A resource-based conceptual model of plant diversity that reassesses causality in the productivity–diversity relationship

Biogeographical studies frequently reveal positive correlations between species richness and estimates of environmental water and/or energy. A popular interpretation of this relationship relates the supply of water and energy to productivity, and then, in turn, to richness. Productivity–diversity theories are now legion, yet none has proved sufficiently intuitive to gain broad acceptance. Like productivity, heterogeneity is known to influence diversity at fine spatial scales, yet the possibility that richness might relate to water–energy dynamics at coarse spatial scales via a heterogeneity‐generating mechanism has received little attention. In this paper we outline such a conceptual model for plants that is internally consistent and testable. We believe it may help to explain the capacity of environments receiving different inputs of water and energy to support variable numbers of species at a range of spatial scales, the pervasive correlation between productivity and richness, some exceptions to the productivity–diversity relationship, the form of productivity–diversity curves and the link between richness and environmental ‘harshness’. The model may also provide an answer to one of the most venerable puzzles in the field of diversity studies: why high inputs of water and energy correspond to more species rather than simply more individuals.     

喀斯特常绿落叶阔叶混交林物种多度与丰富度空间分布的尺度效应

物种多样性的空间分布格局及其尺度效应是生态学研究的重点,对于理解物种多样性的形成和维持机制以及生物多样性的管理和保护均具有重要意义。以贵州茂兰国家级自然保护区分布的亚热带原生性喀斯特常绿落叶阔叶混交林为研究对象,分析了2个1hm2(100m×100m)样地中物种多度和丰富度的空间分布特征及其与取样尺度的关系,采用方差和变异系数描述多度和丰富度在5个尺度(5m×5m,10m×10m,20m×20m,25m×25m,50m×50m)上的空间变异性。结果表明:(1)两个样地的物种多度和丰富度具有尺度依赖性特征;(2)由于多度具有叠加性,物种多度的方差随着尺度的增加呈线性增加,而变异系数呈线性下降;(3)丰富度的方差随尺度的增加表现出单峰分布的特征,在25 m×25 m尺度上达到最大值,变异系数则随取样尺度的增加而呈线性下降。研究表明,物种多度具有尺度推演规律,而丰富度却没有,因此,应慎重进行物种丰富度的尺度推演。在分析喀斯特森林物种多样性时,应注重尺度效应带来的影响。     

Local–regional diversity relationship varies with spatial scale in lotic diatoms

Aim Ecologists have shown increasing interest in the relative roles of local and regional factors in structuring biotic communities. One approach to studying this is to examine the relationship between local species richness (LSR) and regional species richness (RSR). We examined the LSR–RSR relationship in stream diatoms, using two data sets that varied in spatial extent. At broad spatial extent ranging across drainage systems, we expected climatic and dispersal‐related factors to constrain LSR, thus resulting in a linear LSR–RSR relationship. However, at small spatial scales dispersal across sites should be unconstrained, resulting in strong local interactions and a weak or asymptotic LSR–RSR relationship. Location Boreal streams in Finland. Methods For data set 1, we sampled 15 stream riffles (localities) in each of eight drainage systems (regions), with the latitudinal gradient between the southernmost and northernmost sites being almost 1100 km. For data set 2, a locality for estimating LSR was a single stone, and each riffle represented a region for estimating RSR. We sampled 20 stones in each of eight riffles. We used linear regressions to examine the relationship between LSR and RSR across regions. We used both observed richness values, as well as values estimated with the Chao1 estimator. Results We found a relatively strong linear relationship between the Chao1‐estimated mean LSR and RSR (R² = 0.654, P = 0.015) across drainage systems. The slope of the regression was 0.643 and it did not differ from 1.0, thus indicating linearity. At the riffle scale, however, LSR and RSR were not linearly related, and the slope of the regression (0.039) differed significantly from 1.0, indicating curvilinearity. Main conclusions These results suggest that the relationship between mean LSR and RSR varies across spatial scales in diatoms – from significantly linear at large scales to curvilinear at small scales. These plots imply strong regional enrichment in stream diatoms across drainage systems. Their diversity is thus determined largely by the composition of the regional species pool, as also in many macroorganisms. In contrast, at small spatial scales the LSR–RSR relationship implied a hard limit to local diversity, reflecting the primacy of local processes.     

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.     

Climate-induced temporal variation in the productivity–diversity relationship

Spatial scales are known to influence the form of the productivity–diversity relationship, but less attention has been given to the influence of temporal scales. Interannual climatic variation in semi-arid ponderosa pine–bunchgrass ecosystems causes significant year-to-year differences in herbaceous production. We hypothesized that unimodal (or 'hump-backed') relationships would be detected between herbaceous production and species richness in wet years, whereas positive logarithmic relationships would be detected in dry years. We analyzed nine years of herbaceous production and species richness data and used Akaike's information criterion (AICc) to determine the weight of evidence for each model in each year. As predicted, species richness exhibited a unimodal relationship to herbaceous production in wet years; however, richness exhibited a logarithmic relationship with herbaceous production in dry years. These results suggest that competitive exclusion occurred within this semi-arid plant community in years of high production when enough moisture was available to drive abundant plant growth. Thus, just as it is important to sample broad spatial variation in production to detect the full unimodal productivity–diversity relationship, it is also important to recognize that the full unimodal curve may be undetectable in less productive dry years in semi-arid ecosystems.     

Species' traits predict the effects of disturbance and productivity on diversity

Disturbance is an important factor influencing diversity patterns. Ecological theory predicts that diversity peaks at intermediate levels of disturbance, but this pattern is not present in a majority of empirical tests and can be influenced by the level of ecosystem productivity. We experimentally tested the effects of disturbance on diversity and show that species' autecological traits and community relations predicted species loss. We found that – alone or in concert – increasing disturbance intensity or frequency, or decreasing productivity, reduced diversity. Our species did not exhibit a clear competition-colonization trade-off, and intrinsic growth rate was a more important predictor of response to disturbance and productivity than measures of competitive ability. Furthermore, competitive ability was more important in predicting responses when, in addition to killing individuals, disturbance returned nutrients to the ecosystem. Our results demonstrate that species' traits can help resolve conflicting patterns in the response of diversity to disturbance and productivity.     

Mycorrhizal fungal identity and richness determine the diversity and productivity of a tallgrass prairie system

We investigated the effects of arbuscular mycorrhizal fungal (AMF) species richness and composition on plant community productivity and diversity, and whether AMF mediate plant species coexistence by promoting niche differentiation in phosphorus use. Our experiment manipulated AMF species richness and identity across a range of P conditions in tallgrass prairie mesocosms. We showed that increasing AMF richness promoted plant diversity and productivity, but that this AMF richness effect was small relative to the effects of individual AMF species. We found little support for AMF-facilitated complementarity in P use. Rather, the AMF richness effect appeared to be caused by the inclusion of particular diversity- and productivity-promoting AMF (a sampling effect). Furthermore, the identity of the diversity-promoting fungi changed with P environment, as did the relationship between the diversity-promoting and productivity-promoting benefits of AMF. Our results suggest that plant diversity and productivity are more responsive to AMF identity than to AMF diversity per se, and that AMF identity and P environment can interact in complex ways to alter community-level properties.     

青藏高原高寒草甸物种多样性的海拔梯度分布格局及对地上生物量的影响

植物物种多样性在海拔梯度上的变化规律以及物种多样性与生产力的关系是生态学研究的热点, 至今还没有得出一般性规律。本文以青海省海南藏族自治州贵德县的拉脊山(36°21′ N, 101°27′ E, 海拔3,389-3,876 m)和果洛藏族自治州的玛沁县军牧场山体(34°22′ N, 100°30′ E, 海拔4,121-4,268 m)为研究对象, 对植物高度、盖度、地上生物量和物种多样性随海拔高度的变化进行调查和统计分析, 以探讨青藏高原高寒草甸的物种多样性和地上生物量在海拔梯度上的变化规律及两者的关系。结果表明: (1)两条山体样带上地上生物量与物种多样性随海拔的变化规律一致: 随着海拔的升高, 地上生物量线性降低; Shannon-Wiener指数、Simpson指数和物种丰富度都呈单峰曲线, 在中间海拔最大, 而Pielou指数随海拔的升高线性增加。结合目前针对青藏高原高寒草甸的研究数据, 发现物种丰富度随海拔高度的变化均呈单峰曲线, 说明随着海拔的升高物种多样性先升高后降低可能是青藏高原物种多样性分布的普遍规律。(2)地上生物量与物种多样性的关系在两条山体样带上表现一致: 地上生物量随Shannon- Wiener指数、Simpson指数和Pielou指数的升高而线性降低, 但与物种丰富度不相关。综合两条山体样带所有样方数据, 发现地上生物量与Shannon-Wiener指数、Simpson指数不相关, 而随物种丰富度的升高线性增加。结合目前在青藏高原的相关研究数据, 发现地上生物量与物种丰富度呈S型曲线(logistic model)。     

Resource quantity, not resource heterogeneity, maintains plant diversity

Resource heterogeneity has often been proposed to explain the maintenance of plant species diversity and patterns of species diversity along productivity gradients. Resource heterogeneity should maintain biodiversity by preventing competitive exclusion because different species are superior competitors in different parts of a heterogeneous environment. In natural systems, however, resource heterogeneity covaries with average resource supply rate, making the effect of heterogeneity difficult to isolate. Using a novel experimental approach, we tested the independent effects of resource heterogeneity and average supply rate on plant species diversity. We show that the average supply rate of the most limiting resource controlled species diversity, whereas heterogeneity of this resource had virtually no effect. These findings also suggest that biodiversity declines with increasing productivity because at high enough levels of productivity one resource may always be driven to sufficiently short supply to exclude many species.     

外来种入侵与物种多样性

在入侵生态学研究方面 ,物种多样性与生物入侵之间的关系已成为当前研究和争论的焦点。自Elton的经典假说提出以来 ,物种丰富度高的群落比物种贫乏的群落更能抵抗外来种入侵的观点得到广泛接受。一些理论模型和多样性处理实验支持了该假说。但现在越来越多的野外观测和实验研究开始对这一经典假说提出异议 ,甚至反对。同时 ,在入侵生态学广泛受到关注的今天 ,大量的实验研究也提出了一些新的观点。本文对Elton经典假说提出以来全球有关物种多样性与生物入侵关系的主要研究及其观点进行了评述 ,以期为我国有关研究工作的开展提供参考。     

Connectivity and cladoceran species richness in a metacommunity of shallow lakes

SUMMARY 1. The shallow ponds of the nature reserve 'De Maten' form a metacommunity, in which individual ponds are highly interconnected via a system of overflows and rivulets. This study reports on the relations between cladoceran species richness and (a) connectivity patterns and (b) local environmental variables.
2. No relation was found between local species richness and three connectivity variables or dispersal pathways.
3. Spatial configuration was related to richness, but was confounded by environmental variables for 2 of 3 years. In those 2 years, there was a significant linear relation between Secchi disc depth and species richness, suggesting an important impact of the clearwater/turbid state alternative equilibria in shallow lakes in determining cladoceran richness. Only in the year in which environmental variables were unimportant did connectivity between the ponds influence species richness.
4. These results suggest that local environmental variables related to the clearwater/turbid state alternative equilibria in shallow lakes are important in determining cladoceran species richness. Connectivity and dispersal of individuals between the different ponds only act secondarily by increasing the general species richness within a pond through dispersal from ponds with different environmental conditions.     

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。     

物种多样性和功能群多样性与生态系统生产力的关系——以内蒙古短花针茅草原为例

生物多样性与生态系统生产力之间的关系是当前生态学领域的热点问题。短花针茅(Stipa breviflora)草原是内蒙古荒漠草原的主要类型, 生态系统脆弱, 气候波动剧烈, 研究内蒙古短花针茅草原生物多样性与生产力的关系具有十分重要的意义。该研究在内蒙古短花针茅草原区设置了202个样地进行群落调查, 在干旱区及半干旱区两种资源供给下, 分析了物种丰富度、功能群丰富度与生产力的关系, 旨在解决两个科学问题: 1)物种多样性和功能群多样性中, 哪一种与生产力关系更为密切?2)资源供给对多样性和生产力关系的影响。结果表明: 1)物种丰富度、群落生产力与年降水量呈正相关关系, 而功能群丰富度与年降水量之间不存在显著相关性; 2)群落生产力随物种丰富度的增加而增加, 且两者间呈正线性关系, 功能群丰富度与生产力之间不存在显著相关关系; 3)资源供给会影响多样性与生产力之间的关系, 资源供给低时, 多样性对生产力贡献较低, 资源供给高时, 多样性对生产力的贡献较高。该研究丰富了多样性与生产力关系的研究, 同时, 考虑到植物功能性状的研究在近几年受到生态学家的重视, 且多数研究集中于小尺度的人工控制实验, 因此, 在大尺度自然生态系统中开展功能性状多样性与生态系统功能关系的研究将十分必要。     

On the relationship between plant species diversity and genetic diversity of Plantago lanceolata (Plantaginaceae) within and between grassland communities

Aims and Methods The relationship between genetic diversity and species diversity and the underlying mechanisms are of both fundamental and applied interest. We used amplified fragment length polymorphism (AFLP) and vegetation records to investigate the association between genetic diversity of Plantago lanceolata and plant species diversity using 15 grassland communities in central Germany. We used correlation and partial correlation analyses to examine whether relationships between genetic and species diversity were direct or mediated by environmental differences between habitats.Important findings Both within- and between-population genetic diversity of P. lanceolata were significantly positively correlated with plant species diversity within and between sites. Simple and partial correlations revealed that the positive correlations indirectly resulted from the effects of abiotic habitat characteristics on plant species diversity and, via abundance, on genetic diversity of P. lanceolata. Thus, they did not reflect a direct causal relationship between plant species diversity and genetic diversity of P. lanceolata, as would have been expected based on the hypothesis of a positive relationship between plant species diversity and niche diversity.     

Effects of fertilization and clipping on species diversity,productivity and their relationship in subalpine meadow

施肥和刈割分别对植物群落物种多样性和生产力有重要的影响。采用不同施肥水平和刈割频度, 研究了施肥和刈割对亚高山草甸植物群落物种多样性与生产力及其关系的影响。结果显示, 5年的施肥和刈割处理对群落地上生物量均有显著影响, 但对物种多样性影响不显著; 物种多样性与生产力之间的关系因施肥和刈割次数的不同而不同, 有负相关、二次函数关系和不相关等几种类型, 多数为不相关。此结果表明施肥和刈割并不总是一致性地影响群落物种多样性与生产力。因此, 不同施肥和刈割处理下的亚高山草甸植物群落生产力与物种多样性之间并没有确定的关系。     

Towards a more general species–area relationship: diversity on all islands, great and small

Aim

To demonstrate a new and more general model of the species–area relationship that builds on traditional models, but includes the provision that richness may vary independently of island area on relatively small islands (the small island effect).

Location

We analysed species–area patterns for a broad diversity of insular biotas from aquatic and terrestrial archipelagoes.

Methods

We used breakpoint or piecewise regression methods by adding an additional term (the breakpoint transformation) to traditional species–area models. The resultant, more general, species–area model has three readily interpretable, biologically relevant parameters: (1) the upper limit of the small island effect (SIE), (2) an estimate of richness for relatively small islands and (3) the slope of the species–area relationship (in semi‐log or log–log space) for relatively large islands.

Results

The SIE, albeit of varying magnitude depending on the biotas in question, appeared to be a relatively common feature of the data sets we studied. The upper limit of the SIE tended to be highest for species groups with relatively high resource requirements and low dispersal abilities, and for biotas of more isolated archipelagoes.

Main conclusions

The breakpoint species–area model can be used to test for the significance, and to explore patterns of variation in small island effects, and to estimate slopes of the species–area (semi‐log or log–log) relationship after adjusting for SIE. Moreover, the breakpoint species–area model can be expanded to investigate three fundamentally different realms of the species–area relationship: (1) small islands where species richness varies independent of area, but with idiosyncratic differences among islands and with catastrophic events such as hurricanes, (2) islands beyond the upper limit of SIE where richness varies in a more deterministic and predictable manner with island area and associated, ecological factors and (3) islands large enough to provide the internal geographical isolation (large rivers, mountains and other barriers within islands) necessary for in situ speciation.

     

施肥和刈割对亚高山草甸物种多样性与生产力及其关系的影响

施肥和刈割分别对植物群落物种多样性和生产力有重要的影响。采用不同施肥水平和刈割频度, 研究了施肥和刈割对亚高山草甸植物群落物种多样性与生产力及其关系的影响。结果显示, 5年的施肥和刈割处理对群落地上生物量均有显著影响, 但对物种多样性影响不显著; 物种多样性与生产力之间的关系因施肥和刈割次数的不同而不同, 有负相关、二次函数关系和不相关等几种类型, 多数为不相关。此结果表明施肥和刈割并不总是一致性地影响群落物种多样性与生产力。因此, 不同施肥和刈割处理下的亚高山草甸植物群落生产力与物种多样性之间并没有确定的关系。