三江源区不同建植年代人工草地群落演替与土壤养分变化

研究了了三源区不同建植期人工修复草地在不同演替阶段毒杂草[主要是甘肃马先蒿(Pedicularis kansuensis)]的入侵规律、数量特征,植物群落物种组成、生物苗和草地质最以及土壤养分、微生物活性的变化规律.结果表明,不同建植期人工修复草地植物群落的种类组成、植物功能群组成和群落数量特征存在显著差异.随着演替时间的推移,人工草地群落盖度、高度、物种数、生物最和多样性指数均表现出"V"字型变化规律,杂类草--甘肃马先蒿的数量特征变化尤为明显,在4 a的人工草地群落中开始局部入侵,在5～6 a的人工草地群落中大面积入侵,其入侵速度、入侵面积达到高峰期.土壤的含水量、容重、土壤中有机质、氮素和磷素在演替过程(7 a、9 a草地)中逐渐降低,到一定时期又逐步增加;随着演替的进行,不同建植期人工草地的土壤微牛物生物量碳和酶活性均呈"V"字型,变化.对于退化生态系统的恢复首先是植被恢复,其次是土壤肥力的恢复.土壤有机质等养分的积累、微生物活性的改善不仅能使土壤-植物复合系统的功能得以恢复,同时也能促进物种多样性的形成,有利于人工草地群落稳定性的提高.在试验区尽管植被恢复演替进行得比较缓慢,但从土壤发展的角度看,仍属进展演替.所以,在退化高寒草甸的恢复过程中,若降低和有效控制外界的干扰(如围栏封育),可为退化草地恢复提供繁殖体与土壤环境,实现人工草地逐步向恢复(正向)演替进行.图3表6参34     

Prioritizing Conservation Effort through the Use of Biological Soil Crusts as Ecosystem Function Indicators in an Arid Region

Abstract: Conservation prioritization usually focuses on conservation of rare species or biodiversity, rather than ecological processes. This is partially due to a lack of informative indicators of ecosystem function. Biological soil crusts (BSCs) trap and retain soil and water resources in arid ecosystems and function as major carbon and nitrogen fixers; thus, they may be informative indicators of ecosystem function. We created spatial models of multiple indicators of the diversity and function of BSCs (species richness, evenness, functional diversity, functional redundancy, number of rare species, number of habitat specialists, nitrogen and carbon fixation indices, soil stabilization, and surface roughening) for the 800,000‐ha Grand Staircase‐Escalante National Monument (Utah, U.S.A.). We then combined the indicators into a single BSC function map and a single BSC biodiversity map (2 alternative types of conservation value) with an unweighted averaging procedure and a weighted procedure derived from validations performance. We also modeled potential degradation with data from a rangeland assessment survey. To determine which areas on the landscape were the highest conservation priorities, we overlaid the function‐ and diversity‐based conservation‐value layers on the potential degradation layer. Different methods for ascribing conservation‐value and conservation‐priority layers all yielded strikingly similar results (r= 0.89–0.99), which suggests that in this case biodiversity and function can be conserved simultaneously. We believe BSCs can be used as indicators of ecosystem function in concert with other indicators (such as plant‐community properties) and that such information can be used to prioritize conservation effort in drylands.     

植被恢复的生态效应研究进展

植被在水土保持、水源涵养及生态系统的固碳过程中起着重要的作用。植被恢复是指运用生态学原理,通过保护现有植被、封山育林或营造人工林、灌、草植被,修复或重建被毁坏或被破坏的森林和其他自然生态系统,恢复其生物多样性及其生态系统功能。目前,植被的自然及人工恢复是改善脆弱生态系统及退化生态系统生态环境现状最有效的措施。植被在恢复过程中对地上植被生态系统,物种多样性的恢复有着重要影响,同时通过凋落物及根系的输入,可以有效改善地下生态系统,增加土壤的养分含量、改善土壤的物理结构、增加土壤生物的生物量及活性。文章以地上及地下生态系统为出发点,综述了植被恢复过程中自然及人工恢复过程中不同的植被类型、不同的恢复时间下植物物种组成和多样性、土壤理化性质及土壤微生物群落的变化。植被的自然及人工恢复在一定程度上均能增加植物物种的多样性,随着恢复年限的增加物种的组成发生改变且多样性呈增加趋势,但一些特殊环境下不当的人工恢复可造成植被演替向退化方向发展,降低生物多样性。不同的植被类型由于其生长方式的不同对土壤理化性质和土壤微生物的影响存在差异,随着恢复年限的增长,土壤理化指标及微生物学指标呈现先增加而后趋于平稳的状态。针对已有的研究进展,提出在未来的研究过程中,一方面应该增加更多的对比研究,对不同环境下,不同的恢复物种,不同的恢复方式进行更深入地探讨;另外一方面应增加不同尺度的研究,现有的研究多集中在样地尺度,未来应在更大尺度上进行分析;再者,地上及地下生态系统之间的相互关系及影响机理一直是土壤学科研究的热点,植被恢复过程中应增加更多该方面的机理研究。     

Biotic contexts alter metal sequestration and AMF effects on plant growth in soils polluted with heavy metals

Investigating how arbuscular mycorrhizal fungi (AMF)-plant interactions vary with edaphic conditions provides an opportunity to test the context-dependency of interspecific interactions. The relationship between AMF and their host plants in the context of other soil microbes was studied along a gradient of heavy metal contamination originating at the site of zinc smelters that operated for a century. The site is currently under restoration. Native C3 grasses have reestablished, and C4 grasses native to the region but not the site were introduced. Interactions involving the native mycorrhizal fungi, non-mycorrhizal soil microbes, soil, one C3 grass (Deschampsia flexuosa), and one C4 grass (Sorghastrum nutans) were investigated using soils from the two extremes of the contamination gradient in a full factorial greenhouse experiment. After 12 weeks, plant biomass and root colonization by AMF and non-mycorrhizal microbes were measured. Plants from both species grew much larger in soil from low-contaminated (LC) origin than high-contaminated (HC) origin. For S. nutans, the addition of a non-AMF soil microbial wash of either origin increased the efficacy of AMF from LC soils but decreased the efficacy of AMF from HC soils in promoting plant growth. Furthermore, there was high mortality of S. nutans in HC soil, where plants with AMF from HC died sooner. For D. flexuosa, plant biomass did not vary with AMF source or the microbial wash treatment or their interaction. While AMF origin did not affect root colonization of D. flexuosa by AMF, the presence and origin of AMF did affect the number of non-mycorrhizal (NMF) morphotypes and NMF root colonization. Adding non-AMF soil biota reduced Zn concentrations in shoots of D. flexuosa. Thus the non-AMF biotic context affected heavy metal sequestration and associated NMF in D. flexuosa, and it interacted with AMF to affect plant biomass in S. nutans. Our results should be useful for improving our basic ecological understanding of the context-dependency of plant-soil interactions and are potentially important in restoration of heavy-metal-contaminated sites.     

Microbial diversity and ecological networks as indicators of environmental quality

Evaluating the quality of ecosystems in terms of biological patrimony and functioning is of critical importance in the actual context of intensified human activities. Microbial diversity is commonly used as a bioindicator of ecosystems functioning. However, there is a lack of sensitivity of microbial diversity indicators in the case of moderate and chronic environmental degradation, such as atmospheric deposition of pollutants, agricultural practices, diffuse pollution by wastewater and climate change. As a consequence, there is a need for alternative bioindicators of soils and water quality. Here, we discuss the interest of adopting a more integrative approach based on biotic interaction networks beyond the simple diversity indicators. We review how the various biotic interactions can be integrated in the various microbial networks such as trophic, mutualistic and co-occurrence networks. Then we discuss the efficiency of microbial networks and associated metrics to detect changes in microbial communities. We conclude that the connectance, the number of links and the average degree of co-occurrence networks could vary from 10 to 50% in response to minor perturbations when microbial diversity parameters remain stable. Finally, we analyze studies that aimed at linking microbial networks and activity to evaluate the potential of such networks for providing simple and operational indicators of ecosystem quality and functioning.     

不同森林恢复类型对土壤生物学特性的影响

比较研究了南方红壤侵蚀区4种主要森林恢复类型下土壤中有机质、微生物结构与功能以及酶活性.结果表明,不同的森林恢复类型导致了土壤生物学性状的明显差异.4种森林类型的土壤生物学性状均比长期干扰下CK的高.人工林土壤生物学性状相对比天然次生林土壤差.整合上述指标的土壤生物学肥力指数分别为:天然次生林(0.752)、油茶林(0.611)、杉木林(0.422)、湿地松林(0.439)、对照(0.124).在森林恢复初期,采用自然恢复有效地提高了土壤生物学肥力.导致天然次生林土壤生物学活性相对较高的主要因素是较高的凋落物产量和质量、较高的根系生物量、较丰富的植物种类组成、较优越的土壤生态条件和快速的植物生长.在反映土壤生物学活性的指标选择方面,培养基平均颜色变化率(AWCD)、培养基丰富度和培养基Shannon-Wiener多样性指数与反映土壤生物学活性的大部分指标具有较好的相关关系,是反映土壤生物学活性的较好指标.图3表6参68     

植物多样性对土壤微生物的影响

生物多样性强烈地影响生态系统的过程.生态系统过程的变化可导致生物多样性衰减并因此导致生态系统功能衰退.植物种丰度和植物功能多样性对土壤细菌群落的代谢活性和代谢多样性有成正比的影响.土壤细菌的代谢活性和代谢多样性随植物种数量的对数和植物功能组的数量而直线上升.其原因可能是由植被流入土壤的物质和能量的多样性和数量的增加,也可能是由土壤动物区系起作用的土壤微生境的多样性的增加造成的.由于植物多样性的丧失所引起的植物生物量的减少对分解者群落有强烈的影响微生物生物量将可能减少,因为在大多数陆地生态系统中,有机碳源限制着土壤微生物的活性.     

Interactive effects of plant species diversity and elevated CO2 on soil biota and nutrient cycling

Terrestrial ecosystems consist of mutually dependent producer and decomposer subsystems, but not much is known on how their interactions are modified by plant diversity and elevated atmospheric CO2 concentrations. Factorially manipulating grassland plant species diversity and atmospheric CO2 concentrations for five years, we tested whether high diversity or elevated CO2 sustain larger or more active soil communities, affect soil aggregation, water dynamics, or nutrient cycling, and whether plant diversity and elevated CO2 interact. Nitrogen (N) and phosphorus (P) pools, symbiotic N2 fixation, plant litter quality, soil moisture, soil physical structure, soil nematode, collembola and acari communities, soil microbial biomass and microflora community structure (phospholipid fatty acid [PLFA] profiles), soil enzyme activities, and rates of C fluxes to soils were measured. No increases in soil C fluxes or the biomass, number, or activity of soil organisms were detected at high plant diversity; soil H2O and aggregation remained unaltered. Elevated CO2 affected the ecosystem primarily by improving plant and soil water status by reducing leaf conductance, whereas changes in C cycling appeared to be of subordinate importance. Slowed-down soil drying cycles resulted in lower soil aggregation under elevated CO2. Collembola benefited from extra soil moisture under elevated CO2, whereas other faunal groups did not respond. Diversity effects and interactions with elevated CO2 may have been absent because soil responses were mainly driven by community-level processes such as rates of organic C input and water use; these drivers were not changed by plant diversity manipulations, possibly because our species diversity gradient did not extend below five species and because functional type composition remained unaltered. Our findings demonstrate that global change can affect soil aggregation, and we advocate that soil aggregation should be considered as a dynamic property that may respond to environmental changes and feed back on other ecosystem functions.     

Self-organized spatial patterns of vegetation in alpine grasslands

There is increased interest in vegetation spatial pattern as an indicator of transition shifts following catastrophes. Much, however, remains unknown about the mechanisms that underlie spatial pattern formations. In this study, we examined how the spatial heterogeneity of species distributions in the grasslands of the Central Pyrenees and Middle Atlas Mountains is associated with plant species diversity and the importance of self-organizing processes in the control of pattern formations. In the grasslands of the Central Pyrenees and Middle Atlas, the spatial heterogeneity of species distributions increased along a habitat degradation gradient defined by an increase in bare soil. In Central Pyrenees grasslands, however, the increase in heterogeneity was associated with self-organizing bare soil formations, rather than the self-organizing distribution of plant species. In Middle Atlas grasslands, the increased heterogeneity of species spatial distributions was a consequence of the self-organizing capacity of the composing species; the increase in bare soil was randomly distributed. In the more heavily grazed grasslands (Middle Atlas), but not in the more lightly grazed and better preserved ecosystem (Central Pyrenees), plant species richness and diversity declined significantly with an increase in grazing pressure because fewer species were able to colonize empty space. On the contrary, the colonization of bare soil by new species increased the diversity and spatial organization of new colonizing species in Central Pyrenees grassland.     

Successional changes in plant resistance and tolerance to herbivory

Despite considerable research on plant defenses, we know very little about how temporal changes in the environment may influence resistance and tolerance levels, or the costs and benefits of these defense strategies for long-lived plant species. We hypothesized that, in successional habitats, predictable environmental changes should favor strong plasticity in defense phenotypes and that the costs, benefits, and levels of tolerance and resistance will change with environmental context. Using a widely distributed, old-field perennial, late goldenrod (Solidago altissima), we conducted a field experiment to test these predictions. We planted goldenrod genets exhibiting varying levels of resistance and tolerance into three early-successional and three late-successional fields (approximately three and 15 years in age, respectively) and experimentally measured resistance and tolerance levels and their associated costs and selection coefficients. We found a significant effect of successional stage but no effect of genotype or stage-genotype interaction on defense levels. Genets planted in early-successional fields appeared to be more resistant and less tolerant to herbivory than those same genets planted in late-successional fields. There were significant trade-offs between resistance and tolerance in early-successional fields but not in late-successional fields. Each late-successional field exhibited a significant cost or selection gradient for resistance, but there was no general pattern of resistance costs or selection gradients specific to a successional stage class. In contrast, there was evidence of stage-specific costs of tolerance; late-successional fields exhibited significant costs of tolerance whereas early-successional fields did not. There was no evidence of direct selection for or against tolerance in either stage. Our results suggest that defense phenotypes might change in qualitative ways during succession. High resistance in early stages may be attributed to associational effects of the early-successional community, reducing the probability of damage, and despite a cost of tolerance in late stages, tolerance may be beneficial in mitigating the effects of both herbivory and environmental stresses (i.e., low light availability) that limit fitness in these fields. This study provides experimental evidence that succession can strongly influence defense phenotypes and promote temporal variability in relative resistance and tolerance levels.     

Ecological consequences of carbon substrate identity and diversity in a laboratory study

Plants return a wide range of carbon (C) substrates to the soil system. The decomposition rate of these substrates is determined by their chemical nature, yet few studies have examined the relative ecological role of specific substrates (i.e., substrate identity) or mixtures of substrates. Carbon substrate identity and diversity may alter soil chemistry and soil community composition, resulting in changes in belowground ecosystem functions such as decomposition and nutrient transfer, creating feedbacks that may affect plant growth and the aboveground community. A laboratory experiment was set up in which eight C substrates of varying chemical complexity were added to a base soil singly, in pairs, fours, or with all eight together every four days over a 92-day period. After 92 days these soils were analyzed for changes in chemistry, microbial community structure, and components of ecosystem functioning. The identity of the added C substrates significantly affected soil chemistry, microbial basal and substrate-induced respiration, and soil microbial community structure measured by either the catabolic response profile (CRP) technique or phospholipid fatty acid composition. These belowground changes strongly affected the ability of the soil microflora to decompose cellulose paper, probably because of differential effects of the C substrates on soil energy supplies and enzyme activities. The addition of C substrates to soils also reduced plant growth compared to the unamended control soil, but less so in soils amended with a tannin than those amended with other substrates. Carbon substrate diversity effects saturated at low diversity levels, tended to have neutral or negative effects on ecosystem functions, and depended strongly on which C substrates were added. It increased CRP compound use but had little effect on other measures of the soil microbial community. Overall, results showed that the chemical nature of C substrates added to soil, and sometimes their diversity, can affect the soil microbial community and soil chemistry, which subsequently affect other ecosystem processes such as decomposition and plant growth. The identity and diversity of substrates that plants add to soil may therefore have important consequences for both above- and belowground ecosystem functions.     

Decomposers (Lumbricidae, Collembola) affect plant performance in model grasslands of different diversity

Decomposer invertebrates influence soil structure and nutrient mineralization as well as the activity and composition of the microbial community in soil and therefore likely affect plant performance and plant competition. We established model grassland communities in a greenhouse to study the interrelationship between two different functional groups of decomposer invertebrates, Lumbricidae and Collembola, and their effect on plant performance and plant nitrogen uptake in a plant diversity gradient. Common plant species of Central European Arrhenatherion grasslands were transplanted into microcosms with numbers of plant species varying from one to eight and plant functional groups varying from one to four. Separate and combined treatments with earthworms and collembolans were set up. Microcosms contained 15N labeled litter to track N fluxes into plant shoots. Presence of decomposers strongly increased total plant and plant shoot biomass. Root biomass decreased in the presence of collembolans and even more in the presence of earthworms. However, it increased when both animal groups were present. Also, presence of decomposers increased total N concentration and 15N enrichment of grasses, legumes, and small herbs. Small herbs were at a maximum in the combined treatment with earthworms and collembolans. The impact of earthworms and collembolans on plant performance strongly varied with plant functional group identity and plant species diversity and was modified when both decomposers were present. Both decomposer groups generally increased aboveground plant productivity through effects on litter decomposition and nutrient mineralization leading to an increased plant nutrient acquisition. The non-uniform effects of earthworms and collembolans suggest that functional diversity of soil decomposer animals matters and that the interactions between soil animal functional groups affect the structure of plant communities.     

Cascading effects of long-term land-use changes on plant traits and ecosystem functioning

There is much concern that the functioning of ecosystems will be affected by human-induced changes in biodiversity, of which land-use change is the most important driver. However, changes in biodiversity may be only one of many pathways through which land use alters ecosystem functioning, and its importance relative to other pathways remains unclear. In particular, although biodiversity-ecosystem function research has focused primarily on grasslands, the increases in agricultural inputs (e.g., fertilization, irrigation) and grazing pressure that drive change in grasslands worldwide have been largely ignored. Here we show that long-term (27-year) manipulations of soil resource availability and sheep grazing intensity caused marked, consistent shifts in grassland plant functional composition and diversity, with cascading (i.e., causal chains of) direct, indirect, and interactive effects on multiple ecosystem functions. Resource availability exerted dominant control over above-ground net primary production (ANPP), both directly and indirectly via shifts in plant functional composition. Importantly, the effects of plant functional diversity and grazing intensity on ANPP shifted from negative to positive as agricultural inputs increased, providing strong evidence that soil resource availability modulates the impacts of plant diversity and herbivory on primary production. These changes in turn altered litter decomposition and, ultimately, soil carbon sequestration, highlighting the relevance of ANPP as a key integrator of ecosystem functioning. Our study reveals how human alterations of bottom-up (resources) and top-down (herbivory) forces together interact to control the functioning of grazing systems, the most extensive land use on Earth.     

Sinks for nitrogen inputs in terrestrial ecosystems: a meta-analysis of 15N tracer field studies

Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (< 1 week after 15N tracer application), total ecosystem 15N recovery was negatively correlated with fine-root and soil 15N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3-18 months after 15N tracer application), total ecosystem 15N retention was negatively correlated with foliar natural-abundance 15N but was positively correlated with mineral soil C and N concentration and C:N, showing that plant and soil natural-abundance 15N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem 15N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for 15N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N x ha(-1) x yr(-1) above which most ecosystems showed net losses of applied 15N tracer in response to N fertilizer addition.     

Competition-colonization trade-offs and disturbance effects at multiple scales

The competition-colonization trade-off has long been a mechanism explaining patterns of species coexistence and diversity in nonequilibrium systems. It forms one explanation of the intermediate disturbance hypothesis (IDH) for local communities--specifically that diversity should be maximized at intermediate disturbance frequencies, yet only a fraction of empirical studies support IDH predictions. Similarly, this trade-off is also a powerful explanation of coexistence at larger spatial scales. I show, with a microbial experimental system, that the diversity-disturbance relationship is dependent on the relative distribution of species along this trade-off. Here I show that, when species are skewed toward late-successional habits, local diversity declines with disturbance. Yet, despite this trait skew, diversity at scales larger than the patch appears insensitive to the trade-off distribution. Intermediate disturbance frequencies produce the greatest diversity in patch successional stage, thus benefiting the maximum number of species at larger scales.     

Janzen-Connell effects are widespread and strong enough to maintain diversity in grasslands

Crop rotation schemes are believed to work by preventing specialist soil-borne pests from depressing the future yields of similar crops. In ecology, such negative plant-soil feedbacks may be viewed as a type of Janzen-Connell effect, which promotes species coexistence and diversity by preventing the same species from repeatedly occupying a particular site. In a controlled greenhouse experiment with 24 plant species and using soils from established field monocultures, we reveal community-wide soil-based Janzen-Connell effects between the three major functional groups of plants in temperate European grasslands. The effects are much stronger and more prevalent if plants are grown in interspecific competition. Using several soil treatments (gamma irradiation, activated carbon, fungicide, fertilizer) we show that the mechanism of the negative feedback is the buildup of soil pathogens which reduce the competitive ability of nearly all species when grown on soils they have formerly occupied. We further show that the magnitude of the change in competitive outcome is sufficient to stabilize observed fitness differences between functional groups in reasonably large communities. The generality and strength of this negative feedback suggests that Janzen-Connell effects have been underestimated as drivers of plant diversity in temperate ecosystems.     

Plant-soil feedback: testing the generality with the same grasses in serpentine and prairie soils

Plants can alter soil properties in ways that feed back to affect plant performance. The extent that plant-soil feedback affects co-occurring plant species differentially will determine its impact on plant community structure. Whether feedback operates consistently across similar plant communities is little studied. Here, the same grasses from two eastern U.S. serpentine grasslands and two midwestern tallgrass prairie remnants were examined for plant-soil feedback in parallel greenhouse experiments. Native soils were homogenized and cultured (trained) for a year with each of the four grasses. Feedback was evaluated by examining biomass variation in a second generation of (tester) plants grown in the trained soils. Biomass was lower in soils trained by conspecifics compared to soils trained by heterospecifics in seven of 15 possible cases; biomass was greater in conspecific soils in one other. Sorghastrum nutans exhibited lower biomass in conspecific soils for all four grasslands, so feedback may be characteristic of this species. Three cases from the Hayden prairie site were explained by trainer species having similar effects across all tester species so the relative performance of the different species was little affected; plants were generally larger in soils trained by Andropogon gerardii and smaller in soils trained by S. nutans. Differences among sites in the incidence of feedback were independent of serpentine or prairie soils. To explore the causes of the feedback, several soil factors were measured as a function of trainer species: nutrients and pH, arbuscular mycorrhizal (AM) spore communities, root colonization by AM fungi and putative pathogens, and functional diversity in bacterial communities as indicated by carbon substrate utilization. Only variation in nutrients was consistent with any patterns of feedback, and this could explain the greater biomass in soils trained by A. gerardii at Hayden. Feedback at Nottingham (one of the serpentine sites) differed, most notably for A. gerardii, from that of similar past studies that used different experimental protocols. To understand the consequences of feedback for plant community structure, it is important to consider how multiple species respond to the same plant-induced soil variation as well as differences in the feedback detected between greenhouse and field settings.     

Direct and indirect control of grassland community structure by litter, resources, and biomass

Multiple factors linked through complex networks of interaction including fertilization, aboveground biomass, and litter control the diversity of plant communities. The challenge of explaining plant diversity is to determine not only how each individual mechanism directly influences diversity, but how those mechanisms indirectly influence diversity through interactions with other mechanisms. This approach is well established in the study of plant species richness, but surprisingly little effort has been dedicated toward understanding the controls of community evenness, despite the recognition that this aspect of diversity can influence a variety of critical ecosystem functions. Similarly, studies of diversity have predominantly focused on the influence of shoot, rather than root, biomass, despite the fact that the majority of plant biomass is belowground in many natural communities. In this study, I examine the roles of belowground biomass, live aboveground biomass, litter, and light availability in controlling the species richness and evenness of a rough fescue grassland community using structural equation modeling. Litter was the primary mechanism structuring grassland diversity, with both richness and evenness declining with increasing litter cover. There were few relationships between shoot biomass, shading, and diversity, and more importantly, no relationship between root biomass and diversity. The lack of relationship between root biomass and species richness and evenness suggests that, even though root competition in grasslands is intense, belowground interactions may not play an important role in structuring community diversity or composition.     

Soil microbes drive the classic plant diversity-productivity pattern

Ecosystem productivity commonly increases asymptotically with plant species diversity, and determining the mechanisms responsible for this well-known pattern is essential to predict potential changes in ecosystem productivity with ongoing species loss. Previous studies attributed the asymptotic diversity-productivity pattern to plant competition and differential resource use (e.g., niche complementarity). Using an analytical model and a series of experiments, we demonstrate theoretically and empirically that host-specific soil microbes can be major determinants of the diversity-productivity relationship in grasslands. In the presence of soil microbes, plant disease decreased with increasing diversity, and productivity increased nearly 500%, primarily because of the strong effect of density-dependent disease on productivity at low diversity. Correspondingly, disease was higher in plants grown in conspecific-trained soils than heterospecific-trained soils (demonstrating host-specificity), and productivity increased and host-specific disease decreased with increasing community diversity, suggesting that disease was the primary cause of reduced productivity in species-poor treatments. In sterilized, microbe-free soils, the increase in productivity with increasing plant species number was markedly lower than the increase measured in the presence of soil microbes, suggesting that niche complementarity was a weaker determinant of the diversity-productivity relationship. Our results demonstrate that soil microbes play an integral role as determinants of the diversity-productivity relationship.