Trophic interactions and the relationship between species diversity and ecosystem stability

Several theoretical studies propose that biodiversity buffers ecosystem functioning against environmental fluctuations, but virtually all of these studies concern a single trophic level, the primary producers. Changes in biodiversity also affect ecosystem processes through trophic interactions. Therefore, it is important to understand how trophic interactions affect the relationship between biodiversity and the stability of ecosystem processes. Here we present two models to investigate this issue in ecosystems with two trophic levels. The first is an analytically tractable symmetrical plant-herbivore model under random environmental fluctuations, while the second is a mechanistic ecosystem model under periodic environmental fluctuations. Our analysis shows that when diversity affects net species interaction strength, species interactions--both competition among plants and plant-herbivore interactions--have a strong impact on the relationships between diversity and the temporal variability of total biomass of the various trophic levels. More intense plant competition leads to a stronger decrease or a lower increase in variability of total plant biomass, but plant-herbivore interactions always have a destabilizing effect on total plant biomass. Despite the complexity generated by trophic interactions, biodiversity should still act as biological insurance for ecosystem processes, except when mean trophic interaction strength increases strongly with diversity.     

Dominant species maintain ecosystem function with non-random species loss

Loss of species caused by widespread stressors, such as drought and fragmentation, is likely to be non‐random depending on species abundance in the community. We experimentally reduced the number of rare and uncommon plant species while independently reducing only the abundance of dominant grass species in intact, native grassland. This allowed us to simulate a non‐random pattern of species loss, based on species abundances, from communities shaped by natural ecological interactions and characterized by uneven species abundance distributions. Over two growing seasons, total above‐ground net primary productivity (ANPP) declined with reductions in abundance of the dominant species but was unaffected by a threefold decline in richness of less common species. In contrast, productivity of the remaining rare and uncommon species decreased with declining richness, in part due to loss of complementary interactions among these species. However, increased production of the dominant grasses offset the negative effects of species loss. We conclude that the dominant species, as controllers of ecosystem function, can provide short‐term resistance to reductions in ecosystem function when species loss is nonrandom. However, the concurrent loss of complementary interactions among rare and uncommon species, the most diverse component of communities, may contribute to additional species loss and portends erosion of ecosystem function in the long term.     

Two approaches towards the relationship between plant species diversity and ecosystem functioning

Abstract. The main question to be dealt with in the papers published in this Special Feature is to which extent plant species richness can be applied as a parameter in restoration projects to qualify the ecosystem's state. Before considering this problem, it should be recognized that this approach illuminates only one side of the coin; the other side is touched by the opposite question, asking which plant species are essential components of an ecosystem. These two approaches towards the relationship between species richness and ecosystem functioning are not mutually exclusive, but should not be confused either. In view of ecosystem functioning certain species may be considered redundant, while in view of evolutionary processes certain ecosystem processes may be considered redundant. Where do the two approaches meet and when should they be separated? This paper touches upon this question by referring to the dual hierarchy of ecological systems.     

Phenological complementarity, species diversity, and ecosystem function

Increasing species diversity frequently enhances ecosystem function. Phenological complementarity, the asynchrony of species resource use and growth, may explain how species diversity influences ecosystem function but remains largely untested. We used an early successional plant community containing species with a variety of phenologies to test whether increasing species diversity enhances ecosystem function by increasing phenological complementarity. Over a two-year period, we increased environmental heterogeneity within an abandoned field with variation in disturbance, soil nutrients, water, light availability, and disturbance in 160 permanent plots, and measured percent cover of each plant species three times in each growing season. We did not manipulate species composition directly, and thus diversity and complementarity in each plot were the result of pre-existing conditions and responses of individuals to experimental treatments. Species diversity was measured in two ways, as the total number of species per plot and as the evenness of species abundances. Phenological complementarity was measured as the negative logarithm of the variance ratio. We tested whether the number of plant species per plot, species evenness, and their phenological complementarity in the first year predicted total annual cover in the second year. Total annual cover increased only moderately with number of species and evenness, consistent with studies that randomize species composition among replicate plots. Any effect that species number or evenness had on total annual cover, however, was not due to phenological complementarity. Rather, diversity was unrelated to phenological complementarity. These results indicate that naturally occurring variation in species diversity had little effect on whether phenological complementarity can enhance ecosystem function.     

Chemical diversity – highlighting a species richness and ecosystem function disconnect

The lack of predictability in litter-mix studies may result from the low correlation between species number and the traits that drive the processes under observation. From the standpoint of litter-quality-dependent ecological processes, we propose that litter chemical qualities are functional traits and introduce a multivariate index of chemical diversity (CD_Q) based on Rao's quadratic entropy to describe the compositional heterogeneity of litter and foliar mixtures. Using published data from temperate and tropical forest systems to illustrate the relationship between species richness and chemical diversity, we show the variation of chemical diversity based on profiles of total nutrient concentrations (N, P, K, Ca and Mg) with species richness. We discuss how this behavior may explain the idiosyncratic responses exhibited in litter-mix experiments and how it may contribute to the observed dominance of species identity over species diversity. As a summary of resource heterogeneity relevant to detritivore and microbial processes, the chemical diversity index is potentially a better predictor of diversity effects on nutrient dynamics than species richness. Finally, we propose the use of infrared spectroscopy techniques for a rapid and more comprehensive determination of foliar and litter chemical composition to provide a more information-rich index.     

羌塘高寒草地物种多样性与生态系统多功能关系格局

传统的生物多样性-生态系统功能研究大多侧重于单一生态系统功能与物种多样性的关系,忽略了生态系统的重要价值在于其能够同时提供多种功能或服务,即生态系统的多功能性。基于藏北羌塘高寒草地样带调查数据,选取植被地上生物量、地下生物量、土壤全氮、硝态氮及铵态氮含量、土壤全磷含量、土壤有机碳储量等7个与植物生长、养分循环、土壤有机碳蓄积相关的参数来表征生态系统多功能性。采用上述参数转换为Z分数后的平均值计算多功能性指数(M)。分析了不同生物多样性指数与生态系统多功能指数的关系以及年降水量和年均温度对物种多样性和生态系统多功能性指数的影响。结果表明,物种丰富度指数与生态系统多功能性之间呈极显著的正相关关系,Shannon-wiener和Simpson物种多样性指数也与多功能性指数间呈显著的正相关,但多功能性指数与Pielou均匀度指数没有表现出明显的相关关系。物种丰富度与表征植物生长、养分循环以及土壤有机碳蓄积的生态系统功能指数间也均呈极显著的正相关关系。降水格局显著影响羌塘高原物种丰富度和生态系统多功能指数,二者均随年降雨量的增加而显著增加,但物种多样性指数并未与年降水量呈现显著相关关系。研究强调了群落物种丰富度即群落物种数量对维持生态系统多功能性的重要意义,这意味着由于人类活动导致的物种丧失可能会给藏北高寒草地生态系统多功能和生态服务带来更为严重的后果。就退化草地恢复或草地可持续管理而言,在藏北羌塘地区,本地植物种的物种丰富度恢复和维持应作为重要目标之一。     

物种多样性与生态系统功能的关系研究进展

物种的空前丧失促使人们越来越多地开始研究物种多样性与生态系统功能的关系,并探讨其潜在的作用机制.本文根据最新研究进展,归纳了微宇宙实验、"生态箱"实验、Cedar Creek草地多样性实验和欧洲草地实验等代表性实验中关于物种多样性与生产力、稳定性、抗入侵性等生态系统功能的焦点问题,介绍了去除实验在多样性与生态系统功能研究中的应用.在此基础上,提出未来研究所面临的挑战,并对研究趋势进行了展望.主要挑战和趋势有:将小尺度上开展的实验研究扩展到较大的时空尺度上;综合考虑生物因素和非生物因素对多样性变化、生态系统功能的交互作用;营养级之间的相互作用、物种共存机制对物种多样性与生态系统功能关系的影响.     

Resource availability dominates and alters the relationship between species diversity and ecosystem productivity in experimental plant communities

Experimental evidence that plant species diversity has positive effects on biomass production appears to conflict with correlations of species diversity and standing biomass in natural communities. This may be due to the confounding effects of a third variable, resource availability, which has strong control over both diversity and productivity in natural systems and may conceal any positive effects of diversity on productivity. To test this hypothesis, I independently manipulated resource availability (soil fertility) and sown species diversity in a field experiment and measured their individual and interactive effects on productivity. Although fertility was a far stronger predictor of productivity than diversity, the effect of diversity on productivity significantly increased with fertility. Relative yield analyses indicated that plant mixtures of high fertility treatments significantly "overyielded," or were more productive than expected based on monoculture yields of component species. In contrast, plant mixtures of low fertility treatments had significantly lower-than-expected yields. The effect of diversity on productivity was also driven by sampling effects, where more species-rich mixtures were more likely to include particularly productive species. Unexpectedly, the strength of sampling effects was largely insensitive to fertility, although the particular species most responsible for sampling effects did change with fertility. These results suggest that positive effects of species diversity on ecosystem productivity in natural systems are likely to be masked by variation in environmental factors among habitats.     

物种多样性与生态系统功能时间变异性的关系研究进展

近年来物种多样性的急剧丧失使得物种多样性与生态系统功能的时间变异性的关系及其机制问题的研究成为了生态学研究的一个热点。综述了物种多样性与群落集合性质变异性以及种群性质变异性的关系及其机制的最新研究成果：1、理论上探讨造成物种多样性与群落集合性质变异性负相关关系的机制包括：抽样效应、资源利用分化假说、统计平均效应、保险假说、种群变异性的均匀度效应等;但实验研究对理论预期的支持并不是普遍的;2．多样性与种群变异性之间的关系主要依赖于均值-方差尺度系数Z;理论上大部分自然群落是种群变异性应该随着多样性的增加而增加;但有研究表明：在变动环境中多样性对单个组分物种的种群水平有稳定性作用;而经验研究并不能得出多样性对种群变异性效应的清晰模式。讨论了目前的理论和实验研究中存在的和今后研究中需要认真思考的问题。     

Successional diversity and forest ecosystem function

Forest inventory data was used to examine the relationship between successional diversity and forest ecosytem function. The inventory data show that stands composed of early successional species are more productive than stands composed of late successional species, whereas stands composed of late successsional species have lower turnover than stands composed of early successional species. Taken alone, these results would suggest that forests should be managed in a way that favors the most productive early successional species or longest-lived late successional species, depending on whether the goal is to maximize productivity or maximize carbon storage. However, the inventory data also show that stands with low successional diversity fix and store less carbon than stands with high successional diversity. This result suggests that forests should be managed in such a way as to retain species diversity while also favoring species that maximize the ecosystem function of interest.     

Dominant ants can control assemblage species richness in a South African savanna

1. Competition is considered a key factor structuring many communities, and has been described as the 'hallmark' of ant ecology. Dominant species are thought to play a key role structuring local ant assemblages through competitive exclusion. 2. However, while there have been many studies demonstrating competitive exclusion and consequently reduced richness at baits, it is not clear whether such regulation of 'momentary' diversity at clumped food resources can scale up to the regulation of richness at the site or assemblage level. 3. In this study, ant assemblages were sampled in three different savanna habitats in South Africa using both baiting and pitfall trapping. 4. As has been found in previous studies, there was a unimodal relationship between dominant ants and species richness at baits, with high abundances of dominant ants regulating species richness through competitive exclusion. Analysis of pitfall samples revealed strong convergence in pattern, and results from null model co-occurrence analyses supported the findings. 5. The importance of competition in structuring local ant assemblages was, however, only apparent at one of the three savanna habitats suggesting that a full range of extreme environments is needed to produce the full unimodal relationship at the assemblage level. 6. Although the relative importance of competition varied with habitat type, the study demonstrated that in some habitats, dominant ants can control species richness at the assemblage level.     

Restoring plant species diversity and community composition in a ponderosa pine-bunchgrass ecosystem

Monitoring of ecological restoration treatments often focuses on changes in community structure and function. We suggest that long-term changes in community composition also need to be explicitly considered when evaluating the success of restoration treatments. In 1992, we initiated an experiment in a ponderosa pine-bunchgrass ecosystem to evaluate responses to restoration treatments: (a) thinning the overstory vegetation (‘thinning’), (b) thinning plus forest floor manipulation with periodic prescribed burning (‘composite’), and (c) untreated ‘control.’ Treatments were further stratified by forest patch type: presettlement tree clumps (trees that established prior to the onset of fire exclusion in 1876), patches of retained postsettlement trees, patches where all postsettlement trees were removed, and remnant grass openings. Species richness did not differ among treatments for 10 years, but was highest in the composite treatment in 11th and 12th year after initial treatment. Community composition diverged among treatments 5 years after initial treatment, and compositional changes were greatest in the composite treatment. Species richness and composition differed among patch types prior to treatment. Remnant grass patches were the most diverse and presettlement patches were the least diverse. Following treatment, species richness in the postsettlement removed and retained patches, gradually approached levels found in remnant grass patches. Compositional differences among patch types changed a little by 2005. Species richness at the 2 m² scale increased only where the overstory was thinned and the understory was burned. However, these changes may not be detectable for many years, and can vary temporally in response to events such as severe droughts. Nonnative species establishment may be reduced by scheduling longer burn intervals or by refraining from burning where fuel loads are not hazardous, though these options may hinder goals of increasing diversity. Restoring species diversity and community composition continues to be more difficult than restoring ecosystem structure and function.     

试论生态系统与生物体之间的全息关系

提出生态系统与生物体之间存在全息关系的观点,并认为生态系统与生物体都具有保护、支撑、运动、同化、呼吸、循环、排泄、繁殖和调控功能,生态系统的次生演替与生物体的再生修复过程存在着共同点。根据全息胚重演过程中的滞育性、可简化性,可以对生态演替的多方向、多途径问题作出新的解释。     

Soil microbes regulate ecosystem productivity and maintain species diversity

One of the major goals in ecology is to determine the mechanisms that drive the asymptotic increase in ecosystem productivity with plant species diversity. Niche complementarity, the current paradigm for the asymptotic diversity-productivity pattern, posits that the addition of species to a community increases productivity because each species specializes on different resources and thus can more thoroughly utilize the available resources. At higher diversity the increase in productivity decreases because resources become limiting, resulting in the classic asymptotic diversity-productivity pattern. An alternative but less tested explanation is that density-dependent disease from species-specific soil microbes drive the diversity-productivity relationship by increasing disease and thus decreasing productivity at low diversity. At higher diversity, productivity asymptotes because disease decreases with increasing diversity until it reaches a uniformly low level. Using a series of field experiments, we found that the classic asymptotic diversity-productivity pattern existed only when soil microbes were present. Soil microbes created the well-known pattern by depressing plant growth at low productivity though negative density dependent disease. In contrast, niche complementarity played only a weak role in explaining the diversity-productivity relationship because productivity remained high at low abundance in the absence of soil microbes. Based on our findings, the ongoing loss of species in natural ecosystems will likely increase per capita plant disease and lower ecosystem productivity. Furthermore, recent evidence suggests that negative density dependent disease maintains plant species diversity, and thus this single mechanism appears to link diversity maintenance to the diversity-productivity curve—two important ecological processes.Key words: density dependence, diversity-productivity, negative feedback, pathogens, species richness, soil microbesThe asymptotically saturating increase in ecosystem productivity with increasing diversity is a well know pattern in nature^1–4 (Fig. 1). The pattern has been used as an argument for the importance of species diversity,⁵ and understanding the mechanisms that drive the pattern is critical to determine the potential loss in productivity with ongoing and accelerating species loss in many ecosystems. The cause of the diversity-productivity pattern can be explained by either bottom-up control, such as plant resource competition, or top-down control from plant herbivores or pathogens. Most contemporary explanations for the pattern are centered on the bottom-up concept of niche-based resource competition, in which different species utilize different resources. The commonly accepted explanation, the niche complementarity hypothesis, states that the increase in species diversity increases productivity because each additional species uses a differ set of resources (e.g., nutrients) and thus more thoroughly utilizes whole-ecosystem resources.^3,4,6 At high diversity, however, the resource requirements of additional species overlap with existing ones and thus productivity no longer increases with diversity, resulting in the asymptotic diversity-productivity pattern (Fig. 1).Open in a separate windowFigure 1Theoretical relationship between species number and biomass. As diversity increases, total biomass increases asymptotically.Top-down control from plant enemies may also produce the asymptotic diversity-productivity pattern if the enemies are species-specific and have a strong negative density-dependent effect at low diversity. One general group of enemies is plant pathogens and parasites (bacterial, fungal, viral) that live in the soil and infect plant roots (hereafter referred to as soil pathogens). The specificity of soil pathogens has been shown in various studies and is now generally accepted.^1,7,8 The negative density dependent effect of plant pathogens at low diversity is likely because when diversity is low the relative abundance of each remaining species is high,^9–11 which leads to most individuals growing in close proximity of conspecifics and thus a greater probability of species-specific disease transmission. Unlike other plant enemies, such as foliar pathogens or insect and mammalian herbivores, which can be broadly dispersed, soil-borne pathogens may be a particularly effective driver of negative density dependent effects because they have low mobility and thus are more likely to infect nearby conspecifics, which causes increased disease at low diversity.^9–11 As diversity increases, the effect of soil-borne pathogens decreases because there is a lower likelihood of growing near a conspecific and there are lower concentrations of host-specific soil enemies.¹⁰ Consequently, soil-borne, species-specific disease may limit ecosystem productivity through top-down density-dependent regulation, even in the absence of niche-based explanations. Few studies, however, have considered the role of plant soil pathogens in driving the classic diversity-productivity relationship¹ (see also ref. ²) and, until now, no study has compared the two potential drivers simultaneously.¹We used a modeling approach to first demonstrate that both niche complementarity and species-specific soil pathogens can both theoretically drive the well-known diversity-productivity pattern.¹ We then used a series of complementary field experiments in grasslands in North America (Ontario, Canada and Minnesota, USA) to determine how plant disease and productivity change over a gradient of plant species richness in the presence and absence of soil microbes, and whether feedback between plants and their species-specific soil biota influenced the diversity-productivity pattern.¹ We first tested whether the asymptotic diversity-ecosystem productivity relationship arose in the presence of soil pathogens (a test of the negative density dependence hypothesis) or in the absence of soil pathogens (a test of the niche complementarity hypothesis). We then confirmed that soil biota were species specific and examined the decrease in plant disease and increase in productivity with increasing plant diversity.     

The relationship between nested subsets,habitat subdivision,and species diversity

Biotic assemblages are said to be nested when the species making up relatively species-poor biotas comprise subsets of the species present at richer sites. Because species number and site area are often correlated, previous studies have suggested that nestedness may be relevant to questions of how habitat subdivision affects species diversity, particularly with respect to the question of whether a single large, contiguous patch of habitat will generally contain more species than collections of smaller patches having the same total combined area. However, inferences from analyses of nestedness are complicated by (1) variability in degrees of nestedness measured in natural communities, (2) variability in species-area relationships, and (3) the fact that nestedness statistics do not account for the size of habitat patches, only in the degree of overlap among sites with different numbers of species. By comparing various indices of nestedness with a saturation index that more directly measures the effect of habitat subdivision, it is shown that the first two of these factors are not as important as the third. Whether a single large site or several smaller ones having the same total combined area maximizes species diversity is dependent on (1) overlap in species composition among sites and (2) the number of species per unit area in the different sites. Because nestedness indices do not account for species number at a site, they cannot accurately predict how habitat subdivision affects species diversity patterns. Still, nestedness analyses are important in that they indicate the degree to which rare species tend to be found in the largest, or the most species-rich, sites, patterns not revealed by the saturation index. Both types of analysis are important in order to obtain a more complete picture of how species richness and compositional patterns are influenced by habitat subdivision.     

Connections between species diversity and genetic diversity

Species diversity and genetic diversity remain the nearly exclusive domains of community ecology and population genetics, respectively, despite repeated recognition in the literature over the past 30 years of close parallels between these two levels of diversity. Species diversity within communities and genetic diversity within populations are hypothesized to co‐vary in space or time because of locality characteristics that influence the two levels of diversity via parallel processes, or because of direct effects of one level of diversity on the other via several different mechanisms. Here, we draw on a wide range of studies in ecology and evolution to examine the theoretical underpinnings of these hypotheses, review relevant empirical literature, and outline an agenda for future research. The plausibility of species diversity–genetic diversity relationships is supported by a variety of theoretical and empirical studies, and several recent studies provide direct, though preliminary support. Focusing on potential connections between species diversity and genetic diversity complements other approaches to synthesis at the ecology–evolution interface, and should contribute to conceptual unification of biodiversity research at the levels of genes and species.     

Links between tree species, symbiotic fungal diversity and ecosystem functioning in simplified tropical ecosystems

We studied the relationships among plant and arbuscular mycorrhizal (AM) fungal diversity, and their effects on ecosystem function, in a series of replicate tropical forestry plots in the La Selva Biological Station, Costa Rica. Forestry plots were 12 yr old and were either monocultures of three tree species, or polycultures of the tree species with two additional understory species. Relationships among the AM fungal spore community, host species, plant community diversity and ecosystem phosphorus-use efficiency (PUE) and net primary productivity (NPP) were assessed. Analysis of the relative abundance of AM fungal spores found that host tree species had a significant effect on the AM fungal community, as did host plant community diversity (monocultures vs polycultures). The Shannon diversity index of the AM fungal spore community differed significantly among the three host tree species, but was not significantly different between monoculture and polyculture plots. Over all the plots, significant positive relationships were found between AM fungal diversity and ecosystem NPP, and between AM fungal community evenness and PUE. Relative abundance of two of the dominant AM fungal species also showed significant correlations with NPP and PUE. We conclude that the AM fungal community composition in tropical forests is sensitive to host species, and provide evidence supporting the hypothesis that the diversity of AM fungi in tropical forests and ecosystem NPP covaries.