Seasonal changes in the relationship between plant species richness and community biomass in early succession

We analysed the relationship between plant species richness and productivity on first-year-old fields at two similar sites in central Europe. At both sites, a wide range of productivity levels was available resulting from different long-term fertilisation. In order to identify underlying mechanisms of the species richness–productivity relationship we included the seasonal dynamics and the number of individuals of each species in our analysis. We sampled 10 and 21 plots, respectively, at the two sites in May, June and July by harvesting all aboveground parts of vascular plants in 0.25 m² subplots. Species richness, number of individuals of each species and community biomass as a surrogate of productivity were recorded in each sample.At one site, the relationship between species richness and biomass was significantly positive in the May and June harvest. This relationship disappeared in the July harvest due to a reduction in species richness at high productivity levels. The relations between species richness and number of individuals and between number of individuals and biomass paralleled the species richness–productivity relation but the individual number–biomass relationship remained positive until the last harvest. Between-species differences in individual number–community biomass relationships and their seasonal dynamics revealed “interspecific competitive exclusion” even though the species richness–biomass relationships were not negative or hump-shaped. At the second site, species richness was not related to productivity or to number of individuals. Our study demonstrated the importance of temporal dynamics and regional processes in understanding species richness–productivity patterns.     

Competitive exclusion and the No‐Interaction model operate simultaneously in microcosm plant communities

Question: Is the relation between productivity and species richness due to an increase in plant size and hence a reduced plant density? Location: Glasshouse experiment. Methods: Productivity was manipulated with fertilizer and irrigation in a microcosm experiment. The ‘sampling effect’ was removed using rarefaction to a common density of individual plants per pot. Results: Fertility increased community biomass towards an asymptotic maximum, and reduced the light passing through plant canopies towards an asymptotic minimum. As biomass increased, so did species richness. However, this did not seem to be a direct effect of productivity on species richness, but rather one mediated by plant density, since: (1) the richness/ density relation was stronger than the richness/biomass one; (2) adding biomass to the richness/density regression did not increase its predictivity; (3) the richness/biomass relation was removed by rarefaction to 200 individuals per pot. It is therefore concluded that the richness/biomass relation observed was due to the sampling effect. Rarefaction to a small number of plants gave a quite different trend: lower richness at high biomass. This seems to be due to an increased number of subordinate species at high community biomass, and a more uneven distribution of abundance. Conclusion: The Competitive Exclusion and No‐Interaction hypotheses have been seen as alternatives. We suggest that they can operate simultaneously.     

Impact of Land-Use Intensity and Productivity on Bryophyte Diversity in Agricultural Grasslands

While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m²) and central Germany (6.4 species per 16 m²) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.     

Responses of subalpine dwarf‐shrub heath to irrigation and fertilization

Question: Our study aimed at testing to what extent water and/or nutrients affect community composition in sub‐alpine heath. We hypothesized that nutrient addition will have an overall positive effect on heath vegetation but water addition will have greater effects and will probably reinforce the effects of fertilization in drier habitat conditions. Location: Monte Rondinaio, northern Apennines, Italy (44°08’ N; 10°35’ E, ca. 1850‐1930 m a.s.l.). Methods: Nutrients and water were added during five growing seasons in two communities (HV community, moister; EV community, drier) and the biomass of all species was estimated non‐destructively by the point‐quadrat method. Results: Total above‐ground biomass increased in both communities with fertilization due to increased graminoid biomass and decreased moss biomass, but was unaffected by irrigation. In the HV community species richness decreased as an effect of fertilization while in the EV community species richness was raised by irrigation. Conclusions: Our study partly supported our hypothesis since water addition had a stronger effect in the drier community. However, this effect was not revealed by changes in above‐ground biomass but rather by differing responses of individual species in the two communities.     

Effects of resource additions on species richness and ANPP in an alpine meadow community

Aims Theories based on resource additions indicate that plant species richness is mainly determined by the number of limiting resources. However, the individual effects of various limiting resources on species richness and aboveground net primary productivity (ANPP) are less well understood. Here, we analyzed potential linkages between additions of limiting resources, species loss and ANPP increase and further explored the underlying mechanisms.Methods Resources (N, P, K and water) were added in a completely randomized block design to alpine meadow plots in the Qinghai-Tibetan Plateau. Plant aboveground biomass, species composition, mean plant height and light availability were measured in each plot. Regression and analysis of variance were used to analyze the responses of these measures to the different resource-addition treatments.Important findings Species richness decreased with increasing number of added limiting resources, suggesting that plant diversity was apparently determined by the number of limiting resources. Nitrogen was the most important limiting resource affecting species richness, whereas P and K alone had negligible effects. The largest reduction in species richness occurred when all three elements were added in combination. Water played a different role compared with the other limiting resources. Species richness increased when water was added to the treatments with N and P or with N, P and K. The decreases in species richness after resource additions were paralleled by increases in ANPP and decreases in light penetration into the plant canopy, suggesting that increased light competition was responsible for the negative effects of resource additions on plant species richness.     

The influence of vines on an oligohaline marsh community: results of a removal and fertilization study

The effects of competitive suppression by vines on the non-vine plant community have received little attention in temperate habitats. This study investigated the impact vines have on their herbaceous hosts in a wetland community at two soil fertility levels. Plots in an oligohaline marsh were treated in a 2 × 2 factorial design with vine removal and fertilization over two growing seasons. There was no significant interaction between removal and fertilization treatments on any of the measured variables. Vine removal initially caused an increase in light penetration through the canopy, but by the end of the study, plots with vines removed had less light due to a 25% increase in biomass by the plants released from competition with vines. For plots with vines removed, species richness was higher during a brief period in the spring of the second year, but by the end of the study, richness in removal plots decreased relative to controls. Fertilization caused a 40% increase in biomass overall, although only two species, Sagittaria lancifolia L. and Polygonum punctatum Ell., showed dramatic increases. Despite fertilization causing a 40% decrease in light penetration to the ground, no change in species richness was observed. Overall, these results show that vine cover in this wetland suppresses non-vine species and reduces community biomass. Removal of vines increased biomass of non-vine dominants but resulted in only an ephemeral change in species richness. Fertilization did not increase the effects of vines on the non-vine community. Received: 14 November 1996 / Accepted: 10 June 1997     

From plant neighbourhood to landscape scales: how grazing modifies native and exotic plant species richness in grassland

The interactive effect of grazing and soil resources on plant species richness and coexistence has been predicted to vary across spatial scales. When resources are not limiting, grazing should reduce competitive effects and increase colonisation and richness at fine scales. However, at broad scales richness is predicted to decline due to loss of grazing intolerant species. We examined these hypotheses in grasslands of southern Australia that varied in resources and ungulate grazing intensity since farming commenced 170 years ago. Fine-scale species richness was slightly greater in more intensively grazed upper slope sites with high nutrients but low water supply compared to those that were moderately grazed, largely due to a greater abundance of exotic species. At broader scales, exotic species richness declined with increasing grazing intensity whether nutrients or water supply were low or high. Native species richness declined at all scales in response to increasing grazing intensity and greater resource supply. Grazing also reduced fine-scale heterogeneity in native species richness and although exotics were also characterised by greater heterogeneity at fine scales, grazing effects varied across scales. In these grasslands patterns of plant species richness did not match predictions at all scales and this is likely to be due to differing responses of native and exotic species and their relative abundance in the regional species pool. Over the past 170 years intolerant native species have been eliminated from areas that are continually and heavily grazed, whereas transient, light grazing increases richness of both exotics and natives. The results support the observation that the processes and scales at which they operate differ between coevolved ungulate—grassland systems and those in transition due to recent invasion of herbivores and associated plant species.     

Staged invasions across disparate grasslands: effects of seed provenance,consumers and disturbance on productivity and species richness

Exotic plant invasions are thought to alter productivity and species richness, yet these patterns are typically correlative. Few studies have experimentally invaded sites and asked how addition of novel species influences ecosystem function and community structure and examined the role of competitors and/or consumers in mediating these patterns. We invaded disturbed and undisturbed subplots in and out of rodent exclosures with seeds of native or exotic species in grasslands in Montana, California and Germany. Seed addition enhanced aboveground biomass and species richness compared with no‐seeds‐added controls, with exotics having disproportionate effects on productivity compared with natives. Disturbance enhanced the effects of seed addition on productivity and species richness, whereas rodents reduced productivity, but only in Germany and California. Our results demonstrate that experimental introduction of novel species can alter ecosystem function and community structure, but that local filters such as competition and herbivory influence the magnitude of these impacts.     

Interactions among irradiance,nutrients, and herbivores constrain a stream algal community

Using stream-side, flow-through channels, I tested for the effects of nutrients (NU) (nitrogen plus phosphorus), irradiance (L), and snail grazing (G) on a benthic algal community in a small, forested stream. Grazed communities were-dominated by a chlorophyte (basal cells ofStigeoclonium) and a cyanophyte (Chamaesiphon investiens), whereas ungrazed communities were comprised almost entirely of diatoms, regardless of nutrient and light levels. Snails maintained low algal biomass in all grazed treatments, presumably by consuming increased algal production in treatments to which L and NU were increased. When nutrients were increased, cellular nutrient content increased under ambient conditions (shaded, grazed) and biomass and productivity increased when snails were removed and light was increased. Together, nutrients and light had positive effects and grazing had negative effects on biomass (chlorophylla, AFDM, algal biovolume) and chlorophyll-and areal-specific productivity in ANOVAs. However, in most cases, only means from treatments in which all three factors were manipulated (ungrazed, +NU&L treatments) were significantly different from controls; effects of single factors were generally undetectable. These results indicate that all three factors simultaneously limited algal biomass and productivity in this stream during the summer months. Additionally, the effects of these factors in combination were in some cases different from the effects of single factors. For example, light had slight negative effects on some biomass parameters when added at ambient snail densities and nutrient concentrations, but had strong positive effects in conjunction with nutrient addition and snail removal. This study demonstrates that algal biomass and productivity can be under multiple constraints by irradiance, nutrients, and herbivores and indicates the need to employ multifactor experiments to test for such interactive effects.     

Dispersal and seed limitation affect diversity and productivity of montane grasslands

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity–ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m⁻² and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m⁻². One year after seed addition, local plant species richness had increased on average by six species m⁻² (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m⁻² (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump‐shaped model appears to be the limiting outline of the natural diversity–productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems.     

Effect of functional group richness and species richness in manipulated productivity–diversity studies: a glasshouse pot experiment

Species and functional group (grasses, legumes, creeping nonlegume forbs, rosette nonlegume forbs) richness of species assemblages composed of 16 species from four functional plant groups were manipulated to evaluate the productivity-diversity relationships in a greenhouse pot experiment. Pots were filled with sand, and supplied at two levels of nutrients. The plants were grown in monocultures, two, four, eight and 16 species mixtures. Individual two, four, and eight species mixtures differed in the richness of functional groups. Although the two characteristics of biodiversity, i.e. species and functional group richness, were necessarily correlated, it was shown that it is possible to separate their effect statistically, and also test for their common effect without pronounced loss of test power. There was a pronounced increase of average aboveground biomass and a mild increase in belowground biomass with biodiversity. The effect of functional group richness was more pronounced than the effect of the number of species. By using the method of Loreau and Hector (Nature 411 (2001) 72), selection and complementarity effects were statistically separated, and the overyielding index was calculated as a ratio of the productivity of a mixture to the productivity of its most productive component (to demonstrate transgressive overyielding). Positive values of complementarity and transgressive overyielding were both found, particularly in some rich communities and under high nutrient levels. Complementarity significantly increased only with functional group richness and mainly under high nutrients in the belowground biomass. Some species, when grown in monocultures, had decreased productivity under higher nutrients, and thus were more productive in mixtures than in monocultures. It seems that those species suffered from too high nutrient levels when grown in monocultures, but not in the presence of other species, which were able to use the nutrients in high concentrations and effectively decrease the nutrient levels. As a consequence, mixtures of high diversity were always more productive under high nutrients. The difference in species proportions between high and low nutrients, characterized by chord distance, increased with species richness. The relative change in productivity decreased with the number of functional groups. This suggests that species richness might lead to stabilization of aggregate characteristics (like total productivity) under changing environmental conditions by changing the proportions of individual species.     

Seed limitation interacts with biotic and abiotic factors to constrain novel species' impact on community biomass and richness

Seed limitation can narrow down the number of coexisting plant species, limit plant community productivity, and also constrain community responses to changing environmental and biotic conditions. In a 10-year full-factorial experiment of seed addition, fertilisation, warming and herbivore exclusion, we tested how seed addition alters community richness and biomass, and how its effects depend on seed origin and biotic and abiotic context. We found that seed addition increased species richness in all treatments, and increased plant community biomass depending on nutrient addition and warming. Novel species, originally absent from the communities, increased biomass the most, especially in fertilised plots and in the absence of herbivores, while adding seeds of local species did not affect biomass. Our results show that seed limitation constrains both community richness and biomass, and highlight the importance of considering trophic interactions and soil nutrients when assessing novel species immigrations and their effects on community biomass.     

Vesicular-arbuscular mycorrhizae in a wastewater-irrigated oldfield ecosystem in Michigan

The incidence of vesicular-arbuscular mycorrhizae (VAM) in wastewater irrigated and non-irrigated oldfield soils in Michigan was studied. Soil and root samples were taken monthly from field plots on the second and third years of consecutive irrigation with municipal wastewater at rates of 0, 5 and 10 cm wk^–1. The oldfield ecosystem contained a high VAM fungal spore population density, but low species diversity. The most common VAM fungal species were Glomus mosseae and G. fasciculatum. Both spore density and root colonization were higher in irrigated than in non-irrigated plots. Irrigation effects were largest early in the growing season. In addition to increasing VAM incidence, wastewater irrigation shifted VAM fungal species composition. Irrigation favored G. mosseae over G. fasciculatum. Bioassays using either Sorghum vulgare or Daucus carota, an oldfield native species, indicated that the VAM systems were still functioning after the third year of consecutive wastewater irrigation. The data from experiments using nutrient solutions at wastewater concentrations suggest that the effects of wastewater irrigation on VAM are due to the effects of both water and nutrients. Since VAM are a very important component of the plant's water and nutrient uptake system and equally important in structuring plant communities under limiting growth conditions, it is suggested that the stimulatory effect of wastewater irrigation on VAM in an oldfield ecosystem enhances the ecosystem's ability to function as a living filter for wastewater clean up.Michigan Agricultural Experimental Station Journal Article No 13137.On sabbatical leave from ESAL, Lavras-MG, Brazil     

Impact of management on biodiversity-biomass relations in Estonian flooded meadows

A negative species richness–productivity relationship is often described in grasslands at smaller scales. We aimed to study the effect of management on this relationship. In particular, we addressed the relative importance of biomass cutting, hay removal and nutrient impoverishment on species richness and growth form structure. We conducted fieldwork in flooded meadows in Alam-Pedja Nature Reserve, central Estonia. We sampled vegetation in managed and abandoned stands of two types of alluvial meadows, sedge and tall forb meadow. Aboveground biomass and litter were harvested, weighed and analysed for major plant nutrients by near infrared reflectance spectroscopy. Three groups of general additive models were developed and compared, addressing the impact of (i) productivity, (ii) nutrients and (iii) management regime on species richness. Management—mowing and hay removal—reduced the amount of litter but not aboveground biomass. Management led to a decrease in nitrogen in the biomass and enhanced species richness, particularly in the tall forb meadow. The strongest determinant of species richness was the amount of litter, exhibiting a hump-shaped relationship with richness. The effect of nitrogen supply was significant, but explained less variation. Management increased the proportion of sedges in the sedge meadow and of small herbs in the tall forb meadow. We conclude that litter removal is the most important management means to support biodiversity. On highly productive sites, reducing nutrients via hay removal is of secondary importance within a timeframe of 10 years.     

Effects of productivity on species–area curves in herbaceous vegetation: evidence from experimental and observational data

The effects of productivity on the parameters of the species–area curve were investigated in this paper using two data sets on terrestrial plant communities: (1) one including 48 plots in 12 experimental sites on ploughed, formerly cultivated fields in the Siena region, Italy, and (2) one including 40 plots in hay meadows in the Bremen region, Germany. In both regions, species presence of vascular plants was recorded in nested plots ranging in size from 0.004 to 256 m² and 0.001 to 1000 m², respectively. Productivity was estimated as dry standing biomass. In the Siena data set, species richness showed a humped‐back relation to biomass in the plot sizes up to 1 m². For the larger plot sizes, no significant correlations were found. In the Bremen data set, positive relation between species number and biomass was observed at the smallest spatial scale (0.001 m²), whereas the relation disappeared or tended to be negative for the larger plot sizes. In general, the slopes z of the log species–log area curves (SAC) were negatively related to biomass in both data sets, while the intercept c increased with biomass in the Siena data set and was unrelated to biomass in the Bremen data set. The relationship between c and z was negative in the Siena data set and positive in the Bremen data set. The above results differed somewhat depending on which plot sizes were considered for the calculation of the SAC. Literature data confirmed that there are no clear patterns in the inter‐correlations between productivity, small scale and large scale species richness. Sites differing in productivity and in the slopes and intercepts of SAC may thus give rise to different species richness–productivity relationships. There can be one possible relation between species richness and biomass at one spatial scale (e.g. humped‐back) and another type of relation, even opposite, at another spatial scale. This suggests that the properties of species–area curves do not respond in a uniform way to the changes in productivity, but depend on the type of habitat or plant community and its particular properties. The parameter of the SAC can then hardly be used as scale‐independent parameter to investigate the effects of ecological factors, such as productivity, on species richness. The lack of clear patterns in the relations between small scale and large scale species richness implies that the predictions of the species‐pool hypothesis may fail when applied to plot sizes as dealt with in this study.     

施氮对空心莲子草(Alternanthera philoxeroides)和莲子草(Alternanthera sessilis)种间关系的影响

自然界的氮素释放总是呈现出空间和时间上的异质性,但关于异质性氮释放对于入侵植物和本地植物种间关系影响的研究相对较少。将入侵植物空心莲子草(Alternanthera philoxeroides)和同属本地植物莲子草(Alternanthera sessilis)分别进行单种种植(12株,无种间竞争)和混种种植(每种6株,有种间竞争),模拟大气氮湿沉降设置由两种不同施氮总量(15g N m~(-2)a~(-1)和30g N m~(-2)a~(-1))和两种不同施氮频率(每5天1次和每15天1次)交叉组成的4种施氮处理,并以不施氮为对照。施氮总量的增加显著促进了两种植物的生长,但对两种植物的种间竞争关系没有显著影响。施氮频率对两种植物的生长以及种间竞争关系都没有显著影响。两种植物在面对竞争时表现出不同的生物量分配策略,空心莲子草将更多的生物量分配到茎,而莲子草将更多的生物量分配到根。在全球变化的背景下,大气氮湿沉降可能会改变两种植物的种群结构和动态,但可能对这两种植物的种间关系影响较小。     

Adaptive survival mechanisms and growth limitations of small-stature herb species across a plant diversity gradient

Several biodiversity experiments have shown positive effects of species richness on aboveground biomass production, but highly variable responses of individual species. The well-known fact that the competitive ability of plant species depends on size differences among species, raises the question of effects of community species richness on small-stature subordinate species. We used experimental grasslands differing in species richness (1-60 species) and functional group richness (one to four functional groups) to study biodiversity effects on biomass production and ecophysiological traits of five small-stature herbs (Bellis perennis, Plantago media, Glechoma hederacea, Ranunculus repens and Veronica chamaedrys). We found that ecophysiological adaptations, known as typical shade-tolerance strategies, played an important role with increasing species richness and in relation to a decrease in transmitted light. Specific leaf area and leaf area ratio increased, while area-based leaf nitrogen decreased with increasing community species richness. Community species richness did not affect daily leaf carbohydrate turnover of V. chamaedrys and P. media indicating that these species maintained efficiency of photosynthesis even in low-light environments. This suggests an important possible mechanism of complementarity in such grasslands, whereby smaller species contribute to a better overall efficiency of light use. Nevertheless, these species rarely contributed a large proportion to community biomass production or achieved higher yields in mixtures than expected from monocultures. It seems likely that the allocation to aboveground plant organs to optimise carbon assimilation limited the investment in belowground organs to acquire nutrients and thus hindered these species from increasing their performance in multi-species mixtures.     

Factors affecting growth rates of young tropical floodplain fishes: seasonality and density-dependence

Synopsis Deviations of growth increments from a model describing average growth of 12 common fish species, mostly juveniles, were compared with hydrological variables and biomass density (ranging from 1 gm^-2 to 83 gm^-2 of potentially competing species of similar size ranges (guilds). Seasonal effects on growth were highly significant for omnivores but not for detritivores. Omnivores exhibited faster growth during the rising water season, and higher growth rates were associated with periods of increased rates of flooding during this season, which could be associated with increased availability of food. No species exhibited density-dependent growth when all seasons were considered together. During the rising-water-level season, no inverse relationship between growth and biomass density was found in any of the 11 species tested at p = 0.05. During the short falling-water season, 2 of 8 species tested did exhibit density-dependent growth but a sign test of correlation coefficients from all species was not significant. These two significant results were from four omnivores tested. However, density-dependent growth was not indicated when the data were pooled within either detritivore or omnivore guilds by hydrological season. It was concluded that with the possible exception of juvenile omnivores during the limited falling-water season, density-dependent growth, and by implication interspecific competition, had no effect in regulating the species' populations investigated in the floodplain environment. Conversely, the importance of a seasonal hydrological regime in maintaining growth rates, at least for omnivores, was evident.     

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。     

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.