Zooplankton biodiversity and lake trophic state: explanations invoking resource abundance and distribution

While empirical studies linking biodiversity to local environmental gradients have emphasized the importance of lake trophic status (related to primary productivity), theoretical studies have implicated resource spatial heterogeneity and resource relative ratios as mechanisms behind these biodiversity patterns. To test the feasibility of these mechanisms in natural aquatic systems, the biodiversity of crustacean zooplankton communities along gradients of total phosphorus (TP) as well as the vertical heterogeneity and relative abundance of their phytoplankton resources were assessed in 18 lakes in Quebec, Canada. Zooplankton community richness was regressed against TP, the spatial distribution of phytoplankton spectral groups, and the relative biomass of spectral groups. Since species richness does not adequately capture ecological function and life history of different taxa, features which are important for mechanistic theories, relationships between zooplankton functional diversity (FD) and resource conditions were examined. Zooplankton species richness showed the previously established tendency to a unimodal relationship with TP, but functional diversity declined linearly over the same gradient. Changes in zooplankton functional diversity could be attributed to changes in both the spatial distribution and type of phytoplankton resource. In the studied lakes, spatial heterogeneity of phytoplankton groups declined with TP, even while biomass of all groups increased. Zooplankton functional diversity was positively related to increased heterogeneity in cyanobacteria spatial distribution. However, a smaller amount of variation in functional diversity was also positively related to the ratio of biomass in diatoms/chrysophytes to cyanobacteria. In all observed relationships, a greater variation of functional diversity than species richness measures was explained by measured factors, suggesting that functional measures of zooplankton communities will benefit ecological research attempting to identify mechanisms behind environmental gradients affecting diversity.     

全尺度人工湿地中植物多样性对生产力与多样性效应的影响

在中国东南部的全尺度复合垂直流人工湿地中开展2年的植物多样性实验,以研究植物多样性（包括植物物种丰富度和植物组成）对群落生产力与多样性效应（即互补效应、选择效应和净多样性效应）的影响及其产生机制。结果表明,2007年物种丰富度与群落生产力呈线形正相关,而2008年显著的单峰格局,其关系式为：y=-0.213x2＋3.455x＋15.192（R=0.215）。2008年物种丰富度与互补效应呈显著地线形负相关,而2007年呈单峰格局,其关系式为：y=-0.389x2＋6.974x-10.707（R=0.247）,而且2007年与2008年的互补效应与生产力都呈显著的正相关,表明互补效应对生产力的提高有重要作用。然而,2007年与2008年物种丰富度与选择效应之间均没有显著相关性,且选择效应与群落生产力之间也没有显著相关性,表明选择效应对生产力的提高作用不显著。2007年与2008年中物种组成对生产力、互补效应、选择效应与净多样性效应均有显著影响,说明人工湿地的植物配置对其生态系统功能的维持尤为重要。2008年物种丰富度与净多样性效应呈极显著地线形负相关,而2007年呈显著单峰格局,其关系式为：y=-0.329 x2＋5.968 x-12.659（R=0.234）,这种趋势主要是由于植物多样性-生态系统功能关系的影响因素（如物种的竞争力和生态位）在2年中有所变化。同时,2007年与2008年的多样性净效应与生产力都呈显著正相关关系,表明生产力与多样性净效应的变化趋势是同步的。与抽样效应假说不同的是,本实验中单种最高产物种（芦竹）在混种时没有表现出高产,主要是由于生长的分配、资源的竞争力与环境的变化等。     

Plankton biodiversity along a gradient of productivity and its mediation by macrophytes

We studied the effect of aquatic vegetation on the process of species sorting and community assembly of three functional groups of plankton organisms (phytoplankton, seston-feeding zooplankton, and substrate-dwelling zooplankton) along a primary productivity gradient. We performed an outdoor cattle tank experiment (n = 60) making an orthogonal combination of a primary productivity gradient (four nutrient addition levels: 0, 10, 100, and 1000 microg P/L; N/P ratio: 16) with a vegetation gradient (no macrophytes, artificial macrophytes, and real Elodea nuttallii). We used artificial plants to evaluate the mere effects of plant physical structure independently from other plant effects, such as competition for nutrients or allelopathy. The tanks were inoculated with species-rich mixtures of phytoplankton and zooplankton. Both productivity and macrophytes affected community structure and diversity of the three functional groups. Taxon richness declined with increasing plankton productivity in each functional group according to a nested subset pattern. We found no evidence for unimodal diversity-productivity relationships. The proportional abundance of Daphnia and of colonial Scenedesmus increased strongly with productivity. GLM analyses suggest that the decline in richness of seston feeders was due to competitive exclusion by Daphnia at high productivity. The decline in richness of phytoplankton was probably caused by high Daphnia grazing. However, partial analyses indicate that these explanations do not entirely explain the patterns. Possibly, environmental deterioration associated with high productivity (e.g., high pH) was also responsible for the observed richness decline. Macrophytes had positive effects on the taxon richness of all three functional plankton groups and interacted with the initial productivity gradient in determining their communities. Macrophytes affected the composition and diversity of the three functional groups both by their physical structure and through other mechanisms. Part of the macrophyte effect may be indirect via a reduction of phytoplankton production. Our results also indirectly suggest that the often reported unimodal relationship between primary productivity and diversity in nature may be partially mediated by the tendency of submerged macrophytes to be most abundant at intermediate productivity levels.     

Contrasting plant productivity-diversity relationships across latitude: the role of evolutionary history

The relationship between net primary productivity and biological diversity has been a central topic in ecology for several decades. The unimodal ("hump-back") relationship has been the most widely accepted for plants with the decrease in diversity at high productivity usually attributed to competitive exclusion. However, the relatively small species pool size under high productivity conditions may account for this pattern as well. Small species pool sizes for highly productive habitats are characteristic of temperate regions, where productive habitats for speciation and species migration have historically been rare. In contrast, productive habitats in the tropics have been relatively common during evolutionary history, resulting in large species pools. We hypothesize that evolutionary history contributes to the observed productivity-diversity relationship of plants, and that the productivity-diversity relationship differs between temperate and tropical regions. We investigated the productivity-diversity relationship patterns from 163 case studies throughout the world. Latitude described approximately 80% of the variation in the shape of the relationships. The unimodal relationship was found to dominate in the temperate zone, whereas the positive relationship was significantly more common in the tropics. We detected no influence due to methods of productivity measurement, but unimodal or positive productivity-diversity relationships were more likely within larger ranges of productivity. The length of the productivity gradient did not affect the latitudinal influence. In summary, the shape of the productivity-diversity relationship differs between temperate and tropical regions and the different evolutionary history of the local species pools is a probable cause for the difference.     

Species richness patterns along environmental gradients in island land molluscan fauna

The relationship between species richness and environmental variables may change depending on habitat structure, dispersal ability, species mixing, and community adaptation to the environment. It is crucial to know how these factors regulate the environment-diversity relationship. The land molluscan fauna of the Ogasawara Islands in the West Pacific is an excellent model system to address this question because of the high species endemicity (> 90%), small area, and simple habitat structure of the islands. I examined relationships among indigenous species composition, richness, and habitat condition, and especially productivity and forest moisture on the island of Anijima. Two major communities of snails could be distinguished by detrended correspondence analysis (DCA): one group dominated in a moist habitat with high productivity, and the other group dominated in a dry habitat with low productivity. However, species richness became highest at the intermediate condition between the habitats in which the two snail communities were dominant, so that species richness showed a hump-shaped relationship with moisture and productivity. In contrast, the species richness of the snail community in the moist habitat showed a monotonically positive correlation, and that in the dry habitat showed a monotonically negative correlation with moisture and productivity. Thus, the greater species richness in intermediate moisture and productivity resulted from the ecotone effect or community overlap at the transitional areas, where faunas with different ecologies can meet in a single site. These findings suggest that hump-shaped productivity-diversity relationships in land Mollusca would reflect the ecotone effect as a result of the mixing of species adapted to either fertile habitats or sterile habitats.     

Spatial scale and species identity influence the indigenous-alien diversity relationship in springtails

Although theory underlying the invasion paradox, or the change in the relationship between the richness of alien and indigenous species from negative to positive with increasing spatial scale, is well developed and much empirical work on the subject has been undertaken, most of the latter has concerned plants and to a lesser extent marine invertebrates. Here we therefore examine the extent to which the relationships between indigenous and alien species richness change from the local metacommunity to the interaction neighborhood scales, and the influences of abundance, species identity, and environmental favorability thereon, in springtails, a significant component of the soil fauna. Using a suite of modeling techniques, including generalized least squares and geographically weighted regressions to account for spatial autocorrelation or nonstationarity of the data, we show that the abundance and species richness of both indigenous and alien species at the metacommunity scale respond strongly to declining environmental favorability, represented here by altitude. Consequently, alien and indigenous diversity covary positively at this scale. By contrast, relationships are more complex at the interaction neighborhood scale, with the relationship among alien species richness and/or density and the density of indigenous species varying between habitats, being negative in some, but positive in others. Additional analyses demonstrated a strong influence of species identity, with negative relationships identified at the interaction neighborhood scale involving alien hypogastrurid springtails, a group known from elsewhere to have negative effects on indigenous species in areas where they have been introduced. By contrast, diversity relationships were positive with the other alien species. These results are consistent with both theory and previous empirical findings for other taxa, that interactions among indigenous and alien species change substantially with spatial scale and that environmental favorability may play a key role in explaining the larger scale patterns. However, they also suggest that the interactions may be affected by the identity of the species concerned, especially at the interaction neighborhood scale.     

地形因子对暖温带森林群落物种丰富度-地上生物量关系的影响

地形因子是森林群落物种丰富度与生产力的重要影响因素,可以修正物种丰富度-生产力两者之间的关系。为了厘清地形因子对物种丰富度-生产力关系的修正效应,以东灵山暖温带天然次生林为研究对象,基于20 hm2样地DEM地图,提取海拔、坡度、坡向、凹凸度等地形因子,用GLM模型探讨了地形因子对群落丰富度与地上生物量的影响。结果表明,群落地上生物量与物种丰富度分别受不同地形因子的影响,群落地上生物量受Ea、El2、El3、Co34个因子的显著影响（P〈0.05）,而物种丰富度则受Sl、El、El2、El34个因子的显著作用（P〈0.05）,两者受海拔相关因子的影响最大,分别解释了两者3.99%和11.09%的变异。地形因子修正了物种丰富度-地上生物量之间的单峰曲线关系,使其朝着单调与直线化趋势变化。以后在探讨森林群落物种丰富度-生产力关系时,地形因子的修正作用应该受到重视。     

Effects of productivity on biodiversity in forest ecosystems across the United States and China

In the global campaign against biodiversity loss in forest ecosystems, land managers need to know the status of forest biodiversity, but practical guidelines for conserving biodiversity in forest management are lacking. A major obstacle is the incomplete understanding of the relationship between site primary productivity and plant diversity, due to insufficient ecosystem‐wide data, especially for taxonomically and structurally diverse forest ecosystems. We investigated the effects of site productivity (the site's inherent capacity to grow timber) on tree species richness across 19 types of forest ecosystems in North America and China through 3 ground‐sourced forest inventory data sets (U.S. Forest Inventory and Analysis, Cooperative Alaska Forest Inventory, and Chinese Forest Management Planning Inventory). All forest types conformed to a consistent and highly significant (P < 0.001) hump‐shaped unimodal relationship, of which the generalized coefficients of determination averaged 20.5% over all the forest types. That is, tree species richness first increased as productivity increased at a progressively slower rate, and, after reaching a maximum, richness started to decline. Our consistent findings suggest that forests of high productivity would sustain few species because they consist mostly of flat homogeneous areas lacking an environmental gradient along which a diversity of species with different habitats can coexist. The consistency of the productivity–biodiversity relationship among the 3 data sets we examined makes it possible to quantify the expected tree species richness that a forest stand is capable of sustaining, and a comparison between the actual species richness and the sustainable values can be useful in prioritizing conservation efforts.     

Diversity-invasibility across an experimental disturbance gradient in Appalachian Forests

Research examining the relationship between community diversity and invasions by nonnative species has raised new questions about the theory and management of biological invasions. Ecological theory predicts, and small-scale experiments confirm, lower levels of nonnative species invasion into species-rich compared to species-poor communities, but observational studies across a wider range of scales often report positive relationships between native and nonnative species richness. This paradox has been attributed to the scale dependency of diversity-invasibility relationships and to differences between experimental and observational studies. Disturbance is widely recognized as an important factor determining invasibility of communities, but few studies have investigated the relative and interactive roles of diversity and disturbance on nonnative species invasion. Here, we report how the relationship between native and nonnative plant species richness responded to an experimentally applied disturbance gradient (from no disturbance up to clearcut) in oak-dominated forests. We consider whether results are consistent with various explanations of diversity-invasibility relationships including biotic resistance, resource availability, and the potential effects of scale (1 m2 to 2 ha). We found no correlation between native and nonnative species richness before disturbance except at the largest spatial scale, but a positive relationship after disturbance across scales and levels of disturbance. Post-disturbance richness of both native and nonnative species was positively correlated with disturbance intensity and with variability of residual basal area of trees. These results suggest that more nonnative plants may invade species-rich communities compared to species-poor communities following disturbance.     

The role of local and landscape level measures of greenness in modelling boreal plant species richness

Measurements of primary productivity and its heterogeneity based on satellite images can provide useful estimates of species richness and distribution patterns. However, species richness at a given site may depend not only on local habitat quality and productivity but also on the characteristics of the surrounding landscape. In this study we investigated whether the predictions of species richness of plant families in northern boreal landscape in Finland can be improved by incorporating greenness information from the surrounding landscape, as derived from remotely sensed data (mean, maximum, standard deviation and range values of NDVI derived from Landsat ETM), into local greenness models. Using plant species richness data of 28 plant families from 440 grid cells of 25 ha in size, generalized additive models (GAMs) were fitted into three different sets of explanatory variables: (1) local greenness only, (2) landscape greenness only, and (3) combined local and landscape greenness. The derived richness–greenness relationships were mainly unimodal or positively increasing but varied between different plant families, and depended also on whether greenness was measured as mean or maximum greenness. Incorporation of landscape level greenness variables improved significantly both the explanatory power and cross-validation statistics of the models including only local greenness variables. Landscape greenness information derived from remote sensing data integrated with local information has thus the potentiality to improve predictive assessments of species richness over extensive and inaccessible areas, especially in high-latitude landscapes. Overall, the significant relationship between plants and surrounding landscape quality detected here suggests that landscape factors should be considered in preserving species richness of boreal environments, as well as in conservation planning for biodiversity in other environments.     

Woody encroachment decreases diversity across North American grasslands and savannas

Woody encroachment is a widespread and acute phenomenon affecting grasslands and savannas worldwide. We performed a meta-analysis of 29 studies from 13 different grassland/savanna communities in North America to determine the consequences of woody encroachment on plant species richness. In all 13 communities, species richness declined with woody plant encroachment (average decline = 45%). Species richness declined more in communities with higher precipitation (r2 = 0.81) and where encroachment was associated with a greater change in annual net primary productivity (ANPP; r2 = 0.69). Based on the strong positive correlation between precipitation and ANPP following encroachment (r2 = 0.87), we hypothesize that these relationships occur because water-limited woody plants experience a greater physiological and demographic release as precipitation increases. The observed relationship between species richness and ANPP provides support for the theoretical expectation that a trade-off occurs between richness and productivity in herbaceous communities. We conclude that woody plant encroachment leads to significant declines in species richness in North American grassland/savanna communities.     

Metacommunity models predict the local-regional species richness relationship in a natural system

Many natural communities exhibit positive relationships between local and regional species richness (LSR-RSR relationships), which can be either linear or curvilinear. Previous models have shown that the form of this relationship depends on the relative rates of colonization and extinction and the sensitivity of these rates to competition. We use simple models to show that the LSR-RSR relationship also depends on the type of metacommunity structure (Levins-like or mainland-island), and our models generate a wider range of realistic forms than do most previous models. We parameterize and test our models with two independent data sets for Daphnia in rock pools on islands in Finland and Sweden. We find that the Levins-like model with competition correctly predicts the observed LSR-RSR relationship and provides the best fit to the average local species richness per island. Simulations show that our models are robust to relaxing our assumption of identical species properties. Our study is one of the first to make and successfully test quantitative predictions for how a widely studied community pattern, the LSR-RSR relationship, arises from metacommunity dynamics.     

Linking richness, community variability, and invasion resistance with patch size

The influence of community dynamics on the success or failure of an invasion is of considerable interest. What has not been explored is the influence of patch size on the outcomes of invasions for communities with the same species pool. Here we use an empirically validated spatial model of a marine epibenthic community to examine the effects of patch size on community variability, species richness, invasion, and the relationships between these variables. We found that the qualitative form of the relationship between community variability and species richness is determined by the size of the model patch. In small patches, variability decreases with species richness, but beyond a critical patch size, variability increases with increasing richness. This occurs because in large patches large, long-lived colonies attain sufficient size to minimize mortality and dominate the community, leading to decreased species richness and community variability. This mechanism cannot operate on smaller patches where the size of colonies is limited by the patch size and mortality is high irrespective of species identity. Further, invasion resistance is strongly correlated with community variability. Thus, the relationship between species richness and invasion resistance is also determined by patch size. These patterns are generated largely by an inverse relationship between colony size and mortality, and they depend on the spatial nature and patch size of the community. Our results suggest that a continuum of possible relationships can exist between species richness, community variability, invasion resistance, and area. These relationships are emergent behaviors generated by the individual properties of the particular component species of a community.     

A unified model of avian species richness on islands and continents

How many species in a given taxon should be found in a delimited area in a specified place in the world? Some recent literature suggests that the answer to this question depends strongly on the geographical, evolutionary, and ecological context. For example, current theory suggests that species accumulate as a function of area differently on continents and islands. Species richness-climate relationships have been examined separately on continents and on islands. This study tests the hypotheses that (1) the functional relationship between richness and climate is the same on continents and islands; (2) the species-area slope depends on distance-based isolation; (3) species-area relationships differ among land bridge islands, oceanic islands, and continents; (4) richness differs among biogeographic regions independently of climate and isolation. We related bird species numbers in a worldwide sample of 240 continental parcels and 346 islands to several environmental variables. We found that breeding bird richness varies similarly on islands and on continents as a function of mean annual temperature, an area x precipitation interaction, and the distance separating insular samples from the nearest continent (R2 = 0.86). Most studies to date have postulated that the slope of the species-area relationship depends upon isolation. In contrast, we found no such interaction. A richness-environment relationship derived using Old World sites accurately predicts patterns of richness in the New World and vice versa (R2 = 0.85). Our results suggest that most of the global variation in richness is not strongly context-specific; rather, it reflects a small number of general environmental constraints operating on both continents and islands.     

Scale-Dependent Effects of Habitat Disturbance on Species Richness in Tropical Forests

Abstract: Despite growing concern, no consensus has emerged over the effects of habitat modification on species diversity in tropical forests. Even for comparatively well-studied taxa such as Lepidoptera, disturbance has been reported to increase and decrease diversity with approximately equal frequency. Species diversity within landscapes depends on the spatial scale at which communities are sampled, and the effects of disturbance in tropical forests have been studied at a wide range of spatial scales. Yet the question of how disturbance affects diversity at different spatial scales has not been addressed. We reanalyzed data from previous studies to examine the relationship between spatial scale and effects of disturbance on tropical-forest Lepidoptera. Disturbance had opposite effects on diversity at large and small scales: as scale decreased, the probability of a positive effect of disturbance on diversity increased. We also explicitly examined the relationship between spatial scale and the diversity of butterflies in selectively logged and unlogged forest in Maluku Province, Indonesia. Species richness increased with spatial scale in both logged and unlogged forest, but at a significantly faster rate in unlogged forest, whereas species evenness increased with scale in unlogged forest but did not increase with scale in logged forest. These data indicate that the effects of habitat modification on species diversity are heavily scale-dependent. As a result, recorded effects of disturbance were strongly influenced by the spatial scale at which species assemblages were sampled. Future studies need to account for this by explicitly examining the effects of disturbance at a number of different spatial scales. A further problem arises because the relationship between scale and diversity is likely to differ among taxa in relation to mobility. This may explain to some extent why the measured effects of disturbance have differed between relatively mobile and immobile taxa.     

Landscape connectivity strengthens local-regional richness relationships in successional plant communities

Local species diversity is maintained over ecological time by a balance between dispersal and species interactions. Local-regional species richness relationships are often used to investigate the relative importance of these two processes and the scales at which they operate. For communities undergoing succession, theory predicts a temporal progression in local-regional species richness relationships: from no relationship to positive linear to saturating. However, observational tests have been mixed, and experiments have been rare. Using a replicated large-scale experiment, we evaluate the impact of two dispersal-governing processes at the regional scale, connectivity and shape of the region (i.e., patches), on the progression of local-regional species richness relationships for plant communities undergoing succession. Regional connectivity accelerates the transition from no relationship to a positive linear relationship, while the shape of the region has no consistent effect nine years post-disturbance. Our results experimentally demonstrate the importance of dispersal in structuring local-regional species richness relationships over time and suggest that conservation corridors among regions can increase local diversity through regional enrichment of plant communities undergoing reassembly.     

Bird metacommunities in temperate South American forest: vegetation structure, area, and climate effects

Spatial structure in metacommunities and their relationships to environmental gradients have been linked to opposing theories of community assembly. In particular, while the species sorting hypothesis predicts strong environmental influences, the neutral theory, the mass effect, and the patch dynamics frameworks all predict differing degrees of spatial structure resulting from dispersal and competition limitations. Here we study the relative influence of environmental gradients and spatial structure in bird assemblages of the Chilean temperate forest. We carried out bird and vegetation surveys in South American temperate forests at 147 points located in nine different protected areas in central Chile, and collected meteorological and productivity data for these localities. Species composition dissimilarities between sites were calculated, as well as three indices of bird local diversity: observed species richness, Chao estimate of richness, and Shannon diversity. A stepwise multiple regression and partial regression analyses were used to select a small number of environmental factors that predicted bird species diversity. Although diversity indices were spatially autocorrelated, environmental factors were sufficient to account for this autocorrelation. Moreover, community dissimilarities were not significantly related to distance between sites. We then tested a multivariate hypothesis about climate, vegetation, and avian diversity interactions using a structural equation modeling (SEM) approach. The SEM showed that climate and area of fragments have important indirect effects on avian diversity, mediated through changes in vegetation structure. Given the scale of this study, the metacommunity framework provides useful insights into the mechanisms driving bird assemblages in this region. Taken together, the weak spatial structure of community composition and diversity, as well as the strong environmental effects on bird diversity, support the interpretation that species sorting has a predominant role in structuring avian assemblages in the region.     

Species richness and stand stability in conifer forests of the Sierra Nevada

Theoretical and empirical studies have long suggested that stability and complexity are intimately related, but evidence from long-lived systems at large scales is lacking. Stability can either be driven by complex species interactions, or it can be driven by the presence/absence and abundance of a species best able to perform a specific ecosystem function. We use 64 years of stand productivity measures in forest systems composed of four dominant conifer tree species to contrast the effect of species richness and abundance on three stability measures. To perform this contrast, we measured the annual growth increments of > 900 trees in mixed and pure forest stands to test three hypotheses: increased species richness will (1) decrease stand variance, (2) increase stand resistance to drought events, and (3) increase stand resilience to drought events. In each case, the alternate hypothesis was that species richness had no effect, but that species composition and abundance within a stand drove variance, resistance, and resilience. In pure stands, the four species demonstrated significant differences in productivity, and in their resistance and resilience to drought events. The two pine species were the most drought resistant and resilient, whereas mountain hemlock was the least resistant and resilient, and red fir was intermediate. For community measures we found a moderately significant (P = 0.08) increase in the community coefficient of variation and a significant (P = 0.03) increase in resilience with increased species richness, but no significant relationship between species richness and community resistance, though the variance in community resistance to drought decreased with species richness. Community resistance to drought was significantly (P = 0.001) correlated to the relative abundance of lodgepole pine, the most resistant species. We propose that resistance is driven by competition for a single limiting resource, with negative diversity effects. In contrast resilience measures the capacity of communities to partition resources in the absence of a single limiting resource, demonstrating positive diversity effects.     

Modifying native and exotic species richness correlations: the influence of fire and seed addition.

An important goal in restoration is to increase the richness of native species while reducing exotic species. However, native species richness is often positively correlated with exotic species richness. In a grassland-savanna system in Michigan (USA), we show that management that focuses on changing the nature of the exotic-native richness relationship can be used to restore native communities. Native and exotic species richnesses were positively correlated, likely due to a shared coupling with aboveground live biomass (a surrogate for productivity). The addition of native seed shifted the exotic-native richness relationship from a linear positive to a monotonic relationship: in areas of intermediate levels of exotic species richness, seed addition increased native diversity without an associated effect on exotic diversity, but in areas of high or low exotic richness, it did not affect native species richness. Prescribed burning broke the correlation between native and exotic richness with no consistent effect on the richness of either group. However, when burning was combined with native-seed addition, the relationship between native and exotic richness was maintained and was shifted upwards, enhancing native recruitment. Although aboveground productivity was strongly related to species richness across the landscape, changes in productivity did not drive these shifts.     

Small-scale grassland assembly patterns differ above and below the soil surface

The existence of deterministic assembly rules for plant communities remains an important and unresolved topic in ecology. Most studies examining community assembly have sampled aboveground species diversity and composition. However, plants also coexist belowground, and many coexistence theories invoke belowground competition as an explanation for aboveground patterns. We used next-generation sequencing that enables the identification of roots and rhizomes from mixed-species samples to measure coexisting species at small scales in temperate grasslands. We used comparable data from above (conventional methods) and below (molecular techniques) the soil surface (0.1 x 0.1 x 0.1 m volume). To detect evidence for nonrandom patterns in the direction of biotic or abiotic assembly processes, we used three assembly rules tests (richness variance, guild proportionality, and species co-occurrence indices) as well as pairwise association tests. We found support for biotic assembly rules aboveground, with lower variance in species richness than expected and more negative species associations. Belowground plant communities were structured more by abiotic processes, with greater variability in richness and guild proportionality than expected. Belowground assembly is largely driven by abiotic processes, with little evidence for competition-driven assembly, and this has implications for plant coexistence theories that are based on competition for soil resources.