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.     

Fertilization effects on species density and primary productivity in herbaceous plant communities

Fertilization experiments in plant communities are often interpreted in the context of a hump-shaped relationship between species richness and productivity. We analyze results of fertilization experiments from seven terrestrial plant communities representing a productivity gradient (arctic and alpine tundra, two old-field habitats, desert, short- and tall-grass prairie) to determine if the response of species richness to experimentally increased productivity is consistent with the hump-shaped curve. In this analysis, we compared ratios of the mean response in nitrogen-fertilized plots to the mean in control plots for aboveground net primary productivity (ANPP) and species density ( D ; number of species per plot of fixed unit area). In general, ANPP increased and plant species density decreased following nitrogen addition, although considerable variation characterized the magnitude of response. We also analyzed a subset of the data limited to the longest running studies at each site (≥4 yr), and found that adding 9 to 13 g N m⁻² yr⁻¹ (the consistent amount used at all sites) increased ANPP in all communities by approximately 50% over control levels and reduced species density by approximately 30%. The magnitude of response of ANPP and species density to fertilization was independent of initial community productivity. There was as much variation in the magnitude of response among communities within sites as among sites, suggesting community-specific mechanisms of response. Based on these results, we argue that even long-term fertilization experiments are not good predictors of the relationship between species richness and productivity because they are relatively small-scale perturbations whereas the pattern of species richness over natural productivity gradients is influenced by long-term ecological and evolutionary processes.     

Resource heterogeneity, soil fertility, and species diversity: effects of clonal species on plant communities

Spatial heterogeneity in soil resources is widely thought to promote plant species coexistence, and this mechanism figures prominently in resource-ratio models of competition. However, most experimental studies have found that nutrient enhancements depress diversity regardless of whether nutrients are uniformly or heterogeneously applied. This mismatch between theory and empirical pattern is potentially due to an interaction between plant size and the scale of resource heterogeneity. Clonal plants that spread vegetatively via rhizomes or stolons can grow large and may integrate across resource patches, thus reducing the positive effect of small-scale resource heterogeneity on plant species richness. Many rhizomatous clonal species respond strongly to increased soil fertility, and they have been hypothesized to drive the descending arm of the hump-shaped productivity-diversity relationship in grasslands. We tested whether clonals reduce species richness in a grassland community by manipulating nutrient heterogeneity, soil fertility, and the presence of rhizomatous clonal species in a 6-year field experiment. We found strong and consistent negative effects of clonals on species richness. These effects were greatest at high fertility and when soil resources were applied at a scale at which rhizomatous clonals could integrate across resource patches. Thus, we find support for the hypothesis that plant size and resource heterogeneity interact to determine species diversity.     

Patterns of species density and productivity at different spatial scales in herbaceous plant communities

A major challenge in evaluating patterns of species richness and productivity involves acquiring data to examine these relationships empirically across a range of ecologically significant spatial scales. In this paper, we use data from herb‐dominated plant communities at six Long‐Term Ecological Research (LTER) sites to examine how the relationship between plant species density and above‐ground net primary productivity (ANPP) differs when the spatial scale of analysis is changed. We quantified this relationship at different spatial scales in which we varied the focus and extent of analysis: (1) among fields within communities, (2) among fields within biomes or biogeographic regions, and (3) among communities within biomes or biogeographic regions. We used species density (D=number of species per m²) as our measure of diversity to have a comparable index across all sites and scales. Although we expected unimodal relationships at all spatial scales, we found that spatial scale influenced the form of the relationship. At the scale of fields within different grassland communities, we detected a significant relationship at only one site (Minnesota old‐fields), and it was negative linear. When we expanded the extent of analyses to biogeographic regions (grasslands or North America), we found significant unimodal relationships in both cases. However, when we combined data to examine patterns among community types within different biogeographic regions (grassland, alpine tundra, arctic tundra, or North America), we did not detect significant relationships between species density and ANPP for any region. The results of our analyses demonstrate that the spatial scale of analysis – how data are aggregated and patterns examined – can influence the form of the relationship between species density and productivity. It also demonstrates the need for data sets from a broad spectrum of sites sampled over a range of scales for examining challenging and controversial ecological hypotheses.     

Species evenness and productivity in experimental plant communities

In nature, plant biomass is not evenly distributed across species, and naturally uncommon species may differ from common species in the probability of loss from the community. Understanding relationships between evenness and productivity is therefore critical to understanding changes in ecosystem functioning as species are lost from communities. We examined data from a large multi-site grassland experiment (BIODEPTH) for relationships between evenness of species composition (proportional abundance of biomass) and total biomass of communities. For plots which started with the same and even species composition, but which diverged in evenness over time, those with lower evenness had a significantly greater biomass. The relationship between evenness and biomass across all plots was also negative. However, for communities where the most common species represented one of the three largest species in monoculture at that site (inclusion of a large dominant species), the relationship was neutral. Path analyses indicated that three paths contributed to this negative relationship. First, higher species richness decreased evenness, but increased biomass (primarily through an increase in maximum plant size). Contrary to predictions, maximum plant size had either no effect on evenness, or a positive effect (in year 3 plots with a large dominant species), thereby reducing this relationship. In year 2, large variation among species in plant size (as measured in monoculture) both decreased evenness and increased biomass, thus increasing the strength of the negative relationship between evenness and biomass. However, the former effect was only found in plots with a large dominant species, the latter only in plots without a large dominant species. When species richness, maximum plant size, and variation in size were accounted for, in year 2 evenness positively affected biomass in plots that included a large dominant species. Our results are consistent with the view that naturally uncommon species may be unaffected by (or even benefit from) the presence of a large naturally common species, and that uncommon plants may have little ability to increase productivity in the absence of such a species. We conclude that the observed negative relationship between evenness and biomass resulted from multiple direct and indirect effects, the relative strength of which depended in part on the presence of large dominant species.     

物种组成、丰富度、播种密度和土壤养分对群落补偿效应的影响

为了明确物种组成、丰富度、播种密度和土壤养分对群落补偿效应的影响,以及群落补偿效应随时间的变化趋势,本文以青藏高原高寒草地常见物种垂穗披碱草(Elymus nutans)、中华羊茅(Festuca sinensis)和羊茅(F.ovina)为材料,通过建植不同物种组合的单、混播草地,连续5年对不同群落的生产力进行了测定,结果表明:物种组成显著影响群落的补偿效应;在施肥情况下,物种丰富群落有较高的补偿效应;实验初期,随着播种密度的增加,群落补偿效应有降低趋势,但随着生长年限的增加,密度对补偿效应的影响变得不显著;施肥显著提高了群落的补偿效应;随着生长年限的延长,群落补偿效应有升高趋势。     

Effect of plant species richness on invasibility of experimental plant communities

Invasion of unsown species to artificially created assemblages of grassland species was investigated in a 3-year field experiment. In the experiment, assemblages varying in species richness (1, 2, 4, 8, and 16 species) and functional group richness (1–4, grasses, legumes, rosette forbs, and creeping forbs) were grown in control and fertilized plots, without any attempt to prevent the invasion of unsown species or to weed them. The relationship between species and functional group diversity and above-ground biomass was positive for sown species in all study years (2003, 2004 and 2005). In the latter 2 years, weed invader biomass decreased significantly with increasing biomass of sown species and their functional group richness, but not with number of species. However, no suppressive effect of species or functional group richness beyond that by increased biomass of residents was found. In fact, slight but significant positive partial effect of species richness was found, suggesting that the negative effect of the same amount of resident biomass on invaders is stronger when the biomass is composed of fewer species. The negative relationship between the number of functional groups of residents and invader biomass suggested that better coverage of functional trait space could be a mechanism promoting the resistance to invasion. In Addition, species composition of invaders were significantly related to initial composition of sown residents.     

干扰对植物群落物种组成及多样性的影响

在介绍了干扰的概念及其性质和干扰对物种多样性影响的有关假说基础上,以森林干扰为主要对象,探讨了干扰对植物群落物种组成的影响,并从干扰类型、干扰强度和干扰频率等几个方面阐述了干扰对植物群落物种多样性的影响;另外,还分析了重要的小尺度干扰——林隙在该方面的影响。分析总结出干扰对植物群落的影响差异主要与干扰特征、植物群落特征及植物的生物学特性和受干扰地点的资源条件有关。最后,总结分析了国内外在该方面研究存在的问题,并对今后研究提出一些建议,为相关工作提供参考。     

Small differences in arrival time influence composition and productivity of plant communities

'Who comes first' is decisive for plant community assembly and ecosystem properties. Early arrival or faster initial development of a species leads to space occupancy both above and below ground and contributes to species success. However, regular disturbance (e.g. biomass removal) might permit later-arriving or slower-developing species to catch up. Here, artificial communities of grassland species belonging to the plant functional types (PFTs) herb, grass and legume were used to test the effect of stepwise arrival (sowing) of PFTs. Dramatic effects were found as a result of a 3 wk arrival difference on composition and above-ground biomass that persisted over four harvests and two seasons. Priority effects, such as unequal germination time (arrival), and thus differences in community age structure, had lasting effects on PFT biomass contribution and associated ecosystem functioning. These effects were robust against above-ground disturbance. Benefits of earlier root formation outweighed above-ground species interaction. Earlier space occupancy and bigger reserve pools are the likely causes. Natural populations commonly exhibit age diversity and asynchrony of development among taxa. In experiments, artificial synchrony of arrival (sowing) may thus induce assembly routes favouring faster-establishing taxa, with consequences for ecosystem functioning (e.g. productivity). Founder effects, such as those observed here, could be even greater in communities of slow-growing species or forests, given their longer generation time and minor disturbance.     

Spatial patterns and species performances in experimental plant communities

The roles of co-occurring herbivores that modify habitat structure and ecosystem processes have seldom been examined in manipulative experiments or explored in early successional communities. In a created marsh in southern California (USA), we tested the individual and combined effects of two epibenthic invertebrates on nutrient and biomass pools, community structure, and physical habitat features. We manipulated snail (Cerithidea californica) and crab (Pachygrapsus crassipes) presence in field enclosures planted with pickleweed (Salicornia virginica) at elevations matching the plant’s lower extent in an adjacent natural marsh. In the 4-month experiment, C. californica altered habitat structure by reducing sediment surface heterogeneity and shear strength (a measure of sediment stability) markedly throughout the enclosures. Both invertebrates had strong negative effects on a group of correlated sediment physicochemical characteristics, including nitrogen and organic matter concentrations and soil moisture. In addition, both invertebrates greatly reduced benthic chlorophyll a, a proxy for biomass of microphytobenthos. Compared to controls, macroalgal cover was up to sixfold lower with crabs present, while snails increased cover at low elevations of enclosures. Unexpectedly, macroalgal cover was eliminated with both species present, perhaps through P. crassipes consumption of larger thalli and C. californica reduction in cover of recruits. Neither species influenced the S. virginica canopy (quantified with an index of branch length and number); however, at the lower elevation of enclosures, the two species together negatively impacted the plant canopy. The two invertebrates’ modifications to our experimental marshes led to distinct suites of biotic and physicochemical features depending on their presence or co-occurrence, with the latter producing several unexpected results. We propose that the roles and interactions of habitat-modifying fauna deserve further attention, particularly in the context of efforts to conserve and restore the processes found in natural systems.     

Local adaptation of mycorrhizae communities changes plant community composition and increases aboveground productivity

Soil microbial communities can have an important role in the adaptation of plants to their local abiotic soil conditions and in mediating plant responses to environmental stress. This has been clearly demonstrated for individual plant species, but it is unknown how locally adapted microbes may affect plant communities. It is possible that the adaptation of microbial communities to local conditions can shape plant community composition. Additionally, it is possible that the effects of locally adapted microorganisms on individual plant species could be altered by co-occurring plant species. We tested these possibilities in plant community mesocosms with soils and mycorrhizal fungi (AMF) from three locations. We found that plant community biomass responded positively to local adaptation of AMF to soil conditions. Plant community composition also changed in response to local adaptation of AMF. Unexpectedly, the strongest benefits of locally adapted AMF went to early successional plant species that have the highest relative growth rates and the lowest responsiveness to the presence of AMF. Late successional plants that responded positively overall to the presence of AMF were often suppressed in communities with local AMF, perhaps because of strong competition from fast growing plant species. These results show that local adaptation of soil microbial communities can shape plant community composition, and the benefits that plants derive from locally adapted microorganisms can be reshaped by the competitive context in which these associations occur.     

Seasonal drought, soil fertility and the species density of tropical forest plant communities

Recent comparisons of plant species densities in tropical forest make it possible to evaluate factors that govern species richness. Contrary to earlier predictions, plant species densities are not greater on soils of relatively low fertility. In fact, the opposite trend is often observed, although the relationship between species densities and soil fertility is highly variable. However, tropical forest plant species densities consistently increase with rainfall. Species coexistence in wetter tropical forests may be facilitated by the absence of competition for moisture combined with year-round pest pressure and low understory light levels, which reduce growth rates and the potential for competition for other resources.     

Resource productivity and availability impacts for food-chain length

Ecologists have focused on food-chain length (FCL) for the past eight decades as an index of food-web structure. Here, I review the hypotheses determining FCL with a focus on resource productivity and availability effects on FCL. First, I introduce the mainstream hypotheses to explain FCL variations: productivity, ecosystem size, and disturbance. For the existing productivity and productive space hypotheses, I stress the importance of using resource availability to estimate the productivity effect on FCL. Using a FCL dataset from 15 ponds, I tested the resource stoichiometry effect on FCL for ponds with between carbon:nitrogen ratio of primary producers and FCL. Moreover, I provide a perspective for studying resource availability and stoichiometry effects on FCL and of the alternative hypotheses of productive-space and ecosystem size. Finally, I suggest the future directions of the FCL study: a resource subsidy and climate change effects on FCL and food-web structure.     

Temporal fluctuations and experimental effects in desert plant communities

In the Chihuahuan Desert of the southwestern United States we monitored responses of both winter and summer annual plant communities to natural environmental variation and to experimental removal of seed-eating rodents and ants for 13 years. Analyses of data on population densities of the species by principal component analysis (PCA) followed by repeated measures analysis of variance (rmANOVA) on PCA scores showed that: (1) composition of both winter and summer annual communities varied substantially from year to year, presumably in response to interannual climatic variation, and (2) community composition of winter annuals was also significantly affected by the experimental manipulations of seed-eating animals, but the composition of the summer annual community showed no significant response to these experimental treatments. Canonical discriminant analysis (CDA) was then applied to the data for winter annuals to more clearly identify the responses to the different classes of experimental manipulations. This analysis showed that removing rodents or ants or both taxa caused distinctive changes in species composition. There was a tendency for large-seeded species to increase on rodent removal plots and to decrease on ant removal plots, and for small-seeded species to change in the opposite direction. In the winter annual community there was a significant time x treatment interaction: certain combinations of species that responded differently to removal of granivores also showed opposite fluctuations in response to long-term climatic variation. The large year-to-year variation in the summer annual community was closely and positively correlated across all experimental treatments. The use of multivariate analysis in conjunction with long-term monitoring and experimental manipulation shows how biotic interactions interact with variation in abiotic conditions to affect community dynamics.     

Prediction of species composition of plant communities in a rural landscape based on species traits

The species composition of a community is a subset of the regional species pool, and predicting the species composition of a community from ecological traits of organisms is an important objective in ecology. If such a prediction can be made feasible, we could assess the risk of invasion of locally new species (alien species and genetically modified species) into natural communities. We developed and tested statistical models to predict a community’s species composition from ecological traits of the species pool. Various types of communities (forest, meadow, and weed communities) exist in a small area of traditional rural landscape in Japan, and have been maintained by human activities. These communities and the tracheophytes species pool in the 1-km² research area were considered. We used logistic regression and decision-tree analysis to construct predictive models of community species composition based on plant traits, using the presence or absence of species in a community as the dependent variable and ecological traits as independent variables. Plant traits were grouped by cluster analysis, and the average in each trait group was used for model building to avoid multiple collinearity. Statistical prediction models were significant in all communities. About 60–75% of species composition could be predicted from the measured plant traits in forest communities, but 33–56% in the meadow and weed communities. Our results showed the possibility of predicting the species composition of plant communities from the ecological traits of the plant species together with the information on local species pool.     

Impact of plant functional group, plant species, and sampling time on the composition of nirK-type denitrifier communities in soil

We studied the influence of eight nonleguminous grassland plant species belonging to two functional groups (grasses and forbs) on the composition of soil denitrifier communities in experimental microcosms over two consecutive years. Denitrifier community composition was analyzed by terminal restriction fragment length polymorphism (T-RFLP) of PCR-amplified nirK gene fragments coding for the copper-containing nitrite reductase. The impact of experimental factors (plant functional group, plant species, sampling time, and interactions between them) on the structure of soil denitrifier communities (i.e., T-RFLP patterns) was analyzed by canonical correspondence analysis. While the functional group of a plant did not affect nirK-type denitrifier communities, plant species identity did influence their composition. This effect changed with sampling time, indicating community changes due to seasonal conditions and a development of the plants in the microcosms. Differences in total soil nitrogen and carbon, soil pH, and root biomass were observed at the end of the experiment. However, statistical analysis revealed that the plants affected the nirK-type denitrifier community composition directly, e.g., through root exudates. Assignment of abundant T-RFs to cloned nirK sequences from the soil and subsequent phylogenetic analysis indicated a dominance of yet-unknown nirK genotypes and of genes related to nirK from denitrifiers of the order Rhizobiales. In conclusion, individual species of nonleguminous plants directly influenced the composition of denitrifier communities in soil, but environmental conditions had additional significant effects.     

Non-native grass invasion alters native plant composition in experimental communities

Invasions of non-native species are considered to have significant impacts on native species, but few studies have quantified the direct effects of invasions on native community structure and composition. Many studies on the effects of invasions fail to distinguish between (1) differential responses of native and non-native species to environmental conditions, and (2) direct impacts of invasions on native communities. In particular, invasions may alter community assembly following disturbance and prevent recolonization of native species. To determine if invasions directly impact native communities, we established 32 experimental plots (27.5 m²) and seeded them with 12 native species. Then, we added seed of a non-native invasive grass (Microstegium vimineum) to half of the plots and compared native plant community responses between control and invaded plots. Invasion reduced native biomass by 46, 64, and 58%, respectively, over three growing seasons. After the second year of the experiment, invaded plots had 43% lower species richness and 38% lower diversity as calculated from the Shannon index. Nonmetric multidimensional scaling ordination showed a significant divergence in composition between invaded and control plots. Further, there was a strong negative relationship between invader and native plant biomass, signifying that native plants are more strongly suppressed in densely invaded areas. Our results show that a non-native invasive plant inhibits native species establishment and growth following disturbance and that native species do not gain competitive dominance after multiple growing seasons. Thus, plant invaders can alter the structure of native plant communities and reduce the success of restoration efforts.     

高寒草甸不同草地类型功能群多样性及组成对植物群落生产力的影响

对不同类型草地功能群多样性和组成与植物群落生产力之间的关系进行了探讨。结果表明：(1)在矮嵩草(Kobresia humlis)草甸和金露梅(Potentilla froticosa)灌丛中,豆科植物的作用比较明显,而其他功能群植物的作用较弱。(2)在藏嵩草(Kobresia tibetica)沼泽化草甸和小嵩草(K．pygmaca)草甸中,虽然杂类草、C3植物和莎草科植物功能群的生产力占群落初级生产力的比例较大,但二者在统计上没有显著性差异,这表明群落生产力除受物种多样性的影响外,也受物种本身特征和环境资源的影响,更主要的是受到功能群内物种密度和均匀度的影响,即功能群组成比功能群多样性更能说明对生态系统过程的影响。(3)不同类型草地群落植物功能群盖度与群落初级生产力呈显著的线性相关。(4)不同类型草地群落生产力与功能群内物种数的变化均表现为单峰曲线关系,即功能群内物种数处于中间水平时,群落生产力最高。     

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.