Ecosystem allometry: the scaling of nutrient stocks and primary productivity across plant communities

A principal challenge in ecology is to integrate physiological function (e.g. photosynthesis) across a collection of individuals (e.g. plants of different species) to understand the functioning of the entire ensemble (e.g. primary productivity). The control that organism size exerts over physiological and ecological function suggests that allometry could be a powerful tool for scaling ecological processes across levels of organization. Here we use individual plant allometries to predict how nutrient content and productivity scale with total plant biomass (phytomass) in whole plant communities. As predicted by our model, net primary productivity as well as whole community nitrogen and phosphorus content all scale allometrically with phytomass across diverse plant communities, from tropical forest to arctic tundra. Importantly, productivity data deviate quantitatively from the theoretically derived prediction, and nutrient productivity (production per unit nutrient) of terrestrial plant communities decreases systematically with increasing total phytomass. These results are consistent with the existence of pronounced competitive size hierarchies. The previously undocumented generality of these 'ecosystem allometries' and their basis in the structure and function of individual plants will likely provide a useful quantitative framework for research linking plant traits to ecosystem processes.     

Size-dependent species richness: trends within plant communities and across latitude

We examine how species richness and species‐specific plant density (number of species and number of individuals per species, respectively) vary within community size frequency distributions and across latitude. Communities from Asia, Africa, Europe, and North, Central and South America were studied (60°4′N–41°4′S latitude) using the Gentry data base. Log–log linear stem size (diameter) frequency distributions were constructed for each community and the species richness and species‐specific plant density within each size class were determined for each frequency distribution. Species richness in the smallest stem size class correlated with the Y‐intercepts (β‐values) of the regression curves describing each log–log linear size distributions. Two extreme community types were identified (designated as type A and type B). Type A communities had steep size distributions (i.e. large β‐values), log–log linear species‐richness size distributions, low species‐specific plant density distributions, and a small size class (2–4 cm) containing the majority of all species but rarely conspecifics of the dominant tree species. Type B communities had shallow size distributions (i.e. small β‐values), more or less uniform (and low) size class species‐ richness and species‐specific density distributions and size‐dominant species resident in the smallest size class. Type A communities were absent in the higher latitudes but increased in number towards the equator, i.e. in the smallest size class, species richness increased (and species‐specific density decreased) towards the tropics. Based on our survey of type A and type B communities (and their intermediates), species richness evinces size‐dependent and latitudinal trends, i.e. species richness increased with decreasing body size and most species increasingly reside in the smallest plant size class towards the tropics. Across all latitudes, a trade‐off exists between the number of species and the number of individuals per species residing in the smaller size classes.     

Food webs obscure the strength of plant diversity effects on primary productivity

Plant diversity experiments generally find that increased diversity causes increased productivity; however, primary productivity is typically measured in the presence of a diverse food web, including pathogens, mutualists and herbivores. If food web impacts on productivity vary with plant diversity, as predicted by both theoretical and empirical studies, estimates of the effect of plant diversity on productivity may be biased. We experimentally removed arthropods, foliar fungi and soil fungi from the longest‐running plant diversity experiment. We found that fungi and arthropods removed a constant, large proportion of biomass leading to a greater reduction of total biomass in high diversity plots. As a result, the effect of diversity on measured plant productivity was much higher in the absence of fungi and arthropods. Thus, diversity increases productivity more than reported in previous studies that did not control for the effects of heterotrophic consumption.     

Unanticipated impacts of spatial variance of biodiversity on plant productivity

Experiments on biodiversity have shown that productivity is often a decelerating monotonic function of biodiversity. A property of nonlinear functions, known as Jensen's inequality, predicts negative effects of the variance of predictor variables on the mean of response variables. One implication of this relationship is that an increase in spatial variability of biodiversity can cause dramatic decreases in the mean productivity of the system. Here I quantify these effects by conducting a meta‐analysis of experimental data on biodiversity–productivity relationships in grasslands and using the empirically derived estimates of parameters to simulate various scenarios of levels of spatial variance and mean values of biodiversity. Jensen's inequality was estimated independently using Monte Carlo simulations and quadratic approximations. The median values of Jensen's inequality estimated with the first method ranged from 3.2 to 26.7%, whilst values obtained with the second method ranged from 5.0 to 45.0%. Meta‐analyses conducted separately for each combination of simulated values of mean and spatial variance of biodiversity indicated that effect sizes were significantly larger than zero in all cases. Because patterns of biodiversity are becoming increasingly variable under intense anthropogenic pressure, the impact of loss of biodiversity on productivity may be larger than current estimates indicate.     

A comparison of methods for measuring primary productivity and community respiration in streams

Carbon dioxide and oxygen exchange procedures for measuring community metabolism (two open stream methods and three chamber methods) were compared on the same reach of a third-order stream. Open stream methods were complicated by high diffusion rates and yielded net community primary productivity estimates lower than those obtained with chamber methods. Chamber methods yielded variable productivity and respiration data. However, when normalized for chlorophyll a, productivity estimates from the chamber methods were within an expected range for the system. Balances of photosynthesis and respiration from the chamber methods were similar between methods and indicated that autotrophic or heterotrophic processes could dominate the system. Considerations in applying the various procedures are discussed.     

Species richness and canopy productivity of Australian plant communities

The species richness (number of vascular plants per hectare) of Australian plant communities (containing a mosaic of gap, regeneration, maturation and senescent phases) is correlated with the annual biomass productivity of the overstorey canopy.The annual production of leaves and stem in the canopy of the plant community is shown to be limited by the requirements of photosynthesis (particularly light and the availability of water) and the length of the growing season.The species richness of Australian plant communities is the product of the blance between the dominance of the overstorey and the response of the understorey to the shading of the overstorey. For all climatic regions and zones the species richness of the overstorey of the plant community is shown to be exponentially related to the annual shoot growth of the overstorey canopy, until the latitudinal or altitudinal tree line is reached. With latitudinal increase outside the tropics, overstorey canopies of forest communities absorb increasingly more of the incident solar radiation. markedly reducing the species richness of the understorey strata. In contrast, in these latitudes the overstorey of plant communities with widely spaced trees or tall shrubs will absorb far less solar radiation, thus enabling the species richness of the understorey to be maintained.     

Functional trait variation of forest understorey plant communities across Europe

Global environmental changes are expected to alter the functional characteristics of understorey herb-layer communities, potentially affecting forest ecosystem functioning. However, little is known about what drives the variability of functional traits in forest understories. Here, we assessed the role of different environmental drivers in shaping the functional trait distribution of understorey herbs in fragmented forests across three spatial scales. We focused on 708 small, deciduous forest patches located in 16 agricultural landscape windows, spanning a 2500-km macroclimatic gradient across the temperate forest biome in Europe. We estimated the relative effect of patch-scale, landscape-scale and macroclimatic variables on the community mean and variation of plant height, specific leaf area and seed mass. Macroclimatic variables (monthly temperature and precipitation extremes) explained the largest proportion of variation in community trait means (on average 77% of the explained variation). In contrast, patch-scale factors dominated in explaining community trait variation (on average 68% of the explained variation). Notably, patch age, size and internal heterogeneity had a positive effect on the community-level variability. Landscape-scale variables explained only a minor part of the variation in both trait distribution properties. The variation explained by shared combinations of the variable groups was generally negligible. These findings highlight the importance of considering multiple spatial scales in predictions of environmental-change effects on the functionality of forest understories. We propose that forest management sustainability could benefit from conserving larger, historically continuous and internally heterogeneous forest patches to maximise ecosystem service diversity in rural landscapes.     

陕西省礼泉县耕地产能核算

基于农用地分等成果,在建立关中渭河平原区理论和可实现单产模型的基础上,以乡镇为研究单元,计算了礼泉县理论产能、可实现产能和实际产能.结果表明,2008年礼泉县理论产能为39.60万t,可实现产能为29.82万t,实际产能为18.69万t;理论单产为19071 kg·hm-2,可实现单产为14745 kg·hm-2,实际单产为9042 kg.hm-2;理论利用强度为0.764,可实现利用强度为0.639;理论利用潜力为4376 kg·hm-2,可实现利用潜力5115kg·hm-2.礼泉县三大产能在空间上差异明显,理论单产、可实现单产和实际单产在空间上从南向北呈现差异明显阶梯状分布,南部乡镇单产最大、中部乡镇单产居中、北部乡镇单产最小.造成单产差异明显的原因是区域的地形地貌、土壤类型、投入水平、技术等,未来南部和中部的各乡镇是该县粮食增产的重点区域.     

Effects of plant functional group removal on structure and function of soil communities across contrasting ecosystems

Loss of plant diversity has an impact on ecosystems worldwide, but we lack a mechanistic understanding of how this loss may influence below‐ground biota and ecosystem functions across contrasting ecosystems in the long term. We used the longest running biodiversity manipulation experiment across contrasting ecosystems in existence to explore the below‐ground consequences of 19 years of plant functional group removals for each of 30 contrasting forested lake islands in northern Sweden. We found that, against expectations, the effects of plant removals on the communities of key groups of soil organisms (bacteria, fungi and nematodes), and organic matter quality and soil ecosystem functioning (decomposition and microbial activity) were relatively similar among islands that varied greatly in productivity and soil fertility. This highlights that, in contrast to what has been shown for plant productivity, plant biodiversity loss effects on below‐ground functions can be relatively insensitive to environmental context or variation among widely contrasting ecosystems.     

Diversity partitioning of moorland plant communities across hierarchical spatial scales

Understanding of the scaling of diversity is critical to enhance conservation strategies for subalpine moorland ecosystems vulnerable to future environmental changes. However, a paucity of quantitative data strongly limits such attempts. In this study, we used an additive diversity partitioning framework and quantified diversity patterns of moorland plant communities across hierarchical spatial scales, within- and between-sample transects, and between sites (corresponding to α and two levels of β diversity). Moorland sites markedly differed in size (range 1,000–160,000 m²) and were isolated from each other to varying extents within an inhospitable matrix (i.e., forests). We found that β diversity components were consistently higher, whereas the local α diversity component was consistently lower than expected by chance. We observed substantial contribution at the between-site scale to total species richness. By focusing on diversity patterns of moorland plant communities across multiple hierarchical spatial scales, we could thus identify the scale at which regional diversity is maximized. Our results suggest that protection of as many moorland sites as possible, to ensure beta diversity between sites, will effectively conserve total diversity. The use of additive diversity partitioning is a major step forward in providing strategies for the biological conservation of subalpine moorland ecosystems vulnerable to future environmental changes.     

Overyielding and the role of complementary use of nitrogen in wetland plant communities

Biodiversity and ecosystem functioning experiments have demonstrated that plant biomass of species grown in mixtures is often greater than plant biomass of monocultures (i.e., mixtures over yield). While we understand that plant species utilize resources differently, how a combination of species increases resource use and productivity is not well known, especially in wetland ecosystems. Here, we used a mesocosm experiment to explore diversity effects on plant biomass production and to examine the role of N partitioning as a mechanism for overyielding in wetland ecosystems. Plant functional groups (FGs) represented the unit of diversity, and we included five levels of diversity (0-4 FGs). To test for N partitioning, we used a stable isotope technique to determine niche breadth and proportion similarity of inorganic N use (NO₃⁻ and NH₄⁺) for individual FGs as well as mixtures containing 3 and 4 FGs. We found that total plant biomass increased in the first season from an average of 290 ± 60 SE g ash-free dry mass (AFDM) m⁻² at the 1 FG level to 490 ± 70 g AFDM m⁻² at the 4 FG level and in the second season from an average of 560 ± 80 g AFDM m⁻² at the 1 FG level to 1000 ± 90 g AFDM m⁻² at the 4 FG level indicating overyielding. Plant species comprising the majority of mesocosm biomass demonstrated preferential uptake of ¹⁵NO₃⁻, while species with relatively less biomass (e.g., Acorus calamus and Carex crinita) preferred ¹⁵NH₄⁺. Concentrations of ¹⁵N in biomass increased with FG richness, but only in the ¹⁵NO₃⁻ treatment. Niche breadth did not vary among levels of FG richness. We observed a greater niche overlap with an increase of FGs, with species taking up greater proportion of ¹⁵NO₃⁻ than ¹⁵NH₄⁺. Our results indicate that plant overyielding in wetland mesocosms is not the result of niche partitioning of N chemical forms, but is associated with greater uptake of NO₃.     

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.     

高寒草甸不同草地群落物种多样性与生产力关系研究

生态系统的结构和功能、生物多样性与生产力的关系问题是近年来群落生态学中研究的中心问题,其中,生态系统生产力水平是其功能的重要表现形式,用4种不同草地类型探讨自然群落的物种多样性与生产力关系.结果表明,矮嵩草草甸、小嵩草草甸和金露梅灌丛群落中物种多样性与生产力的关系呈线性增加关系,藏嵩草沼泽化草甸群落中线性增加关系不显著,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.不同的环境资源和环境异质性是形成群落结构特征、物种多样性分布格局差异的主要原因之一.     

Forest plant communities of the Baohua Mountains,eastern China

Abstract. In the forest of the Baohua Reserve in the Baohua Mountains, eastern China, six plant communities and 11 sub‐communities were described using the Braun‐Blanquet method. These communities can be assigned to four already described Chinese vegetation types. Curtis’ Compositional Index (CI) and Shannon's diversity index were used to determine the successional stage and species diversity of the communities. The community of Brachystachyum densiflorum and Pinus massoniana and the subcommunity of Fraxinus chinensis and Quercus acutissima with Pueraria lobata had the lowest CI and would be an initial successional stage; the community of Phoebe sheareri and Cyclobalanopsis glauca had the highest CI and would be the final stage. The community of Brachystachyum densiflorum and Pinus massoniana and the sub‐community of Fraxinus chinensis and Quercus acutissima with Pueraria lobata, both pioneer successional stages, had the lowest species diversity. The climax communities had the highest diversity.     

高寒草地植物群落地上-地下净初级生产力权衡

植物生产力分配和权衡是植物生态学研究中的热点,反映植物对环境的适应性,是了解植物响应全球气候变化的关键。青藏高原作为气候变化敏感区,研究其植物地上与地下部分权衡对了解高寒草地植被生存策略和生态系统可持续发展具有重要意义。目前,生物量分配调控机制已被广泛研究,但主要使用植物根冠比和地上-地下生物量比等方法来表征植物分配模式,缺乏考虑因植物生长周期导致的差异。使用青藏高原高寒草地103个样点的地上和地下净初级生产力数据,分析高寒草甸和高寒草原两种主要草地类型的地上-地下净初级生产力权衡关系。利用气候因素和土壤因素等相关数据,结合方差分析、相关分析、相对重要性分析和结构方程模型的方法,探究环境因素对两种草地类型地上-地下净初级生产力权衡的影响机制。研究发现：（1）高寒草甸的地上净初级生产力、地下净初级生产力和土壤养分含量显著高于高寒草原（P<0.05）;（2）高寒草地植被生产力均向地下权衡（0.0199）,且高寒草原（0.0354）的权衡值高于高寒草甸（0.0173）;（3）结构方程模型发现,年平均降水量、土壤容重和土壤速效氮含量是影响高寒草甸生产力权衡的主导因子,而年平均温度和年平均降水量是影响高寒草原生产力权衡的主导因子。研究表明高寒草甸的生产力权衡主要受气候和土壤因素共同影响,而高寒草原主要受气候因素调节。研究为理解植物地上-地下生物量分配调控机制提供了新的视角和方法,对系统了解高寒草地生物量分配模式和准确预测高寒草地植被动态过程具有指导意义。     

Spatial patterns of plant diversity and communities in Alpine ecosystems of the Hengduan Mountains,Northwest Yunnan,China

Aims Conduct a quantitative, but rapid, regional-level assessment of the alpine flora across northwest Yunnan (NWY) to provide a broad-based understanding of local and regional patterns of the composition, diversity and health of alpine ecosystems across NWY.Methods A stratified random sampling design was employed to select sites across the different mountain ranges of NWY. Vegetation was sampled by stratifying each site by the three major alpine vegetation community types: meadow, dwarf shrub and scree. Two 50-m transects were randomly located within each community type at each sampling site with 10 1-m 2 subplots systematically placed along each transect. Environmental variables were recorded at each transect. Multivariate analyses were used to classify the major plant community assemblages and link community patterns to environmental and habitat variables.Important findings Forb species richness varied from 19 to 105 species per site (21 sites total) with an average of 59 species per site (60 m 2 sampled per site). Most species were patchily distributed with narrow distributions and/or small population sizes; over half the species occurred at only one or two sites. Distinct species assemblages were identified in the meadow vegetation that was strongly aggregated by geographic location suggesting the presence of distinct phytogeographic zones of the meadow alpine flora. Elevation and geographic location were the dominant environmental gradients underlying the variations in species composition. Jaccard's coefficient of similarity averaged only 10% among sites indicating there was little similarity in the alpine flora across the region. The alpine vegetation is highly heterogeneous across the complex landscape of the Hengduan Mountains of NWY. Conservation strategies need to take into account the large geographic differences in the flora to maximize protection of biodiversity.     

Diversity of arbuscular mycorrhizal fungi across a fragmented forest in Panama: insular spore communities differ from mainland communities

It is now understood that alterations in the species composition of soil organisms can lead to changes in aboveground communities. In this study, we assessed the importance of spatial scale and forest size on changes in arbuscular mycorrhizal fungal (AMF) spore communities by sampling AMF spores in soils of forested mainland and island sites in the vicinity of Gatun Lake, Republic of Panama. We encountered a total of 27 AMF species or morphospecies, with 17, 8, 1 and 1 from the genera Glomus, Acaulospora, Sclerosystis, and Scutellospora, respectively. At small scales (<100 m²), we found little evidence for spatial structuring of AMF communities (decay of Morisita-Horn community similarity with distance). However, at large spatial scales, we found that the AMF spore community of a mainland plot was more similar to other mainland plots several kilometers (>5) away than to nearby island plots (within 0.7 km). Likewise, most island plots were more similar to other island plots regardless of geographic separation. There was no decay in AMF species richness (number of species), or Shannon diversity (number of species and their spore numbers) either with decreasing forest-fragment size, or with decreasing plant species richness. Of the six most common species that composed almost 70% of the total spore volume, spores of Glomus tsh and G. clavisporum were more common in soils of mainland plots, while spores of Glomus small brown and Acaulospora mellea were more abundant in soils of island plots. None of these common AMF species showed significant associations with soil chemistry or plant diversity. We suggest that the convergence of common species found in AMF spore communities in soils of similar forest sizes was a result of forest fragmentation. Habitat-dependent convergence of AMF spore communities may result in differential survival of tree seedlings regenerating on islands versus mainland.