分别占全部根系生物量的89.81%、96.95%和81.42%、93.62%, 表明地下生物量主要集中在0-20 cm和0-30 cm土壤表层。     

Above- and below-ground impacts of introduced predators in seabird-dominated island ecosystems

Predators often exert multi-trophic cascading effects in terrestrial ecosystems. However, how such predation may indirectly impact interactions between above- and below-ground biota is poorly understood, despite the functional importance of these interactions. Comparison of rat-free and rat-invaded offshore islands in New Zealand revealed that predation of seabirds by introduced rats reduced forest soil fertility by disrupting sea-to-land nutrient transport by seabirds, and that fertility reduction in turn led to wide-ranging cascading effects on belowground organisms and the ecosystem processes they drive. Our data further suggest that some effects on the belowground food web were attributable to changes in aboveground plant nutrients and biomass, which were themselves related to reduced soil disturbance and fertility on invaded islands. These results demonstrate that, by disrupting across-ecosystem nutrient subsidies, predators can indirectly induce strong shifts in both above- and below-ground biota via multiple pathways, and in doing so, act as major ecosystem drivers.     

Competitive dominance mediates the effects of topography on plant richness in a mountain grassland

Small-scale landforms influence plant species richness, but their mechanisms and effects in semi-natural dry grasslands have been poorly investigated. In this study we compared vascular plant richness, species composition, plant traits, soil properties and biomass nutrient content of convex (hillocks) and concave (hollows) karst landforms in a mountain pasture of the Central Apennines (Italy), at a small spatial scale (1 m² plots). We found hillocks had significantly higher species richness than hollows. On hillocks, smaller Specific Leaf Area and Lateral Width, together with greater allocation of resources to Below-Ground Organs, indicated lower water availability, whereas hollows had deeper (thus moister), more acidic and more fertile soils, with aboveground plant biomass displaying higher nutrient levels. Partial correlation and regression tree models suggested that fine-scale richness patterns were not directly determined by abiotic properties, but were rather the result of competition levels associated with the cover of Agrostis capillaris (=A. tenuis) – a calcifuge and drought-sensitive grass able to achieve dominance only in hollows. The higher functional convergence exhibited by hollows suggests that A. capillaris is a strong competitor both above- and below-ground, mediating the effects of topography by imposing a biotic filter. On hillocks, competition is released by lower levels of available soil water in summer and higher soil pH, resulting in higher species richness and a more functionally divergent assemblage.     

氮素添加对青藏高原高寒草甸植物群落物种丰富度及其与地上生产力关系的影响

植物种群对有限资源的竞争是决定植物群落物种组成、多样性和生产力等群落结构和功能的主要因素。该文以青藏高原高寒草甸为研究对象, 研究了短期内不同水平的氮素添加对高寒草甸植物群落的影响。结果表明: 1)氮素添加提高了土壤中NO₃^--N等可利用资源的含量, 增加了植物群落植被的盖度, 减小了植被的透光率, 随着施氮量的增加, 群落中物种丰富度显著降低(p < 0.001); 2)氮素添加显著改变了植物群落的地上生产力(p < 0.05), 随着施氮量的增加, 地上生产力呈先增加后降低的变化趋势, 各功能群中禾草生物量显著增加, 而杂类草和豆科植物生物量随施氮量的增加逐渐减少; 3)物种多样性与植被透光率呈线性正相关(p < 0.05); 地上生产力与土壤NO₃^--N含量呈线性正相关(p < 0.05); 物种丰富度与地上生产力之间呈负相关关系。这说明短期内氮素添加通过改变土壤中NO₃^--N等可利用资源的含量而对植物群落物种组成和地上生产力产生影响。     

黄土高原半干旱区退化草地恢复与利用过程研究

草地退化表现为土壤和植被遭到彻底破坏,草地演替过程受到强烈抑制.实验采用长期(30年)封禁措施,定位监测退化草地从次生半裸地演变为近似原生植被(进展演替)的变化过程.结果表明,随着封禁时间的变化,退化草地恢复演替经历了4个阶段,群落盖度、植株密度、物种丰富度和多样性指数、地上生物量和地下生物量在草地群落恢复过程中逐渐增加,其特征变化出现的峰值均在封禁第20年(地下生物量峰值在第15年),其中地上生物量最高达520.5 g/m2;直到封禁的第20~25年,以本氏针茅为建群种的草原群落衰败退化现象明显,而大针茅种群密度剧增;在封禁的第26年以上以大针茅为优势的群落生长较为稳定,从目前群落演替进程看,大针茅有替代本氏针茅的趋势.另外,在草原沟道两侧以斑块状聚集分布有中旱生灌木,群落的演替进入了一个新的阶段.随着封禁时间的延续,退化草地从自然封禁恢复的0~26年,通过侵入-竞争-扩散-定居的几个演替阶段,目前形成以大针茅为建群种相对稳定的"亚顶级".虽然草地生物量有一定下降,但草地质量提高,物种多样性丰富,促进草地的进展演替.草地植物群落主要由禾本科、豆科和菊科组成;多年生植物、C3和旱生物种可以作为草地演替过程和植被恢复的指示物种.长期封育对草地物种更新和生态系统稳定性有负面影响,因此,合理的封育时间是草地生态恢复中非常重要的一个因素.本研究提出,在黄土区退化草地封育10~15年后可以开始进行合理的利用,例如通过两年一次刈割和轻度放牧(2只羊/hm2).本研究可为干旱区、半干旱区相似的退化草地恢复提供理论依据.     

Nitrogen deposition causes widespread loss of species richness in British habitats

We use national scale data to test the hypothesis that nitrogen (N) deposition is strongly negatively correlated with plant species richness in a wide range of ecosystem types. Vegetation plots from a national ecological surveillance programme were drawn from heathland, acid, calcareous and mesotrophic grassland habitats. Mean species number and mean plant traits were calculated for each plot and related to atmospheric N deposition. There was a significant reduction in species richness with N deposition in acid grassland and heathland even after fitting covarying factors. In acid grassland and heathland, evidence from trait changes suggested that acidification rather than increased fertility was responsible for species loss. In contrast, calcareous grassland showed evidence of eutrophication in response to increasing N deposition. Loss of species richness from chronic N deposition is apparent in infertile grasslands and heathland. Mechanisms associated with loss of species richness differ between habitats so mitigation of N deposition should be targeted to habitat type.     

Constraints on colonization and species richness along a grassland productivity gradient: the role of propagule availability

An important goal in ecology is to discern under what habitat conditions community structure is primarily regulated by local ecological interactions and under what conditions community structure is more regulated by the pool of available colonists. I conducted a seed addition experiment in successional grassland to evaluate the relative significance of neighbourhood biotic interactions and propagule availability in regulating plant colonization and species richness along a natural gradient of grassland productivity. In undisturbed field plots, seed additions of 34 species led to an increase in species richness in locations of low productivity, an effect that declined in magnitude as productivity increased. In disturbed plots, seed additions led to a relatively constant increase in species richness at all levels of productivity. The results support the hypothesis that the role of propagule availability in regulating colonization dynamics and species richness declines in significance relative to local-scale competitive interactions as habitat productivity increases.