高寒草甸植物群落物种多样性和生产力关系的光竞争研究

通过施肥形成的生产力由低到高的过程中,物种多样性往往降低。总体竞争假说认为对所有资源的竞争作用对多样性的影响随着生产力提高而加剧,导致物种多样性的下降;光竞争假说则认为随着生产力提高,种间竞争从低生产力时的地下竞争转向高生产力时的光竞争,是光竞争导致了物种多样性的下降。为了验证这两种假说,本文通过在甘南玛曲高寒草甸的均匀施肥实验,研究了光竞争对高寒草甸植物群落物种多样性和生产力关系的影响。结果表明：(1)随着施肥梯度的增加,大部分植物的生长速率加快,高度和叶面积增加;(2)随着施肥梯度的增加,植物群落地上总的生物量提高,叶面积指数增加,透光率降低,物种多样性减少;(3)个体大小不对称的光竞争导致了高寒草甸植物群落物种多样性随施肥梯度的增加而减少。     

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.     

Biogeography of body size in terrestrial isopods (Crustacea: Oniscidea)

This study tries to unveil the contribution of climatic shift in shaping the extreme body size diversity in terrestrial isopods (Oniscidea). Trying to explain size variation at an interspecific level, we test five hypotheses: (1) Bergmann's Rule and the temperature‐size rule postulate large size in cold areas; (2) The metabolic cold adaptation theory postulates small animal sizes in cold environments; (3) The primary productivity hypothesis predicts size increase in resource‐rich areas; (4) The aridity resistance hypothesis predicts large size in arid regions; and (5). The acidosis hypothesis predicts smaller size with decreasing soil pH. Globally, Bergmann's rule and the aridity hypothesis are weakly supported. Among families and genera, results are variable and idiosyncratic. Conglobating species sizes provide weak support for the acidosis hypothesis. Overall, size is strongly affected by familial affiliation. Isopod size evolution seems to be mainly affected by phylogenetically constrained life‐history traits.     

Relatedness and resource diversity interact to influence the intensity of competition

When resource competition occurs between close relatives, the negative effects of competition are potentially amplified. However, kin selection theory predicts that natural selection should promote the evolution of mechanisms that minimize the intensity of competition between kin. Experimental tests of these hypotheses are mixed, however. Moreover, there is little consensus regarding the generality of either outcome, suggesting that the conditions important in determining the effects of competition between kin are likely complex and not fully understood. We performed two experiments using spadefoot toad tadpoles (Spea multiplicata) to evaluate the hypothesis that individuals can minimize the negative effects of exploitative competition by using alternative resources when competing with close relatives. Supporting our hypothesis, we found that only when individuals had access to alternative resources were the negative effects of competition between siblings less than between unrelated competitors. We suggest that mechanisms to lessen kin competition may be more likely to evolve in environments where alternative resources are available, and that selection to minimize exploitative competition between kin may promote the evolution of resource polyphenism. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 689–695.     

草甸和沼泽植物群落功能多样性与生产力的关系

功能多样性-生产力关系研究结果支持质量比假说和多样性假说, 但对于这两种假说的适用条件尚有争议。通过对吉林省西部草甸和沼泽植物群落的地上生物量、2个物种多样性指标(物种丰富度和Shannon-Weaver指数)、7种植物性状的两类功能多样性指标(群落权重均值和Rao二次熵), 以及土壤环境因子进行调查测量, 研究了群落功能多样性与生产力的关系。结果表明: 1)功能多样性与生产力的关系比物种多样性与生产力的关系更为密切; 2)功能群落权重均值解释生产力变异的能力好于Rao二次熵, 即优势物种对群落生产力的影响作用更大; 3)水淹条件影响着功能多样性与生产力的关系, 以群落权重均值为基础的质量比假说适于解释草甸群落功能多样性与生产力的关系, 而以Rao二次熵为基础的多样性假说适于解释有强烈环境筛(水淹)的沼泽群落功能多样性与生产力的关系。     

Energy, Density, and Constraints to Species Richness: Ant Assemblages along a Productivity Gradient

Species richness describes the number of species of a given taxon in a given time and space. The energy limitation hypothesis links the species richness of consumer taxa to net primary productivity (NPP) through two relationships: NPP limits a taxon's density, and taxon density limits species richness. We study both relationships with a survey of 15 ground ant assemblages, along a productivity gradient from deserts to rain forests. Ant density (colonies m-2) was a positive, decelerating function of net aboveground productivity (NAP). A stepwise regression suggests that the efficiency with which NAP is converted to ant colonies increases with maximum summer temperature and decreases with precipitation. Ant species richness was a positive decelerating function of density at three spatial scales. This supports the energy limitation hypothesis' assumption that average population densities are higher in environments that are more productive. These two nonlinear functions (NAP-density and density-species richness) combine to create, at a variety of scales, positive, decelerating, productivity-diversity curves for a common, ecologically dominant taxon across the terrestrial productivity gradient. However, variance in the density and diversity explained by NAP decreases with scale, suggesting that energy limitation of diversity predominates at small spatial scales (<1 ha).     

Different categories of biodiversity explain productivity variation after fertilization in a Tibetan alpine meadow community

The relationship between productivity and biodiversity has long been an important issue in ecological research. However, in recent decades, most ecologists have primarily focused on species diversity while paying little attention to functional diversity and phylogenetic diversity (PD), especially in alpine meadow communities following fertilization. In this study, a fertilization experiment involving the addition of nitrogen, phosphorus, and a mixture of both was implemented in an alpine meadow on the Tibetan Plateau. Species diversity, functional diversity, and PD were measured, and the responses of these parameters to the variation in productivity were analyzed. We found that the productivity of alpine plant communities was colimited by N and P, with N being the principal and P being the secondary limiting nutrient. Our results supported the prediction of both the mass ratio hypothesis and niche complementarity hypothesis in fertilized communities, but these hypotheses were not mutually exclusive. The combination of different aspects of biodiversity not only provides a crucial tool to explain the variation in productivity and to understand the underlying mechanisms but also plays an important role in predicting the variation in productivity of alpine meadow communities, which are sensitive to nutrient enrichment in the context of global change.     

Relationship between functional diversity and productivity in meadow and marsh plant communities

功能多样性-生产力关系研究结果支持质量比假说和多样性假说, 但对于这两种假说的适用条件尚有争议。通过对吉林省西部草甸和沼泽植物群落的地上生物量、2个物种多样性指标(物种丰富度和Shannon-Weaver指数)、7种植物性状的两类功能多样性指标(群落权重均值和Rao二次熵), 以及土壤环境因子进行调查测量, 研究了群落功能多样性与生产力的关系。结果表明: 1)功能多样性与生产力的关系比物种多样性与生产力的关系更为密切; 2)功能群落权重均值解释生产力变异的能力好于Rao二次熵, 即优势物种对群落生产力的影响作用更大; 3)水淹条件影响着功能多样性与生产力的关系, 以群落权重均值为基础的质量比假说适于解释草甸群落功能多样性与生产力的关系, 而以Rao二次熵为基础的多样性假说适于解释有强烈环境筛(水淹)的沼泽群落功能多样性与生产力的关系。     

Ecological and evolutionary constraints on regional avifauna of passerines in China

Strong correlations between species diversity and climate have been widely observed, but the mechanism underlying this relationship is unclear. Here, we explored the causes of the richness–climate relationships among passerine birds in China by integrating tropical conservatism and diversification rate hypotheses using path models. We found that assemblages with higher species richness southwest of the Salween–Mekong–Pearl River Divide are phylogenetically overdispersed and have shorter mean root distances (MRDs), while species-rich regions northeast of this divide (e.g., north Hengduan Mountains–south Qinling Mountains) are phylogenetically clustered and have longer MRDs. The results of the path analyses showed that the direct effect of climatic factors on species richness was stronger than their indirect effects on species richness via phylogenetic relatedness, indicating that neither tropical conservatism nor diversification rate hypotheses can well explain the richness–climate relationship among passerines in China. However, when path analyses were conducted within subregions separately, we found that the tropical conservatism hypothesis was well supported in the southwestern Salween–Mekong–Pearl River Divide, while the diversification rate hypothesis could explain the richness–climate relationship well in the northeastern divide. We conclude that the diversity patterns of passerines in different subregions of the Eastern Himalayas-Mountains of Southwest China may be shaped by different evolutionary processes related to geological and climatic histories, which explains why the tropical conservatism or diversification rate hypothesis alone cannot fully explain the richness–climate relationships.     

Relationships among soil fertility,native plant diversity and exotic plant abundance inform restoration of forb‐rich eucalypt woodlands

Aim Water and nutrient availability are major limits to productivity in semi‐arid ecosystems; hence, ecological restoration often focuses on conserving or concentrating soil resources. By contrast, nutrient enrichment can promote invasion by exotic annuals, leading to restoration approaches that target reduction of soil nutrients. We aimed to explore potential biodiversity trade‐offs between these approaches by investigating relationships among soil nutrients, exotic annuals and native plant diversity and composition. In particular, we investigated the hypothesis that native plant diversity in semi‐arid to temperate woodlands reflects the productivity–diversity hypothesis, leading to hump‐backed relationships with soil nutrients such that (1) native plant diversity declines with increasing nutrient enrichment and (2) native diversity is limited at the lowest levels of soil fertility. Location Fragmented, long‐ungrazed Eucalyptus loxophleba subsp. loxophleba (York gum)–Acacia acuminata (jam) woodlands in the wheatbelt of South‐Western Australia. Methods We conducted stratified surveys of floristic composition and topsoil nutrient concentrations in 112 woodland patches. We used generalized linear models, structural equation models and ordinations to characterize relationships among soil nutrients, rainfall, exotic annuals and patch‐scale (100 m²) native plant composition and diversity. Results Patch‐scale native plant diversity declined strongly with increasing exotic abundance. This was partly related to elevated soil nutrient concentrations, particularly total nitrogen and available phosphorus. By contrast, there was little evidence for positive correlations between soil nutrients and native diversity, even at very low soil nutrient concentrations. Main conclusions Minimizing weed invasions is crucial for maximizing native plant diversity in E. loxophleba woodlands and could include nutrient‐depleting treatments without substantially compromising the functional capacity of soils to maintain native plant richness and composition. More broadly we emphasize that understanding relationships among ecosystem productivity, plant diversity and exotic invasions in the context of associated theoretical frameworks is fundamental for informing ecological restoration.     

Productivity–diversity patterns in arctic tundra vegetation

Productivity has long been argued to be a major driver of species richness patterns. In the present study we test alternative productivity–diversity hypotheses using vegetation data from the vast Eurasian tundra. The productivity–species pool hypothesis predicts positive relationships at both fine and coarse grain sizes, whereas the productivity–interaction hypothesis predicts unimodal patterns at fine grain size, and monotonic positive patterns at coarse grain size. We furthermore expect to find flatter positive (productivity–species pool hypothesis) or more strongly negative (productivity–interaction hypothesis) relationships for lichens and bryophytes than for vascular plants, because as a group, lichens and bryophytes are better adapted to extreme arctic conditions and more vulnerable to competition for light than the taller‐growing vascular plants. The normalised difference vegetation index (NDVI) was used as a proxy of productivity. The generally unimodal productivity–diversity patterns were most consistent with the productivity–interaction hypothesis. There was a general trend of decreasing species richness from moderately to maximally productive tundra, in agreement with an increasing importance of competitive interactions. High richness of vascular plants and lichens occurred in moderately low productive tundra areas, whereas that of bryophytes occurred in the least productive tundra habitats covered by this study. The fine and coarse grain richness trends were surprisingly uniform and no variation in beta diversity along the productivity gradient was seen for vascular plants or bryophytes. However, lichen beta diversity varied along the productivity gradient, probably reflecting their sensitivity to habitat conditions and biotic interactions. Overall, the results show evidence that productivity–diversity gradients exist in tundra and that these appear to be largely driven by competitive interactions. Our results also imply that climate warming‐driven increases in productivity will strongly affect arctic plant diversity patterns.     

Biomass-diversity responses and spatial dependencies in disturbed tallgrass prairies

Monotonic, hump-shaped and zero-correlation productivity-diversity relationships have been found to date in many ecosystems. This diversity of responses has puzzled ecologists in their search for general principles on ecosystem functioning. Some state that the scale of observation is crucial in defining this relationship. We have developed a spatial model of tallgrass prairies where biomass and litter dynamics are defined by uncoupled difference equations. In this system, we periodically apply prescribed fire as a disturbance that propagates through neighboring cells. The model shows percolation thresholds at points where small-scale spatial heterogeneity and large-scale, global correlation coexist, resulting in power-law distributions in available areas for non-dominant species. These points maximize the biomass-diversity relationship. Our results suggest that spatial dependencies and the disturbance heterogeneity hypothesis are the cornerstone processes accounting for unimodality in productivity-diversity relationships.     

Allelopathy prevents competitive exclusion and promotes phytoplankton biodiversity

It has been hypothesized that allelopathy can prevent competitive exclusion and promote phytoplankton diversity in aquatic ecosystems, where numerous species coexist on a limited number of resources. However, experimental proof‐of‐principle is not available to support this hypothesis. Here we present the first experimental evidence to support this hypothesis by demonstrating that allelopathy promotes the coexistence of two phytoplankton species, Ankistrodesmus falcatus and Oscillatoria sp., that compete for a single limiting nutrient. By performing long‐term competition experiments in nitrate‐limited continuous cultures, and by describing the population dynamics using a mechanistic model, we demonstrate that when allelopathy comes into play, one of the following outcomes is possible depending on the relative initial abundances of the species: dominance of the stronger competitor for nitrate (the non‐allelopathic species), oscillatory coexistence, or dominance of the weaker competitor (the allelopathic species). Our model analysis revealed that sustained oscillatory coexistence of the two species would be a common outcome of this experiment. Our study confirms for the first time, based on laboratory experiments combined with mechanistic models, that allelopathy can alter the predicted outcome of inter‐specific competition in a nutrient‐limited environment and increase the potential for the coexistence of more species than resources, thereby contributing to the identification of endogenous mechanisms that explain the extreme diversity of phytoplankton communities.     

Biodiversity-productivity relations: an experimental evaluation of mechanisms

To examine the mechanisms underlying productivity-diversity relationships, we manipulated nutrient levels in replicate small-scale artificial habitat units in the marine subtidal zone and followed the process of community assembly. In contrast to most enrichment studies, algal diversity increased in enriched habitats relative to controls along with biomass – a result that may be explained by the low nutrient status of the region. Both the total number of faunal species and the total number of individuals were also significantly greater in enriched habitats, but the relationship between algal resources and faunal diversity did not support the resource heterogeneity hypothesis. Received: 14 June 1999 / Accepted: 29 October 1999     

Resource Heterogeneity Moderates the Biodiversity-Function Relationship in Real World Ecosystems

Numerous recent studies have tested the effects of plant, pollinator, and predator diversity on primary productivity, pollination, and consumption, respectively. Many have shown a positive relationship, particularly in controlled experiments, but variability in results has emphasized the context-dependency of these relationships. Complementary resource use may lead to a positive relationship between diversity and these processes, but only when a diverse array of niches is available to be partitioned among species. Therefore, the slope of the diversity-function relationship may change across differing levels of heterogeneity, but empirical evaluations of this pattern are lacking. Here we examine three important functions/properties in different real world (i.e., nonexperimental) ecosystems: plant biomass in German grasslands, parasitism rates across five habitat types in coastal Ecuador, and coffee pollination in agroforestry systems in Indonesia. We use general linear and structural equation modeling to demonstrate that the effect of diversity on these processes is context dependent, such that the slope of this relationship increases in environments where limiting resources (soil nutrients, host insects, and coffee flowers, respectively) are spatially heterogeneous. These real world patterns, combined with previous experiments, suggest that biodiversity may have its greatest impact on the functioning of diverse, naturally heterogeneous ecosystems.     

Separating the influence of resource 'availability' from resource 'imbalance' on productivity–diversity relationships

One of the oldest and richest questions in biology is that of how species diversity is related to the availability of resources that limit the productivity of ecosystems. Researchers from a variety of disciplines have pursued this question from at least three different theoretical perspectives. Species energy theory has argued that the summed quantities of all resources influence species richness by controlling population sizes and the probability of stochastic extinction. Resource ratio theory has argued that the imbalance in the supply of two or more resources, relative to the stoichiometric needs of the competitors, can dictate the strength of competition and, in turn, the diversity of coexisting species. In contrast to these, the field of Biodiversity and Ecosystem Functioning has argued that species diversity acts as an independent variable that controls how efficiently limited resources are utilized and converted into new tissue. Here we propose that all three of these fields give necessary, but not sufficient, conditions to explain productivity–diversity relationships (PDR) in nature. However, when taken collectively, these three paradigms suggest that PDR can be explained by interactions among four distinct, non-interchangeable variables: (i) the overall quantity of limiting resources, (ii) the stoichiometric ratios of different limiting resources, (iii) the summed biomass produced by a group of potential competitors and (iv) the richness of co-occurring species in a local competitive community. We detail a new multivariate hypothesis that outlines one way in which these four variables are directly and indirectly related to one another. We show how the predictions of this model can be fit to patterns of covariation relating the richness and biomass of lake phytoplankton to three biologically essential resources (N, P and light) in a large number of Norwegian lakes.     

草地植物生产力主要影响因素研究综述

草地是全球分布面积最大的陆地生态系统,植物初级生产力是反映草地功能的重要指标。从植物种多样性、资源有效性、放牧、退化草地恢复和气候变化等方面较系统综述了影响草地植物生产力的关键要素和驱动力。大量研究表明,植物多样性与生产力的关系尚未有一致的结论,依据试验地点、起始状态甚至度量指标不同而不同;特别是资源有效性调节着生产力水平并对植物多样性和生产力关系产生显著影响;放牧改变了植物群落特征和养分有效性进而影响生产力的形成过程,也改变了资源有效性-植物多样性-生产力之间的关系;对于退化生态系统,在退化草地恢复过程中植物与土壤资源有效性的互作效应对植物生产力的变化起着关键作用;而在未来气候变化特别是增温对植物生产力的影响因地点和生态系统的不同而异,但多数研究结果显示增温提高了草地植物生产力。与国外其它草地分布区相比,国内的相关研究不仅在数量上明显不足,更重要是欠缺机理上的深入研究。在放牧和未来气候变化背景下如何维持和提高草地生产力,如何加速退化草地生态系统的恢复,进而实现生态安全建设和经济社会协调发展,是我国当前急需解决的理论和实践问题。     

Do biotic interactions shape both sides of the humped‐back model of species richness in plant communities?

A humped-back relationship between species richness and community biomass has frequently been observed in plant communities, at both local and regional scales, although often improperly called a productivity-diversity relationship. Explanations for this relationship have emphasized the role of competitive exclusion, probably because at the time when the relationship was first examined, competition was considered to be the significant biotic filter structuring plant communities. However, over the last 15 years there has been a renewed interest in facilitation and this research has shown a clear link between the role of facilitation in structuring communities and both community biomass and the severity of the environment. Although facilitation may enlarge the realized niche of species and increase community richness in stressful environments, there has only been one previous attempt to revisit the humped-back model of species richness and to include facilitative processes. However, to date, no model has explored whether biotic interactions can potentially shape both sides of the humped-back model for species richness commonly detected in plant communities. Here, we propose a revision of Grime's original model that incorporates a new understanding of the role of facilitative interactions in plant communities. In this revised model, facilitation promotes diversity at medium to high environmental severity levels, by expanding the realized niche of stress-intolerant competitive species into harsh physical conditions. However, when environmental conditions become extremely severe the positive effects of the benefactors wane (as supported by recent research on facilitative interactions in extremely severe environments) and diversity is reduced. Conversely, with decreasing stress along the biomass gradient, facilitation decreases because stress-intolerant species become able to exist away from the canopy of the stress-tolerant species (as proposed by facilitation theory). At the same time competition increases for stress-tolerant species, reducing diversity in the most benign conditions (as proposed by models of competition theory). In this way our inclusion of facilitation into the classic model of plant species diversity and community biomass generates a more powerful and richer predictive framework for understanding the role of plant interactions in changing diversity. We then use our revised model to explain both the observed discrepancies between natural patterns of species richness and community biomass and the results of experimental studies of the impact of biodiversity on the productivity of herbaceous communities. It is clear that explicit consideration of concurrent changes in stress-tolerant and competitive species enhances our capacity to explain and interpret patterns in plant community diversity with respect to environmental severity.     

Explaining the latitudinal gradient anomaly in ichneumonid species richness: evidence from butterflies

1. Despite the profusion of Lepidoptera and other potential hosts in the tropics, the Ichneumonidae are not more species-rich in the lowland tropics than in temperate communities. Alternate but partially complementary hypotheses emphasizing resource fragmentation, predation, or chemical defence have been proposed to explain this paradox.
2. Because different diversity gradients occur in the various subgroups of the Ichneumonidae, as well as among their hosts, general explanations are unlikely to account for this 'anomalous diversity.' Hypotheses of ecological causation are best tested at the taxonomic level where the pattern of interest applies.
3. Phylogenetic trends in the distribution, abundance, and palatability of butterflies (Papilionoidea), a group that increases dramatically in species richness towards the tropics, are presented as a case study for evaluating hypotheses of parasitoid species richness. Evidence is presented that supports the 'nasty host hypothesis' as an explanation for the lack of diversity of associated parasitoids.
4. An examination of hypotheses and the approaches used to test them suggests that the nasty-host hypothesis is best supported by the limited evidence available, while evidence in favour of other hypotheses is either lacking or ambiguous. In particular, use of the koinobiont/idiobiont dichotomy and parasitoid assemblage-size comparisons may result in contradictory conclusions.     

Resource selection,utilization and seasons influence spatial distribution of ungulates in the western Serengeti National Park

Understanding herbivore selection and utilization of vegetation types is fundamental to conservation of multispecies communities. We tested three hypotheses for how ungulate species select their habitats and how this changes with season: first, resources are distributed as a mosaic of patches so that ungulates are also distributed patchily; this distribution reflects habitat selection, which changes with season, the different ungulates behaving differently. Second, resources become scarcer in the dry season relative to those in the wet season. If interspecific competition prevails, then all species should show a contraction of habitats chosen. Third, if predation is limiting, competition will be minimal, and hence, habitat selection by herbivores will not differ between seasons. We used frequencies of occurrence in four common vegetation types in western Serengeti National Park to determine selection coefficients and utilization patterns and Chi‐square analysis to test the hypotheses. The results showed that selection changes differently in each species, agreeing with the first hypothesis. Herbivores did not all become more selective, as predicted by the competition hypothesis, nor did selection remain the same across seasons, as predicted by the predation hypothesis. These results can be useful in constructing habitat suitability maps for ungulate species with special conservation needs.