Relative Importance Of Facilitation Research Articles

Geomorphology and Species Interactions Control Facilitation Cascades in a Salt Marsh Ecosystem.

Facilitation cascades are chains of positive interactions that occur as frequently as trophic cascades and are equally important drivers of ecosystem function, where they involve the overlap of primary and secondary, or dependent, habitat-forming foundation species [1]. Although it is well recognized that the size and configuration of secondary foundation species' patches are critical features modulating the ecological effects of facilitation cascades [2], the mechanisms governing their spatial distribution are often challenging to discern given that they operate across multiple spatial and temporal scales [1, 3]. We therefore combined regional surveys of southeastern US salt marsh geomorphology and invertebrate communities with a predator exclusion experiment to elucidate the drivers, both geomorphic and biotic, controlling the establishment, persistence, and ecosystem functioning impacts of a regionally abundant facilitation cascade involving habitat-forming marsh cordgrass and aggregations of ribbed mussels. We discovered a hierarchy of physical and biological factors predictably controlling the strength and self-organization of this facilitation cascade across creekshed, landscape, and patch scales. These results significantly enhance our capacity to spatially predict coastal ecosystem function across scales based on easily identifiable metrics of geomorphology that are mechanistically linked to ecological processes. Replication of this approach across vegetated coastal ecosystems has the potential to support management efforts by elucidating the multi-scale linkages between geomorphology and ecology that, in turn, define spatially explicit patterns in community assembly and ecosystem functioning.

Experimental analyses of diversity partitioning in southern hemisphere algal communities.

Understanding how biodiversity and its components of alpha, beta, and gamma vary over spatial and temporal scales and across communities is crucial to mitigating stressors of ecosystems. Marine communities present several problems in partitioning diversity over fine spatial scales, such as tidal zones, and temporal scales relating to seasonal occurrences of species and recovery responses to impacts. This study uses an experimental approach to test disturbance effects on beta diversity in algal communities in southern New Zealand. Dominant canopies in mid-shore Hormosira banksii and low-shore Durvillaea poha communities were removed and diversity metrics assessed, including additive partitioning, permutational dispersion, and nestedness and turnover analyses. Over 2years, 258 species were found. Species richness was greater where canopies remained intact in Hormosira communities compared to removal plots, but, in Durvillaea communities, controls and removals had similar richness. In both communities, β-diversity was 1.5-3.9 times greater than α-diversity, with the temporal component βt being 1.2-2.4 × greater than the spatial component. Hormosira communities exhibited high nestedness, with species in removal plots being a subset of those in controls. In Durvillaea communities, however, turnover was high and nestedness low, because removal plots had a different species assemblage than controls. Multivariate analyses showed that species occurrences and abundances remained different in controls and removals in both communities over 2years. Differences in diversity components between communities were related to environmental differences to which they are exposed, including desiccation and wave forces, and the relative importance of facilitation and competition in the different communities.

Facilitation by Tree Species in Variable Retention Harvesting for the Restoration of Monoculture Plantations in Southern China

Facilitation can drive the successional dynamics and change the restoration trajectory of degraded forests. However, the relative importance of facilitation by tree species after variable retention harvesting is unclear. We used a field experiment to evaluate the effect of two facilitator species, Castanopsis fissa (C. fissa) and Manglietia glauca (M. glauca), managed with variable retention harvesting, on the development of two target species, Castanopsis hystrix (C. hystrix) and Erythrophloeum fordii (E. fordii), in a Masson pine ( Pinus massoniana) monoculture. The following variables were measured for all of the four interplanted tree species: structural growth, regeneration, aboveground biomass accumulation, leaf area index, and soil conditions. The results indicate that the abundance, growth, and aboveground biomass were relatively greater in plots planted with C. fissa compared with M. glauca and that the target species performed best with 50% retention harvesting of C. fissa, with an improved establishment of both target species indicating a positive interaction. In addition, the regeneration, leaf area index and soil conditions differed between the two facilitators in the variable retention harvesting treatments because of the different intrinsic characteristic of the facilitators. In summary, our results imply that managers have considerable flexibility to employ various types of facilitation schemes coupled with different harvesting systems for successful short-term restoration within a monoculture.

Intraspecific interactions shift from competitive to facilitative across a low to high disturbance gradient in a salt marsh

Disturbances are ubiquitous among salt marshes, and disturbances such as litter removal may form stressful environmental conditions. The stress-gradient hypothesis (SGH) predicts that the relative importance of facilitation and competition between species will vary inversely to each other across abiotic stress gradients. However, how intraspecific interactions vary across disturbance levels, and whether they follow the SGH has not been investigated. To test the SGH with intraspecific interactions within Suaeda salsa individuals, we assessed individual survival in established fully mapped salt marsh plots with two treatments, disturbed, litter removal and control, undisturbed plots. Recently developed spatial statistics were applied to distinguish between random processes, intraspecific facilitation, scramble competition, and contest competition underlying the spatial patterns at different spatial scales, growth stages, and disturbance levels. We found evidence that intraspecific interactions among S. salsa individuals across disturbance levels tended to support the SGH, but this support depended on time and space. Intraspecific interactions were more likely to show positive density dependence in disturbed plots, particularly at the seedling period and when individuals were separated by < 9 cm. However, positive density dependence was not detected in undisturbed plots. The protective effects of litter on seedlings, which were lost in undisturbed sites, may have been at least partly simulated at high seedling densities. Mortality in disturbed sites was not random, and facilitation had a more important influence than competition on population dynamics in high disturbance sites. However, the influence of competition on population dynamics, specifically scramble competition was also clearly important.

Changes in spatial point patterns of pioneer woody plants across a large tropical landslide

We assessed whether the relative importance of positive and negative interactions in early successional communities varied across a large landslide on Casita Volcano (Nicaragua). We tested several hypotheses concerning the signatures of these processes in the spatial patterns of woody pioneer plants, as well as those of mortality and recruitment events, in several zones of the landslide differing in substrate stability and fertility, over a period of two years (2001 and 2002). We identified all woody individuals with a diameter >1 cm and mapped them in 28 plots measuring 10 × 10-m. On these maps, we performed a spatial point pattern analysis using univariate and bivariate pair-correlation functions; g (r) and g12 (r), and pairwise differences of univariate and bivariate functions. Spatial signatures of positive and negative interactions among woody plants were more prevalent in the most and least stressful zones of the landslide, respectively. Natural and human-induced disturbances such as the occurrence of fire, removal of newly colonizing plants through erosion and clearcutting of pioneer trees were also identified as potentially important pattern-creating processes. These results are in agreement with the stress-gradient hypothesis, which states that the relative importance of facilitation and competition varies inversely across gradients of abiotic stress. Our findings also indicate that the assembly of early successional plant communities in large heterogeneous landslides might be driven by a much larger array of processes than previously thought.

Moving forward on facilitation research: response to changing environments and effects on the diversity, functioning and evolution of plant communities.

Once seen as anomalous, facilitative interactions among plants and their importance for community structure and functioning are now widely recognized. The growing body of modelling, descriptive and experimental studies on facilitation covers a wide variety of terrestrial and aquatic systems throughout the globe. However, the lack of a general body of theory linking facilitation among different types of organisms and biomes and their responses to environmental changes prevents further advances in our knowledge regarding the evolutionary and ecological implications of facilitation in plant communities. Moreover, insights gathered from alternative lines of inquiry may substantially improve our understanding of facilitation, but these have been largely neglected thus far. Despite over 15 years of research and debate on this topic, there is no consensus on the degree to which plant-plant interactions change predictably along environmental gradients (i.e. the stress-gradient hypothesis), and this hinders our ability to predict how plant-plant interactions may affect the response of plant communities to ongoing global environmental change. The existing controversies regarding the response of plant-plant interactions across environmental gradients can be reconciled when clearly considering and determining the species-specificity of the response, the functional or individual stress type, and the scale of interest (pairwise interactions or community-level response). Here, we introduce a theoretical framework to do this, supported by multiple lines of empirical evidence. We also discuss current gaps in our knowledge regarding how plant-plant interactions change along environmental gradients. These include the existence of thresholds in the amount of species-specific stress that a benefactor can alleviate, the linearity or non-linearity of the response of pairwise interactions across distance from the ecological optimum of the beneficiary, and the need to explore further how frequent interactions among multiple species are and how they change across different environments. We review the latest advances in these topics and provide new approaches to fill current gaps in our knowledge. We also apply our theoretical framework to advance our knowledge on the evolutionary aspects of plant facilitation, and the relative importance of facilitation, in comparison with other ecological processes, for maintaining ecosystem structure, functioning and dynamics. We build links between these topics and related fields, such as ecological restoration, woody encroachment, invasion ecology, ecological modelling and biodiversity-ecosystem-functioning relationships. By identifying commonalities and insights from alternative lines of research, we further advance our understanding of facilitation and provide testable hypotheses regarding the role of (positive) biotic interactions in the maintenance of biodiversity and the response of ecological communities to ongoing environmental changes.

Spatial patterns provide support for the stress-gradient hypothesis over a range-wide aridity gradient

We examined variations in the relative importance of facilitation vs. competition, in light of the Stress-Gradient Hypothesis (SGH) by assessing plant interactions along an aridity gradient over biogeographic scales. We surveyed the relationship between a shrub species (Artemisia tridentata) and pine seedlings (Pinus monophylla) across the Great Basin and the Mojave Desert, USA, encompassing the entire range of P. monophylla. Using 69 sites we evaluated the spatial association between P. monophylla seedlings and A. tridentata shrubs, quantified with an electivity index, and implemented multiple regression analysis on the effects of macro- and micro-environmental factors: precipitation, temperature, monsoonality index, topography, substrate and litter cover. We identified annual precipitation as a main factor, which was negatively related to shrub-seedling association. Additionally, shrub-seedling association was stronger in the hot- than in the cold-desert, and was negatively related to litter cover. Effects of monsoonality, summer temperature, and bedrock type were not significant. We also considered nonlinear functional forms of a precipitation–electivity relationship, but the negative linear model proved most predictive. Our observations match SGH predictions. Studying the role of interspecific interactions in shaping species range shifts may lead to improved predictions of distribution ranges and changes in dryland vegetation under global change scenarios.

Testing the Stress‐Gradient Hypothesis During the Restoration of Tropical Degraded Land Using the Shrub Rhodomyrtus tomentosa as a Nurse Plant

AbstractThe relative importance of facilitation and competition between pairwise plants across abiotic stress gradients as predicted by the stress‐gradient hypothesis has been confirmed in arid and temperate ecosystems, but the hypothesis has rarely been tested in tropical systems, particularly across nutrient gradients. The current research examines the interactions between a pioneer shrub Rhodomyrtus tomentosa (the nurse plant) and seedlings of a transplanted native woody Schima superba (the target species) in a tropical system in which position on a slope corresponds with a nutrient gradient; high soil nutrients at the slope bottom and relatively low soil nutrients at the slope top. In contrast, soil physical traits were more favorable for seedling growth under the shrub than in open spaces. The effect of R. tomentosa on S. superba survival was positive (facilitation) at the top of the slope, as indicated by the relative interaction index (RII), but negative in the bottom (competition). RII indicated a positive effect on seedling height at the top of the slope but was not at the bottom. Seedling survival was positively related to soil nutrient level and negatively related to soil acidity, but seedling growth of S. superba seemed to be enhanced by the shrub canopy. Thus, the results seem to support stress‐gradient hypothesis in terms of target species survival but not growth. We suggest using the shrub as a nurse plant in forest restoration in tropical degraded land with caution because not all of its effects on target species are positive.

Shifting Species Interaction in Soil Microbial Community and Its Influence on Ecosystem Functions Modulating

The supportive and negative evidence for the stress gradient hypothesis (SGH) led to an ongoing debate among ecologists and called for new empirical and theoretical work. In this study, we took various biological soil crust (BSCs) samples along a spatial gradient with four environmental stress levels to examine the fitness of SGH in microbial interactions and evaluate its influence on biodiversity-function relationships in BSCs. A new assessment method of species interactions within hard-cultured invisible soil community was employed, directly based on denaturing gradient gel electrophoresis fingerprint images. The results showed that biotic interactions in soil phototroph community dramatically shifted from facilitation to dominant competition with the improvement of microhabitats. It offered new evidence, which presented a different perspective on the hypothesis that the relative importance of facilitation and competition varies inversely along the gradient of abiotic stress. The path analysis indicated that influence of biotic interactions (r = 0.19, p < 0.05) on ecosystem functions is lower than other community properties (r = 0.62, p < 0.001), including soil moisture, crust coverage, and biodiversity. Furthermore, the correlation between species interactions and community properties was non-significant with low negative influence (r = -0.27, p > 0.05). We demonstrate that the inversion of biotic interaction as a response to the gradient of abiotic stresses existed not only in the visible plant community but also in the soil microbial community.

Distance-dependent shifts in net effects by an unpalatable nettle on a palatable plant species

We examined whether the relative importance of facilitation and competition effects by an unpalatable perennial ( Urtica thunbergiana) on a palatable annual ( Persicaria longiseta) change with the spatial distance between them in a long-term deer-grazing habitat. Survivorship, growth, size, and fecundity of Persicaria were recorded at 1–2-month intervals during growing seasons in 2 successive years and were compared among individuals located in different positions relative to the canopy of Urtica: at the centre, internal edge, external edge, and far from the canopy. Survivorship of Persicaria was significantly higher at the centre of the Urtica canopy than outside the canopy early in the growing season in both years. No significant differences in Persicaria growth were observed among the four positions in most periods, except in one when growth was significantly higher at the centre, internal, and external edges of the canopy compared to outside the canopy. We found spatial shifts in the net effects of Urtica on Persicaria fecundity, from positive effects under the canopy centre to negative effects under the external edge of the canopy in the first year, and from negative effects under the centre to positive effects under the external edge in the second year. These results demonstrate that the relative importance of positive and negative effects of Urtica on Persicaria vary temporally within and among years and spatially around a single Urtica plant. Spatiotemporal variation in plant interactions may be attributable to annual and seasonal variation in vegetation productivity and grazing pressure.

Interactions of Acacia raddiana with herbaceous vegetation change with intensity of abiotic stress

Theoretical models predict that the relative importance of facilitation and competition may vary inversely across gradients of abiotic stress. However, these predictions have not been thoroughly tested in the field, especially in arid and semi-arid environments. In this study, we evaluated how the net effect of Acacia tortilis subsp. raddiana (syn. A. raddiana) trees on the herbaceous species varies across a gradient of water stress in arid Tunisian ecosystems. Our results show that the influence of trees on the herbaceous stratum is beneficial. An Acacia tree improves the richness of herbaceous species around it by two to three times. The positive effect of Acacia raddiana on species composition is characterised in particular by a better development of some species of high pastoral value such as Cenchrus ciliaris, Cynodon dactylon, Eragrostis papposa, Sisymbrium irio and Chenopodium album. We fitted the relationship between seasonal rainfall and the relative neighbour effect index, which varied across this gradient, to a linear model. Our results show that herbaceous plant cover, plant biomass, plant density and diversity are higher under a tree canopy and this positive interaction still increase with higher abiotic stress conditions.

Is the change of plant–plant interactions with abiotic stress predictable? A meta‐analysis of field results in arid environments

1 Theoretical models have predicted that the relative importance of facilitation and competition will vary inversely across gradients of abiotic stress, with facilitation being the dominant interaction under high abiotic stress conditions. A critical reappraisal of current theoretical models is needed because experimental studies both support and refute their predictions. 2 A quantitative meta-analysis of field and common garden studies evaluating the effect of abiotic stress (low vs. high) on the net outcome of plant–plant interactions in arid and semi-arid environments was performed to evaluate the degree of empirical support for these models. We created four separate data sets corresponding to the categories of response variables commonly used to measure plant performance (survival, density, growth and fecundity). 3 The analyses showed that both the selection of the estimator of plant performance and the experimental approach followed have a strong influence on both the net outcome of plant–plant interactions and the effect of abiotic stress on such outcome. The effect of neighbours on the survival and growth of target plants was not significant at either stress level, but that on the density and fecundity of target plants was positive (facilitation) and negative (competition) at the low and high abiotic stress level, respectively. Density data showed that the net effect of neighbours was positive and negative at low and high abiotic stress levels, respectively, whereas other estimators suggested that the net effect of neighbours did not differ with stress level. None of our meta-analyses indicated that the magnitude of the net effect provided by neighbours, whether positive or negative, was higher under high abiotic stress conditions, and facilitation does not therefore appear to increase in importance with abiotic stress. 4 As the predictions of current theoretical models regarding the relationship between the net outcome of a plant–plant interaction and abiotic stress do not hold in arid and semi-arid environments, different models are needed. These should consider different sources of abiotic stress separately, and should be valid for performance measurements, such as survival, that integrate plant responses over time. The incorporation of these features into theoretical models will undoubtedly improve their predictive capabilities.

Do positive interactions increase with abiotic stress? A test from a semi-arid steppe

Theoretical models predict that the relative importance of facilitation and competition may vary inversely across gradients of abiotic stress. However, these predictions have not been thoroughly tested in the field, especially in semi-arid environments. In this study, we evaluated how the net effect of the tussock grass Stipa tenacissima on the shrub Pistacia lentiscus varied across a gradient of abiotic stress in semi-arid Mediterranean steppes. We fitted the relationship between accumulated rainfall and the relative neighbour index (our measures of abiotic stress and of the net effect of S. tenacissima on P. lentiscus, respectively), which varied across this gradient, to a quadratic model. Competitive interactions dominated at both extremes of the gradient. Our results do not support established theory. Instead, they suggest that a shift from facilitation to competition under high abiotic stress conditions is likely to occur when the levels of the most limiting resource are so low that the benefits provided by the facilitator cannot overcome its own resource uptake.

Pattern analysis in successional communities – An approach for studying shifts in ecological interactions

Abstract. Many ecological studies have addressed issues of vegetation spatial patterns in attempts to understand the processes generating them. We investigated changes in ecological processes during succession via the analysis of shrubs’ spatial patterns in a system of linear sand dunes, an arid ecosystem located in the Negev Desert in Israel during three consecutive years. We hypothesized that spatial patterns change from clustered to regular as succession progresses due to changes in the relative importance of facilitation and competition in this environment. In this ecosystem communities of early successional stages are frequently disturbed by high rates of sand movement, whereas in later successional stages sand stability is high. We mapped in the field individual shrubs on high‐resolution aerial photographs, and converted the digital images to a GIS data set. Using Ripley's K‐function we analysed spatial patterns at three levels: the single‐species level, among species and at the individual level, in three communities characterizing different successional stages. In the early successional communities we found clustered spatial patterns, in comparison with stable habitats where spatial patterns tended to be regular. We argue that these shifts in spatial patterns are indicative of the assumption that in this sand‐dune system ecological interactions change from facilitation to competition as succession progresses. Further, we argue that these interactions operate in different spatial scales at the different successional stages, and that the study of these processes should be conducted at the spatial scales specific to each community.

Pattern analysis in successional communities – An approach for studying shifts in ecological interactions

Many ecological studies have addressed issues of vegetation spatial patterns in attempts to understand the proc- esses generating them. We investigated changes in ecological processes during succession via the analysis of shrubs' spatial patterns in a system of linear sand dunes, an arid ecosystem located in the Negev Desert in Israel during three consecutive years. We hypothesized that spatial patterns change from clustered to regular as succession progresses due to changes in the relative importance of facilitation and competition in this environment. In this ecosystem communities of early succes- sional stages are frequently disturbed by high rates of sand movement, whereas in later successional stages sand stability is high. We mapped in the field individual shrubs on high- resolution aerial photographs, and converted the digital im- ages to a GIS data set. Using Ripley's K-function we analysed spatial patterns at three levels: the single-species level, among species and at the individual level, in three communities characterizing different successional stages. In the early suc- cessional communities we found clustered spatial patterns, in comparison with stable habitats where spatial patterns tended to be regular. We argue that these shifts in spatial patterns are indicative of the assumption that in this sand-dune system ecological interactions change from facilitation to competition as succession progresses. Further, we argue that these interac- tions operate in different spatial scales at the different succes- sional stages, and that the study of these processes should be conducted at the spatial scales specific to each community.