Plant communities and landscape diversity along a Canadian Arctic river

Abstract. We analysed the structure and diversity of the vegetation along an Arctic river to determine the relationship between species richness and plant community structure. We examined whether variation in species richness along the corridor is structured as (1) an increase in the number of communities due to increasing landscape heterogeneity, (2) an increase in the floristic distinctiveness (β-diversity) of communities, or (3) an increase in within-community richness (α-diversity) as species-poor communities are replaced by species-rich communities. We described 24 community types and analysed the relationship between site vascular species richness (γ-diversity) and β-diversity, α-diversity, site environmental heterogeneity, and the number of distinct plant communities. We also measured diversity patterns of vascular, bryophyte, and lichen species within communities and examined their relationship to community-level estimates of environmental factors. We found that an increase in site species richness correlated with an increase in the number of communities (r²= 0.323, P= 0.0173) and β-diversity (r²= 0.388, P= 0.0075), rather than an increase in the α-diversity of individual communities. Moisture and pH controlled most of the differences in composition between communities. Measures of species richness and correlations with moisture and pH within communities differed among vascular, bryophyte, and lichen species. Bryophyte richness was positively correlated with moisture (r²= 0.862, P= 0.0010) and lichen richness was negatively correlated with moisture (r²= 0.809, P= 0.0031). Vascular plants had a peak in richness at pH 6.5 (r²= 0.214, P < 0.0001). We conclude that site variation in vascular richness in this region is controlled by landscape heterogeneity, and structured as variation in the number and distinctiveness of recognizable plant communities.     

Patterns of diversity in the strata of boreal montane forest in British Columbia

Abstract. Patterns of diversity were analyzed in a boreal coniferous forest and its strata (tree, shrub, herb and bryophyte layers): number of species per community — α-diversity, total species richness — γ-diversity, mean similarity — β-diversity, and mosaic diversity, a measure of complexity. These four measures of diversity consistently decreased from lower to upper vegetation layers. To study the effect of juveniles of larger life forms on diversity of lower layers, they were removed from the data and the measures of diversity reanalyzed. Number of species per community and mosaic diversity decreased substantially, but β-diversity did not change. So, the effect of juveniles on γ-diversity is due to the greater number of species per community. Multiple regression models revealed that the relationships between α-diversity and the environmental variables were the same for the whole forest and for the herb layer. Elevation and soil pH were the major variables explaining α-diversity in the whole community. Climate was the only environmental gradient related to species richness in all individual strata. Tree and herb richness values were negatively related to soil drainage and acidity, respectively. Species richness of the plant community was affected by environmental variability mostly through the herb layer. Various explanations of the observed diversity patterns included: environmental constraints, resource competition, generation time, and colonization processes.     

The richness–heterogeneity relationship differs between heterogeneity measures within and among habitats

The positive monotonic relationship between habitat heterogeneity and species richness is a cornerstone of ecology. Recently, it was suggested that this relationship should be unimodal rather than monotonic due to a tradeoff between environmental heterogeneity and population sizes, which increases local species extinctions at high heterogeneity levels. Here, we studied the richness–heterogeneity relationship for an avian community using two different environmental variables, foliage‐height diversity and cover type diversity. We analyzed the richness–heterogeneity within different habitat types (grasslands, savannas, or woodlands) and at the landscape scale. We found strong evidence that both positive and unimodal relationships exist at the landscape scale. Within habitats we found positive relationships between richness and heterogeneity in grasslands and woodlands, and unimodal relationships in savannas. We suggest that the length of the environmental heterogeneity gradient (which is affected by both spatial scale and the environmental variable being analyzed) affects the type of the richness–heterogeneity relationship. We conclude that the type of the relationship between species richness and environmental heterogeneity is non‐ubiquitous, and varies both within and among habitats and environmental variables.     

Microhabitat amelioration and reduced competition among understorey plants as drivers of facilitation across environmental gradients: Towards a unifying framework

Studies of facilitative interactions as drivers of plant richness along environmental gradients often assume the existence of an overarching stress gradient that equally affects the performance of all the species in a given community. However, co-existing species differ in their ecophysiological adaptations, and do not experience the same stress level under particular environmental conditions. Moreover, these studies assume a unimodal relationship between richness and biomass, which is not as general as previously thought. We ignored these assumptions to assess changes in plant–plant interactions and their effect on local species richness across environmental gradients in semi-arid areas of Spain and Australia. We aimed to understand the relative importance of direct (microhabitat amelioration) and indirect (changes in the competitive relationships among the understorey species: niche segregation, competitive exclusion or intransitivity) mechanisms that might underlie the effects of nurse plants on local species richness. By jointly studying these direct and indirect mechanisms using a unifying framework, we found that nurse plants (trees, shrubs and tussock grasses) increased local richness not only by expanding the niche of neighbouring species but also by increasing niche segregation among them, though the latter was not important in all cases. The outcome of the competition-facilitation continuum varied depending on the study area, likely because the different types of stress gradient considered. When driven by both rainfall and temperature, or rainfall alone, the community-wide importance of nurse plants remained constant (Spanish sites), or showed a unimodal relationship along the gradient (Australian sites). This study expands our understanding of the relative roles of plant–plant interactions and environmental conditions as drivers of local species richness in semi-arid environments. The results can also be used to refine predictions about the response of plant communities to environmental change, and to clarify the relative importance of biotic interactions as drivers of such responses.     

Effects of area,environmental status and environmental variation on species richness per unit area in Mediterranean wetlands

Abstract. We propose a mechanistic model to relate α- and γ-diversity to area per se, moisture status and environmental variation (local and total), and explored the effects these abiotic variables have on species richness per unit area (α-diversity) for plant communities in a network of wetland habitats located in a Mediterranean mountainous region of central Spain. In this study, environmental status is measured as actual evapotranspiration (as an expression of energy), slope and soil wetness, and environmental variation refers to slope variation and soil wetness variation. Species richness per unit area was related to soil wetness, soil wetness variation, ground slope and ground slope variation. There were also positive correlations among moisture status and environmental variation variables. There is a joint effect of slope and soil wetness variation in explaining species richness per unit area of these wetland habitats, but area effects and energy are relatively unimportant. We conclude that species richness per unit area of wetland vegetation can be explained by moisture status and local environmental variation, and that habitat area may not have an important effect. Area could affect γ-diversity directly through random sampling and/or indirectly through increasing β-diversity, and energy may be important in areas with larger energy ranges. Complete surveys of environmental status, local and total environmental variation, and their associated species assemblages are needed to explain the processes that give rise to the rule that larger areas have larger species richness.     

The relationship between species richness and productivity in four typical grasslands of northern China

The relationship between plant species richness and primary productivity has long been acentral topic in biodiversity research.In this paper,we examine the relationship between species richness and productivity in four typical grasslands of Northern China at different spatial scales.At the community scale,a positive correlation was found for six of seven communities.A unimodal pattern was found only for one community (Stipa glareosa community),while at a large scale (vegetation type or landscape/region),the relationship was also found significantly positive.Species richness ranged from 4 to 35 species,and community aboveground productiand aboveground productivity were found in alpine meadow,followed by meadow steppe,typical steppe and desert steppe.     

Microbial leaf degraders in boreal streams: bringing together stochastic and deterministic regulators of community composition

1. Leaves that fall into the water represent a new habitat for microorganisms to colonise in streams, providing an opportunity to study colonisation and the subsequent regulation of community structure. We explored community composition of bacteria and fungi on decomposing alder leaves in nine streams in central Sweden, and describe their relationship with environmental variables. Succession of the microbial community was studied in one of the streams for 118 days. Microbial community composition was examined by denaturing gradient gel electrophoresis on replicate samples of leaves from each stream. 2. During succession in one stream, maximum taxon richness was reached after 34 days for bacteria and 20 days for fungi respectively. Replicate samples within this stream differed between each other earlier in colonisation, while subsequently such variation among replicate communities was low and remained stable for several weeks. Replicate samples taken from all the nine streams after 34 days of succession showed striking similarities in microbial communities within‐streams, although communities differed more strongly between streams. 3. Canonical analysis of microbial communities and environmental variables revealed that water chemistry had a significant influence on community composition. This influence was superimposed on a statistical relationship between the properties of stream catchments and microbial community composition. 4. The catchment regulates microbial communities in two different ways. It harbours the species pool from which the in‐stream microbial community is drawn and it governs stream chemistry and the composition of organic substrates that further shape the communities. We suggest that there is a random element to colonisation early in succession, whereas other factors such as species interactions, stream chemistry and organic substrate properties, result in a more deterministic regulation of communities during later stages.     

Species diversity in vertical, horizontal, and temporal dimensions of a fruit-feeding butterfly community in an Ecuadorian rainforest

To test the hypotheses that fruit-feeding nymphalid butterflies are randomly distributed in space and time, a community of fruit-feeding nymphalid butterflies was sampled at monthly intervals for one year by trapping 6690 individuals of 130 species in the canopy and understory of four forest habitats: primary, higraded, secondary, and edge. The overall species abundance distribution was well described by a lognormal distribution. Total species diversity (γ-diversity) was partitioned into additive components within and among community subdivisions (α-diversity and β-diversity) in vertical, horizontal and temporal dimensions. Although community subdivisions showed high similarity (1 —β-diversity/γ-diversity), significant β-diversity existed in each dimension. Individual abundance and observed species richness was lower in the canopy than in the understory. However, rarefaction analysis and species accumulation curves revealed that canopy had higher species richness than understory. Observed species richness was roughly equal in all habitats, but individual abundance was much greater in edge, largely due to a single, specialist species. Rarefaction analysis and species accumulation curves showed that edge had significantly lower species richness than all other habitats. Samples from a single habitat, height and time contained only a small fraction of the total community species richness. This study demonstrates the feasibility, and necessity, of large-scale, long-term sampling in multiple dimensions for accurately measuring species richness and diversity in tropical forest communities. We discuss the importance of such studies in conservation biology.     

Unimodal diversity-productivity relationship emerged under stressful environment through sampling effect

Researches on the context dependence of biodiversity and ecosystem functioning (BEF) reveal the variation of diversity-productivity relationship (DPR) under stressful environment. The “habitat sampling effect” (HSE) is proposed as the dominant species interaction mechanism at stressful environment, whereas its potential role in driving the DPR has never been testified before. We constructed an individual-based simulation model to explore the variation of DPR along environmental stress gradient, and evaluated the contribution of HSE in explaining this variation. Our results indicated that DPR changed from positive to negative along environmental stress gradients. An unimodal DPR curve emerged at stressful environment, which was caused by the counterbalance of two opposite impacts of HSE on community productivity. At low richness level, the positive impact of HSE on community productivity dominated through the promotion of community size, which resulted in the positive DPR. Whereas with the increase of richness, the negative impact of HSE dominated instead through the reducing of individual productivity, which caused the decreasing part of unimodal curve. Our results highlight the complex characteristic of BEF relationship at stressful environment, and emphasize the necessity of biodiversity in maintaining community's functioning at stressful environment which are often sparse in species richness.     

Altitude effects on spatial components of vascular plant diversity in a subarctic mountain tundra

Environmental gradients are caused by gradual changes in abiotic factors, which affect species abundances and distributions, and are important for the spatial distribution of biodiversity. One prominent environmental gradient is the altitude gradient. Understanding ecological processes associated with altitude gradients may help us to understand the possible effects climate change could have on species communities. We quantified vegetation cover, species richness, species evenness, beta diversity, and spatial patterns of community structure of vascular plants along altitude gradients in a subarctic mountain tundra in northern Sweden. Vascular plant cover and plant species richness showed unimodal relationships with altitude. However, species evenness did not change with altitude, suggesting that no individual species became dominant when species richness declined. Beta diversity also showed a unimodal relationship with altitude, but only for an intermediate spatial scale of 1 km. A lack of relationships with altitude for either patch or landscape scales suggests that any altitude effects on plant spatial heterogeneity occurred on scales larger than individual patches but were not effective across the whole landscape. We observed both nested and modular patterns of community structures, but only the modular patterns corresponded with altitude. Our observations point to biotic regulations of plant communities at high altitudes, but we found both scale dependencies and inconsistent magnitude of the effects of altitude on different diversity components. We urge for further studies evaluating how different factors influence plant communities in high altitude and high latitude environments, as well as studies identifying scale and context dependencies in any such influences.     

Productivity-diversity relationships in lake plankton communities

One of the most intriguing environmental gradients connected with variation in diversity is ecosystem productivity. The role of diversity in ecosystems is pivotal, because species richness can be both a cause and a consequence of primary production. However, the mechanisms behind the varying productivity-diversity relationships (PDR) remain poorly understood. Moreover, large-scale studies on PDR across taxa are urgently needed. Here, we examined the relationships between resource supply and phyto-, bacterio-, and zooplankton richness in 100 small boreal lakes. We studied the PDR locally within the drainage systems and regionally across the systems. Second, we studied the relationships between resource availability, species richness, biomass and resource ratio (N:P) in phytoplankton communities using Structural Equation Modeling (SEM) for testing the multivariate hypothesis of PDR. At the local scale, the PDR showed variable patterns ranging from positive linear and unimodal to negative linear relationships for all planktonic groups. At the regional scale, PDRs were significantly linear and positive for phyto- and zooplankton. Phytoplankton richness and the amount of chlorophyll a showed a positive linear relationship indicating that communities consisting of higher number of species were able to produce higher levels of biomass. According to the SEM, phytoplankton biomass was largely related to resource availability, yet there was a pathway via community richness. Finally, we found that species richness at all trophic levels was correlated with several environmental factors, and was also related to richness at the other trophic levels. This study showed that the PDRs in freshwaters show scale-dependency. We also documented that the PDR complies with the multivariate model showing that plant biomass is not mirroring merely the resource availability, but is also influenced by richness. This highlights the need for conserving diversity in order to maintain ecosystem processes in freshwaters.     

Unifying the relationships of species richness to productivity and disturbance

Although species richness has been hypothesized to be highest at 'intermediate' levels of disturbance, empirical studies have demonstrated that the disturbance-diversity relationship can be either negative or positive depending on productivity On the other hand, hypothesized productivity diversity relationships can be positive, negative or unimodal, as confirmed by empirical studies. However, it has remained unclear under what conditions each pattern is realized, and there is little agreement about the mechanisms that generate these diverse patterns. In this study, I present a model that synthesizes these separately developed hypotheses and shows that the interactive effects of disturbance and productivity on the competitive outcome of multispecies dynamics can result in these diverse relationships of species richness to disturbance and productivity The predicted productivity diversity relationship is unimodal but the productivity level that maximizes species richness increases with increasing disturbance. Similarly, the predicted disturbance diversity relationship is unimodal but the peak moves to higher disturbance levels with increasing productivity Further, these patterns are well explained by the opposite effects of productivity and disturbance on competitive outcome that are suggested by the change in community composition along these two environmental gradients: higher productivity favours superior competitors while higher disturbance levels favour inferior competitors.     

Species richness of tropical wood-inhabiting macrofungi provides support for species-energy theory

A study was undertaken at the El Verde Field Station in Puerto Rico to determine the effect of energy available from newly dead trees on the species richness of macrofungal communities that inhabit them. It is hypothesized that there is a positive relationship between available energy and species richness. Energy was measured using the volume of the dead trees and the wood density of living trees of the same species. One hundred ninety-four logs of known tree species were surveyed 1 y for fruiting bodies of macrofungi at monthly intervals. For individual logs, log volume had a significant positive effect on macrofungal species richness. Younger logs had significantly higher species richness than older logs, and those with less apparent decay had more species than those with more decay. When logs were grouped by tree species, total wood volume and density of live wood had a significant positive effect and average log diameter had a negative effect on total species richness and abundance of the wood-inhabiting macrofungi. Macrofungal richness and abundance constantly increased with initial wood density; there was no evidence for a unimodal relationship. These results support the proposed relationship between species richness and energy.     

Response of reed community to the environment gradient water depth in the Yellow River Delta, China

We investigated and monitored a reed community in the fields.Data on the bio-ecologieal characteristics and β-diversity of reed communities in different environmental gradients (mainly based on water depth) of the Yellow River Delta were collected through multianalysis,extremum analysis and β-diversity index analysis.In accordance with the square sum of deviations (Ward)cluster analysis,10 sampling plots were divided into six types with the dominant plants in different plots varying according to the change in environmental gradients.The dominant plants in these plots varied from aquatic plants to xerophytes and salt tolerant plants as water depth decreased.The average height and diameter of the reeds at breast level were significantly correlated with the average water depth.The fitness curves of average density and coverage with average water depth were nonlinear.When the average water depth was 0.3 m,the average density and coverage of reeds reached the apex value,while the height and diameter of the reeds at breast level increased with the water depth.There were obvious changes to the environmental gradient in the Yellow River Delta.The transitional communities were also found to exist in the Yellow River Delta by β-diversity analysis.Vicarious species appeared with the change in water depth.The occurrence of substitute species is determined by the function of common species between adjacent belts.The different functions of common species led to differences in community structure and function and differences in dominant plants.The result reflects the variations of species present in different habitats and directly reflects environmental heterogeneity.The values of//-diversity indices of adjacent plots were higher than those of nonadjacent plots.There are transition zones between the xerophytes and aquatic plants in the Yellow River Delta.In an aquatic environment,the similarity of reed community is higher than that of xeromorphic plants.The β-diversity index can reflect plant succession trends caused by the change in environmental gradients in the Yellow River Delta.The β-diversity index reveals plant responses to changes in environmental gradient and is helpful in observing changes in patterns of species diversity in relation to environmental gradient change and evolving trends in the future,which in turn plays a prominent role when environmental water requirements of wetland are discussed.     

Herbaceous plant richness and vegetation cover in Mediterranean grasslands and shrublands

Different types of relationship between herbaceous species richness and several parameters indicating abundance of plant material (herbaceous, woody plants, litter and bare ground cover) are presented. The data were obtained from 50 sites along a 300 km strip running from E to W within Spain and Portugal. Each site was representative of the silvo-pastoral landscape of the Mediterranean type ecosystems of the Iberian peninsula, and contained two neighboring patches, one of grassland and the other of shrubland. 3,600 20 × 20 cm subplots were randomly located (72 per site, 36 per patch) crossing the boundary grassland/shrubland. This approach allowed us to analyze the richness-occupation relationship of the space from different points of view: among and within the sites, and among and within the grassland and shrubland plant communities. We found a unimodal relationship between richness-cover similar to the one generally accepted between richness and biomass. Our results show that the dependence of this relationship varies depending on the spatial scale of the analysis and on the type of data used. When the whole region is taken into account, significant unimodal relationships are found between richness and herbaceous cover, litter and bare ground, and a negative linear relationship with woody plant cover. Within the sites there are mainly linear or non-significant relationships. But the results also depend on the type of communities analyzed. In pastures, the unimodal relationship represents the combination of positive and negative linear responses for low and high cover values, respectively. The value for herbaceous cover in which maximum richness occurs is around 60%. In shrublands, this value for cover also corresponds to maximum species richness, although the possibilities of reaching it are limited by other variables, such as woody plant cover. This implies that, on not considering variability at local scale, the relationship is linear and positive. This paper shows the existence of a common model related to herbaceous cover, but this model has multiple controlling factors that act differently in each type of community.     

Comparing Neutral and Trade-off Community Models in Shaping the Community Biomass-Diversity Relationship Under Different Disturbance Levels

Among numerous mechanisms shaping the unimodal relationship between diversity and community biomass, the trade-off model of “CRS” theory is the most famous one. However, recent researches indicate that this relationship may also emerge under the neutral model where all species are identical with each other. By using an individual-based spatially-explicit model, we evaluated the underlying mechanisms shaping this curve for both models under different disturbance levels. We found unimodal relationships emerged for both models at low and medium disturbance levels; the richness for the trade-off community was lower than the neutral community for most of the environment severity levels, especially at the benign environment due to the strong competitive exclusions among species. Whereas under high disturbance level, the positive relationships emerged for both models; both communities had similar richness with their curves nearly overlapped with each other, that is, because the high disturbance intensity strongly decreased the competitive exclusions within the trade-off community. Our results indicate that although the underlying mechanisms are totally different, both models will produce the similar relationship between diversity and community biomass under different disturbance levels.     

The decoupling of abundance and species richness in lizard communities

1. Patterns of species richness often correlate strongly with measures of energy. The more individuals hypothesis (MIH) proposes that this relationship is facilitated by greater resources supporting larger populations, which are less likely to become extinct. Hence, the MIH predicts that community abundance and species richness will be positively related. 2. Recently, Buckley & Jetz (2010, Journal of Animal Ecology, 79, 358-365) documented a decoupling of community abundance and species richness in lizard communities in south-west United States, such that richer communities did not contain more individuals. They predicted, as a consequence of the mechanisms driving the decoupling, a more even distribution of species abundances in species-rich communities, evidenced by a positive relationship between species evenness and species richness. 3. We found a similar decoupling of the relationship between abundance and species richness for lizard communities in semi-arid south-eastern Australia. However, we note that a positive relationship between evenness and richness is expected because of the nature of the indices used. We illustrate this mathematically and empirically using data from both sets of lizard communities. When we used a measure of evenness, which is robust to species richness, there was no relationship between evenness and richness in either data set. 4. For lizard communities in both Australia and the United States, species dominance decreased as species richness increased. Further, with the iterative removal of the first, second and third most dominant species from each community, the relationship between abundance and species richness became increasingly more positive. 5. Our data support the contention that species richness in lizard communities is not directly related to the number of individuals an environment can support. We propose an alternative hypothesis regarding how the decoupling of abundance and richness is accommodated; namely, an inverse relationship between species dominance and species richness, possibly because of ecological release.     

历山自然保护区猪尾沟森林群落植物多样性研究

采用丰富度指数、物种多样性指数和均匀度指数对山西历山自然保护区猪尾沟森林群落多样性进行研究。结果表明 :1 )同一群落内 ,多样性指数存在一定的波动范围 ;不同群落间 ,物种多样性也存在差异 ,但其并不一定具有统计学意义。由此表明 ,群落之间存在差异 ,同时也存在着连续性。 2 )海拔高度是决定本区多样性分布格局的主导因子 ,随着群落分布海拔高度的增加 ,多样性呈一致的上升趋势 ,即多样性与海拔呈正相关关系。 3)群落物种多样性对海拔的敏感性由大到小的次序为草本层 >乔木层 >灌木层 ,其中乔木层的丰富度指数、草本层均匀度指数与海拔有着极显著的正相关关系 ,而乔木层的多样性指数、草本层的丰富度指数与海拔有着极显著的负相关关系 ,灌木层的多样性与海拔没有显著的相关性。 4)群落中不同结构、不同层次对群落总体多样性的贡献是不同的 ,两种测定方法所产生的总体多样性之间呈显著相关关系 ,表明给定加权参数的测定方法没有影响客观生态意义的反映 ,同时也更好地反映出群落结构对于群落多样性的功能差异     

Effects of predator confusion on functional responses

A number of experiments have addressed how increases in nitrogen availability increase the productivity and decrease the diversity of plant communities. We lack, however, a rigorous mechanistic understanding of how changes in abundance of particular species combine to produce changes in community productivity and diversity. Single experiments cannot provide insight into this issue because each species occurs only once per experiment, and each experiment is conducted in only one location; thus, it is impossible from single experiments to determine whether responses of particular species are consistent across environments or dependent on the particular environmental context in which the experiment was conducted. To address this issue, we assembled a dataset of 20 herbaceous species that were each represented in at least 6 different fertilization experiments and tested whether responses were general across experiments. Of the 20 species, one consistently increased in relative abundance and five consistently decreased across replicate experiments. A partially-overlapping group of 8 species displayed responses to nitrogen that varied predictably among experiments as a function of geographic location, neighboring species, or a handful of other community characteristics (ANPP, precipitation, species richness, relative abundance of focal species in control plots, and community composition). Thus, despite modest replication and a limited number of predictor variables, we were able to identify consistent patterns in response of 10 out of 20 species across multiple experiments. We conclude that the responses of individual species to nitrogen addition are often predictable, but that in most cases these responses are functions of the abiotic or biotic environment. Thus, a rigorous understanding of how plant species respond to nitrogen addition will have to consider not only the traits of individual plant species, but also aspects of the communities in which those plants live.