Biotic interactions limit species richness in an alpine plant community,especially under experimental warming

The determinants of local species richness in plant communities have been the subject of much debate. Is species richness the result of stochastic events such as dispersal processes, or do local environmental filters sort species into communities according to their ecological niches? Recent studies suggest that these two processes simultaneously limit species richness, although their relative importance may vary in space and time. Understanding the limiting factors for species richness is especially important in light of the ongoing global warming, as new species establish in resident plant communities as a result of climate‐driven migration. We examined the relative importance of dispersal and environmental filtering during seedling recruitment and plant establishment in an alpine plant community subjected to seed addition and long‐term experimental warming. Seed addition increased species richness during the seedling recruitment stage, but this initial increase was cancelled out by a corresponding decrease in species richness during plant establishment, suggesting that environmental filters limit local species richness in the long term. While initial recruitment success of the sown species was related to both abiotic and biotic factors, long‐term establishment was controlled mainly by biotic factors, indicating an increase in the relative importance of biotic interactions once plants have germinated in a microhabitat with favourable abiotic conditions. The relative importance of biotic interactions also seemed to increase with experimental warming, suggesting that increased competition within the resident vegetation may decrease community invasibility as the climate warms.     

Spatial variation in seed limitation of plant species richness and population sizes in floodplain tallgrass prairie

The relative importance of seed availability versus biotic interactions that affect early life stages in limiting plant population sizes and determining composition of plant communities is a central debate in plant ecology. We conducted a seed addition experiment in restored tallgrass prairie in central Kansas to determine (1) whether addition of seed of 18 native forb species produced persistent (three growing seasons) increases in the species' population sizes and plant species richness, (2) what properties of recipient communities best explained spatial variation in added species' establishment, and (3) whether seed size explained interspecific patterns in establishment success. Adding seed led to persistent increases in the number of added species present and in plant species richness at one of three sites. Increased species richness at the one site where community composition was structured by seed availability largely resulted from greater densities of four species. Seed size did not predict species' establishment success. Pre-existing plant species richness was correlated with added species' establishment success, but the direction of the relationship (positive vs. negative) varied among sites. Living aboveground plant biomass in experimental plots in the year of seed addition was negatively correlated with the number of added species established three years later. Our results provide further evidence for large spatial variation in seed limitation of plant community composition. Surprisingly, mean light availability and heterogeneity in light, both important parameters in conceptual models of grassland plant coexistence, did not predict the response of the recipient plant community to seed addition as well as pre-existing plant species richness and living aboveground biomass.     

Responses of Plant Community Composition and Biomass Production to Warming and Nitrogen Deposition in a Temperate Meadow Ecosystem

Climate change has profound influences on plant community composition and ecosystem functions. However, its effects on plant community composition and biomass production are not well understood. A four-year field experiment was conducted to examine the effects of warming, nitrogen (N) addition, and their interactions on plant community composition and biomass production in a temperate meadow ecosystem in northeast China. Experimental warming had no significant effect on plant species richness, evenness, and diversity, while N addition highly reduced the species richness and diversity. Warming tended to reduce the importance value of graminoid species but increased the value of forbs, while N addition had the opposite effect. Warming tended to increase the belowground biomass, but had an opposite tendency to decrease the aboveground biomass. The influences of warming on aboveground production were dependent upon precipitation. Experimental warming had little effect on aboveground biomass in the years with higher precipitation, but significantly suppressed aboveground biomass in dry years. Our results suggest that warming had indirect effects on plant production via its effect on the water availability. Nitrogen addition significantly increased above- and below-ground production, suggesting that N is one of the most important limiting factors determining plant productivity in the studied meadow steppe. Significant interactive effects of warming plus N addition on belowground biomass were also detected. Our observations revealed that environmental changes (warming and N deposition) play significant roles in regulating plant community composition and biomass production in temperate meadow steppe ecosystem in northeast China.     

The relative role of dispersal and local interactions for alpine plant community diversity under simulated climate warming

Most studies on factors determining diversity are conducted in temperate or warm regions, whereas studies in climatically harsh and low productivity areas, such as alpine regions, are rare. We examined the relative roles of seed availability and different biotic and abiotic factors for the diversity of an alpine plant community in southern Norway. Furthermore, because climate warming is predicted to be an important driver of alpine species diversity, we assessed how the relative impacts of dispersal and local interactions on diversity might change under experimental warming (open top chambers, OTCs).
Addition of seeds from 27 regional species increased community diversity. The establishment of the species was negatively related both to the diversity of the existing system and the cover of the abundant dwarf shrub Dryas octopetala . These results show that both species dispersal limitation and local biotic interactions are important factors for alpine plant community diversity. Despite relatively harsh environmental conditions and low productivity, competition from the resident vegetation appeared to have a greater role for species establishment and diversity than facilitation and experimental warming. Higher temperature appeared to increase the negative relationship between resident species diversity and species establishment. This may suggest that climate warming can increase the role of interspecific competition for alpine plant community structure, and thus alter the long-term effects of biotic interactions on diversity.     

Warming increases plant biomass and reduces diversity across continents,latitudes, and species migration scenarios in experimental wetland communities

Atmospheric warming may influence plant productivity and diversity and induce poleward migration of species, altering communities across latitudes. Complicating the picture is that communities from different continents deviate in evolutionary histories, which may modify responses to warming and migration. We used experimental wetland plant communities grown from seed banks as model systems to determine whether effects of warming on biomass production and species richness are consistent across continents, latitudes, and migration scenarios. We collected soil samples from each of three tidal freshwater marshes in estuaries at three latitudes (north, middle, south) on the Atlantic coasts of Europe and North America. In one experiment, we exposed soil seed bank communities from each latitude and continent to ambient and elevated (+2.8 °C) temperatures in the greenhouse. In a second experiment, soil samples were mixed either within each estuary (limited migration) or among estuaries from different latitudes in each continent (complete migration). Seed bank communities of these migration scenarios were also exposed to ambient and elevated temperatures and contrasted with a no‐migration treatment. In the first experiment, warming overall increased biomass (+16%) and decreased species richness (?14%) across latitudes in Europe and North America. Species richness and evenness of south‐latitude communities were less affected by warming than those of middle and north latitudes. In the second experiment, warming also stimulated biomass and lowered species richness. In addition, complete migration led to increased species richness (+60% in North America, + 100% in Europe), but this higher diversity did not translate into increased biomass. Species responded idiosyncratically to warming, but Lythrum salicaria and Bidens sp. increased significantly in response to warming in both continents. These results reveal for the first time consistent impacts of warming on biomass and species richness for temperate wetland plant communities across continents, latitudes, and migration scenarios.     

Community structure and composition in response to climate change in a temperate steppe

Climate change would have profound influences on community structure and composition, and subsequently has impacts on ecosystem functioning and feedback to climate change. A field experiment with increased temperature and precipitation was conducted to examine effects of experimental warming, increased precipitation and their interactions on community structure and composition in a temperate steppe in northern China since April 2005. Increased precipitation significantly stimulated species richness and coverage of plant community. In contrast, experimental warming markedly reduced species richness of grasses and community coverage. Species richness was positively dependent upon soil moisture (SM) across all treatments and years. Redundancy analysis (RDA) illustrated that SM dominated the response of community composition to climate change at the individual level, suggesting indirect effects of climate change on plant community composition via altering water availability. In addition, species interaction also mediated the responses of functional group coverage to increased precipitation and temperature. Our observations revealed that both abiotic (soil water availability) and biotic (interspecific interactions) factors play important roles in regulating plant community structure and composition in response to climate change in the semiarid steppe. Therefore these factors should be incorporated in model predicting terrestrial vegetation dynamics under climate change.     

Herbivory and nutrient limitation protect warming tundra from lowland species’ invasion and diversity loss

Herbivory and nutrient limitation can increase the resistance of temperature‐limited systems to invasions under climate warming. We imported seeds of lowland species to tundra under factorial treatments of warming, fertilization, herbivore exclusion and biomass removal. We show that warming alone had little impact on lowland species, while exclusion of native herbivores and relaxation of nutrient limitation greatly benefitted them. In contrast, warming alone benefitted resident tundra species and increased species richness; however, these were canceled by negative effects of herbivore exclusion and fertilization. Dominance of lowland species was associated with low cover of tundra species and resulted in decreased species richness. Our results highlight the critical role of biotic and abiotic filters unrelated to temperature in protecting tundra under warmer climate. While scarcity of soil nutrients and native herbivores act as important agents of resistance to invasions by lowland species, they concurrently promote overall species coexistence. However, when these biotic and abiotic resistances are relaxed, invasion of lowland species can lead to decreased abundance of resident tundra species and diminished diversity.     

The interacting effects of herbivore exclosures and seed addition in a wet meadow

Seed limitation may prevent successful restoration of native plant communities. Seed addition is a common restoration practice but the role of small mammals in affecting seedling recruitment is not well understood. The purpose of this investigation was to test the relative effect of seed introduction in combination with small mammal and bird exclosures in an Ohio wet meadow. We ask whether the ambient population of Microtus pennsylvanicus (1) alters species composition (e.g. forb/grass/sedge, invasive, non-native); (2) influences plant diversity; and (3) reduces the effect of increasing local plant richness through seed introductions. We established a 2×2 factorial design including a seed addition treatment (0 and 20 seed species added) and an exclosure treatment (open and fenced to exclude all mammalian and bird herbivores and granivores). Seeds from twenty native species were selected to represent a broad range of plant life forms typically found in temperate eastern North American wet meadow communities. All species were obligate or facultative wetland species with forbs, grasses and sedges represented. We found that forb species increased inside exclosures, especially in the seed addition treatment. We also found that relative biomass of invasive species was reduced in exclosures and with seed addition. Species richness increased with seed addition; however, exclosures significantly increased species richness and diversity, particularly of those species that were experimentally introduced by seed. Our results support the seed limitation hypotheses. It is also evident that seed and seedling predation are important factors that can control wet meadow community composition and diversity.     

Community diversity outweighs effect of warming on plant colonization

Abiotic environmental change, local species extinctions and colonization of new species often co‐occur. Whether species colonization is driven by changes in abiotic conditions or reduced biotic resistance will affect community functional composition and ecosystem management. We use a grassland experiment to disentangle effects of climate warming and community diversity on plant species colonization. Community diversity had dramatic impacts on the biomass, richness and traits of plant colonists. Three times as many species colonized the monocultures than the high diversity 17 species communities (~30 vs. 10 species), and colonists collectively produced 10 times as much biomass in the monocultures than the high diversity communities (~30 vs. 3 g/m²). Colonists with resource‐acquisitive strategies (high specific leaf area, light seeds, short heights) accrued more biomass in low diversity communities, whereas species with conservative strategies accrued most biomass in high diversity communities. Communities with higher biomass of resident C4 grasses were more resistant to colonization by legume, nonlegume forb and C3 grass colonists, but not by C4 grass colonists. Compared with effects of diversity, 6 years of 3°C‐above‐ambient temperatures had little impact on plant colonization. Warmed subplots had ~3 fewer colonist species than ambient subplots and selected for heavier seeded colonists. They also showed diversity‐dependent changes in biomass of C3 grass colonists, which decreased under low diversity and increased under high diversity. Our findings suggest that species colonization is more strongly affected by biotic resistance from residents than 3°C of climate warming. If these results were extended to invasive species management, preserving community diversity should help limit plant invasion, even under climate warming.     

Herbivore species richness, composition and community structure mediate predator richness effects and top-down control of herbivore biomass

Changes in predator species richness can have important consequences for ecosystem functioning at multiple trophic levels, but these effects are variable and depend on the ecological context in addition to the properties of predators themselves. Here, we report an experimental study to test how species identity, community attributes, and community structure at the herbivore level moderate the effects of predator richness on ecosystem functioning. Using mesocosms containing predatory insects and aphid prey, we independently manipulated species richness at both predator and herbivore trophic levels. Community structure was also manipulated by changing the distribution of herbivore species across two plant species. Predator species richness and herbivore species richness were found to negatively interact to influence predator biomass accumulation, an effect which is hypothesised to be due to the breakdown of functional complementarity among predators in species-rich herbivore assemblages. The strength of predator suppression of herbivore biomass decreased as herbivore species richness and distribution across host plants increased, and positive predator richness effects on herbivore biomass suppression were only observed in herbivore assemblages of relatively low productivity. In summary, the study shows that the species richness, productivity and host plant distribution of prey communities can all moderate the general influence of predators and the emergence of predator species richness effects on ecosystem functioning.     

Non-random recruitment of invader species in experimental grasslands

To assess potential effects of seed limitation, characteristics of invader species and characteristics of established plant communities on recruitment success, we conducted a split-plot experiment factorially combining three weeding treatments corresponding to increasing successional age (regular weeding相似文献   

Staged invasions across disparate grasslands: effects of seed provenance,consumers and disturbance on productivity and species richness

Exotic plant invasions are thought to alter productivity and species richness, yet these patterns are typically correlative. Few studies have experimentally invaded sites and asked how addition of novel species influences ecosystem function and community structure and examined the role of competitors and/or consumers in mediating these patterns. We invaded disturbed and undisturbed subplots in and out of rodent exclosures with seeds of native or exotic species in grasslands in Montana, California and Germany. Seed addition enhanced aboveground biomass and species richness compared with no‐seeds‐added controls, with exotics having disproportionate effects on productivity compared with natives. Disturbance enhanced the effects of seed addition on productivity and species richness, whereas rodents reduced productivity, but only in Germany and California. Our results demonstrate that experimental introduction of novel species can alter ecosystem function and community structure, but that local filters such as competition and herbivory influence the magnitude of these impacts.     

Constraints on colonization and species richness along a grassland productivity gradient: the role of propagule availability

An important goal in ecology is to discern under what habitat conditions community structure is primarily regulated by local ecological interactions and under what conditions community structure is more regulated by the pool of available colonists. I conducted a seed addition experiment in successional grassland to evaluate the relative significance of neighbourhood biotic interactions and propagule availability in regulating plant colonization and species richness along a natural gradient of grassland productivity. In undisturbed field plots, seed additions of 34 species led to an increase in species richness in locations of low productivity, an effect that declined in magnitude as productivity increased. In disturbed plots, seed additions led to a relatively constant increase in species richness at all levels of productivity. The results support the hypothesis that the role of propagule availability in regulating colonization dynamics and species richness declines in significance relative to local-scale competitive interactions as habitat productivity increases.     

Dispersal and seed limitation affect diversity and productivity of montane grasslands

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity–ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m⁻² and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m⁻². One year after seed addition, local plant species richness had increased on average by six species m⁻² (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m⁻² (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump‐shaped model appears to be the limiting outline of the natural diversity–productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems.     

Plant Community Responses to Simultaneous Changes in Temperature,Nitrogen Availability,and Invasion

Background

Increasing rates of change in climate have been observed across the planet and have contributed to the ongoing range shifts observed for many species. Although ecologists are now using a variety of approaches to study how much and through what mechanisms increasing temperature and nutrient pollution may influence the invasions inherent in range shifts, accurate predictions are still lacking.

Methods and Results

In this study, we conducted a factorial experiment, simultaneously manipulating warming, nitrogen addition and introduction of Pityopsis aspera, to determine how range-shifting species affect a plant community. We quantified the resident community using ordination scores, then used structural equation modeling to examine hypotheses related to how plants respond to a network of experimental treatments and environmental variables. Variation in soil pH explained plant community response to nitrogen addition in the absence of invasion. However, in the presence of invasion, the direct effect of nitrogen on the community was negligible and soil moisture was important for explaining nitrogen effects. We did not find effects of warming on the native plant community in the absence of invasion. In the presence of invasion, however, warming had negative effects on functional richness directly and invasion and herbivory explained the overall positive effect of warming on the plant community.

Conclusions and Significance

This work highlights the variation in the biotic and abiotic factors responsible for explaining independent and collective climate change effects over a short time scale. Future work should consider the complex and non-additive relationships among factors of climate change and invasion in order to capture more ecologically relevant features of our changing environment.     

Reducing dispersal limitation via seed addition increases species richness but not above‐ground biomass

Seed dispersal limitation, which can be exacerbated by a number of anthropogenic causes, can result in local communities having fewer species than they might potentially support, representing a potential diversity deficit. The link between processes that shape natural variation in diversity, such as dispersal limitation, and the consequent effects on productivity is less well known. Here, we synthesised data from 12 seed addition experiments in grassland communities to examine the influence of reducing seed dispersal limitation (from 1 to 60 species added across experiments) on species richness and productivity. For every 10 species of seed added, we found that species richness increased by about two species. However, the increase in species richness by overcoming seed limitation did not lead to a concomitant increase in above‐ground biomass production. This highlights the need to consider the relationship between biodiversity and ecosystem functioning in a pluralistic way that considers both the processes that shape diversity and productivity simultaneously in naturally assembled communities.     

Local and regional processes in low-productive mountain plant communities: the roles of seed and microsite limitation in relation to grazing

It is becoming widely accepted that plant community structure is determined not only by local scale factors, but that regional factors may play considerable role. The research studying the associated processes in different environments with different species assemblages is still limited. We conducted a two-year seed sowing experiment to test whether a plant community in a low-productive mountain snowbed is limited by seed or microsite availability and how these variables depend on natural grazing. In a factorial design, half of the plots received a mixture of seeds of fourteen species naturally occurring at the study site and above ground biomass was removed from half of the plots. These treatments were applied to plots with long term grazer exclosures and to plots accessible to grazers. Both sowing and biomass removal increased the number of seedlings, the species richness of seedlings and total species richness. The number of seedlings was higher in open plots than in exclosures in the second year. Both seedling richness and total species richness were higher in open plots. Seedling recruitment was negatively related to the amount of above ground biomass and positively to the initial species richness. These results suggest that even fairly low-productive environments can be both seed and microsite limited and that these depend on grazing pressure. Natural grazing by mammal herbivores (e.g. lemmings and reindeer) favours species colonization and seedling emergence. Low-productive mountain snowbeds are prone to colonization from the local species pool and even high species richness may not constrain ingression of new species.     

The interplay between species' positive and negative interactions shapes the community biomass–species richness relationship

We used an individual-based spatially-explicit model to assess the role of facilitation and plant strategies in shaping the 'community biomass–species richness' relationship. Facilitation had few impacts on community's richness under both the most benign (high community biomass) and the most severe (low community biomass) environments where its intensity was weak. From medium to high environmental severity, facilitation increased community richness, because all plant strategies were facilitated. In contrast, from low to medium environmental severity facilitation decreased community richness, because only the most competitive species were facilitated, which induced a decrease in the richness of the stress-tolerant species overwhelming the increase in richness of the competitive species. Above all, our simulations show how 'strategy-dependent' interactions among species combine to shape the humped-back biomass–species richness relationship. It also demonstrates that facilitative effects might have long-term negative effects on species richness, which result is not included in current facilitation models.     

Impacts of sedimentation and nitrogen enrichment on wetland plant community development

Many factors influence which plant species are found in a particular wetland. The species pool is composed of the species present in the seed bank and species able to disperse into the wetland, and many abiotic and biotic factors interact to influence a species performance and abundance in the plant community. Anthropogenic activities produce specific stressors on wetland systems that alter these abiotic and biotic interactions, potentially altering species composition. We simulated three common wetland hydrogeomorphic (HGM) subclasses in a greenhouse to examine the effects of two stressors-sedimentation and nitrogen (N) enrichment-on the performance of 8 species grown in artificial communities. Species establishment, height, biomass, and foliar N and P concentrations were measured to explore species responses to stressors and competition, as well as the potential impacts of changes in species composition on ecosystem processes. Species were affected differently by sedimentation and N enrichment, and there were differences in overall community sensitivity to stressors between wetland subclasses. Sedimentation generally reduced seedling establishment, while N enrichment produced variable effects on height and biomass. Interspecific competition had little effect on establishment but significantly reduced most species biomass. Sedimentation generally lowered community biomass, diversity, and richness, while enrichment increased community biomass. Establishment, biomass, and foliar nutrient concentrations significantly differed between many species, suggesting that shifts in species composition may impact ecosystem processes such as nutrient cycling and carbon storage. Phalaris arundinacea, an aggressive clonal graminoid, universally dominated all wetland subclasses. This dominance across a range of environmental conditions (sedimentation, fertility, and hydrology) has important implications for both restoration and predicting the impacts of human activities on species composition. Our results suggest that, in regions where P. arundinacea is common, restoration projects that establish communities from seeds and human activities that cause vegetation removal are likely to become dominated by P. arundinacea.