The strength of negative plant–soil feedback increases from the intraspecific to the interspecific and the functional group level

One of the processes that may play a key role in plant species coexistence and ecosystem functioning is plant–soil feedback, the effect of plants on associated soil communities and the resulting feedback on plant performance. Plant–soil feedback at the interspecific level (comparing growth on own soil with growth on soil from different species) has been studied extensively, while plant–soil feedback at the intraspecific level (comparing growth on own soil with growth on soil from different accessions within a species) has only recently gained attention. Very few studies have investigated the direction and strength of feedback among different taxonomic levels, and initial results have been inconclusive, discussing phylogeny, and morphology as possible determinants. To test our hypotheses that the strength of negative feedback on plant performance increases with increasing taxonomic level and that this relationship is explained by morphological similarities, we conducted a greenhouse experiment using species assigned to three taxonomic levels (intraspecific, interspecific, and functional group level). We measured certain fitness‐related aboveground traits and used them along literature‐derived traits to determine the influence of morphological similarities on the strength and direction of the feedback. We found that the average strength of negative feedback increased from the intraspecific over the interspecific to the functional group level. However, individual accessions and species differed in the direction and strength of the feedback. None of our results could be explained by morphological dissimilarities or individual traits. Synthesis. Our results indicate that negative plant–soil feedback is stronger if the involved plants belong to more distantly related species. We conclude that the taxonomic level is an important factor in the maintenance of plant coexistence with plant–soil feedback as a potential stabilizing mechanism and should be addressed explicitly in coexistence research, while the traits considered here seem to play a minor role.     

Intraspecific plant–soil feedback and intraspecific overyielding in Arabidopsis thaliana

Understanding the mechanisms of community coexistence and ecosystem functioning may help to counteract the current biodiversity loss and its potentially harmful consequences. In recent years, plant–soil feedback that can, for example, be caused by below‐ground microorganisms has been suggested to play a role in maintaining plant coexistence and to be a potential driver of the positive relationship between plant diversity and ecosystem functioning. Most of the studies addressing these topics have focused on the species level. However, in addition to interspecific interactions, intraspecific interactions might be important for the structure of natural communities. Here, we examine intraspecific coexistence and intraspecific diversity effects using 10 natural accessions of the model species Arabidopsis thaliana (L.) Heynh. We assessed morphological intraspecific diversity by measuring several above‐ and below‐ground traits. We performed a plant–soil feedback experiment that was based on these trait differences between the accessions in order to determine whether A. thaliana experiences feedback at intraspecific level as a result of trait differences. We also experimentally tested the diversity–productivity relationship at intraspecific level. We found strong differences in above‐ and below‐ground traits between the A. thaliana accessions. Overall, plant–soil feedback occurred at intraspecific level. However, accessions differed in the direction and strength of this feedback: Some accessions grew better on their own soils, some on soils from other accessions. Furthermore, we found positive diversity effects within A. thaliana: Accession mixtures produced a higher total above‐ground biomass than accession monocultures. Differences between accessions in their feedback response could not be explained by morphological traits. Therefore, we suggest that they might have been caused by accession‐specific accumulated soil communities, by root exudates, or by accession‐specific resource use based on genetic differences that are not expressed in morphological traits. Synthesis. Our results provide some of the first evidence for intraspecific plant–soil feedback and intraspecific overyielding. These findings may have wider implications for the maintenance of variation within species and the importance of this variation for ecosystem functioning. Our results highlight the need for an increased focus on intraspecific processes in plant diversity research to fully understand the mechanisms of coexistence and ecosystem functioning.     

Spatial heterogeneity of plant–soil feedbacks increases per capita reproductive biomass of species at an establishment disadvantage

Plant–soil feedbacks have been widely implicated as a driver of plant community diversity, and the coexistence prediction generated by a negative plant–soil feedback can be tested using the mutual invasibility criterion: if two populations are able to invade one another, this result is consistent with stable coexistence. We previously showed that two co-occurring Rumex species exhibit negative pairwise plant–soil feedbacks, predicting that plant–soil feedbacks could lead to their coexistence. However, whether plants are able to reproduce when at an establishment disadvantage (“invasibility”), or what drivers in the soil might correlate with this pattern, are unknown. To address these questions, we created experimental plots with heterogeneous and homogeneous soils using field-collected conditioned soils from each of these Rumex species. We then allowed resident plants of each species to establish and added invader seeds of the congener to evaluate invasibility. Rumex congeners were mutually invasible, in that both species were able to establish and reproduce in the other’s resident population. Invaders of both species had twice as much reproduction in heterogeneous compared to homogeneous soils; thus the spatial arrangement of plant–soil feedbacks may influence coexistence. Soil mixing had a non-additive effect on the soil bacterial and fungal communities, soil moisture, and phosphorous availability, suggesting that disturbance could dramatically alter soil legacy effects. Because the spatial arrangement of soil patches has coexistence implications, plant–soil feedback studies should move beyond studies of mean effects of single patch types, to consider how the spatial arrangement of patches in the field influences plant communities.     

Signs of stabilisation and stable coexistence

Many empirical studies motivated by an interest in stable coexistence have quantified negative density dependence, negative frequency dependence, or negative plant–soil feedback, but the links between these empirical results and ecological theory are not straightforward. Here, we relate these analyses to theoretical conditions for stabilisation and stable coexistence in classical competition models. By stabilisation, we mean an excess of intraspecific competition relative to interspecific competition that inherently slows or even prevents competitive exclusion. We show that most, though not all, tests demonstrating negative density dependence, negative frequency dependence, and negative plant–soil feedback constitute sufficient conditions for stabilisation of two‐species interactions if applied to data for per capita population growth rates of pairs of species, but none are necessary or sufficient conditions for stable coexistence of two species. Potential inferences are even more limited when communities involve more than two species, and when performance is measured at a single life stage or vital rate. We then discuss two approaches that enable stronger tests for stable coexistence‐invasibility experiments and model parameterisation. The model parameterisation approach can be applied to typical density‐dependence, frequency‐dependence, and plant–soil feedback data sets, and generally enables better links with mechanisms and greater insights, as demonstrated by recent studies.     

Plant‐soil feedbacks from 30‐year family‐specific soil cultures: phylogeny,soil chemistry and plant life stage

Intraspecific negative feedback effects, where performance is reduced on soils conditioned by conspecifics, are widely documented in plant communities. However, interspecific feedbacks are less well studied, and their direction, strength, causes, and consequences are poorly understood. If more closely related species share pathogens, or have similar soil resource requirements, plants may perform better on soils conditioned by more distant phylogenetic relatives. There have been few empirical tests of this prediction across plant life stages, and none of which attempt to account for soil chemistry. Here, we test the utility of phylogeny for predicting soil feedback effects on plant survival and performance (germination, seedling survival, growth rate, biomass). We implement a full factorial experiment growing species representing five families on five plant family‐specific soil sources. Our experiments exploit soils that have been cultured for over 30 years in plant family‐specific beds at Oxford University Botanic Gardens. Plant responses to soil source were idiosyncratic, and species did not perform better on soils cultured by phylogenetically more distant relatives. The magnitude and sign of feedback effects could, however, be explained by differences in the chemical properties of “home” and “away” soils. Furthermore, the direction of soil chemistry‐related plant‐soil feedbacks was dependent on plant life stage, with the effects of soil chemistry on germination success and accumulation of biomass inversely related. Our results (1) suggest that the phylogenetic distance between plant families cannot predict plant–soil feedbacks across multiple life stages, and (2) highlight the need to consider changes in soil chemistry as an important driver of population responses. The contrasting responses at plant life stages suggest that studies focusing on brief phases in plant demography (e.g., germination success) may not give a full picture of plant–soil feedback effects.     

Contrasting effects of plant inter‐ and intraspecific variation on community trait responses to restoration of a sandy grassland ecosystem

Changes in plant community traits along an environmental gradient are caused by interspecific and intraspecific trait variation. However, little is known about the role of interspecific and intraspecific trait variation in plant community responses to the restoration of a sandy grassland ecosystem. We measured five functional traits of 34 species along a restoration gradient of sandy grassland (mobile dune, semi‐fixed dune, fixed dune, and grassland) in Horqin Sand Land, northern China. We examined how community‐level traits varied with habitat changes and soil gradients using both abundance‐weighted and non‐weighted averages of trait values. We quantified the relative contribution of inter‐ and intraspecific trait variation in specific leaf area (SLA), leaf dry matter content (LDMC), leaf carbon content (LCC), leaf nitrogen content (LNC), and plant height to the community response to habitat changes in the restoration of sandy grassland. We found that five weighted community‐average traits varied significantly with habitat changes. Along the soil gradient in the restoration of sandy grassland, plant height, SLA, LDMC, and LCC increased, while LNC decreased. For all traits, there was a greater contribution of interspecific variation to community response in regard to habitat changes relative to that of intraspecific variation. The relative contribution of the interspecific variation effect of an abundance‐weighted trait was greater than that of a non‐weighted trait with regard to all traits except LDMC. A community‐level trait response to habitat changes was due largely to species turnover. Though the intraspecific shift plays a small role in community trait response to habitat changes, it has an effect on plant coexistence and the maintenance of herbaceous plants in sandy grassland habitats. The context dependency of positive and negative covariation between inter‐ and intraspecific variation further suggests that both effects of inter‐ and intraspecific variation on a community trait should be considered when understanding a plant community response to environmental changes in sandy grassland ecosystems.     

The role of plant–soil feedbacks and land-use legacies in restoration of a temperate steppe in northern China

Plant–soil feedbacks affect plant performance and plant community dynamics; however, little is known about their role in ecological restoration. Here, we studied plant–soil feedbacks in restoration of steppe vegetation after agricultural disturbance in northern China. First, we analyzed abiotic and biotic soil properties under mono-dominant plant patches in an old-field restoration site and in a ‘target’ steppe site. Second, we tested plant–soil feedbacks by growing plant species from these two sites on soils from con- and heterospecific origin. Soil properties generally did not differ between the old-field site and steppe site, but there were significant differences among mono-dominant plant patches within the sites. While soil species origin (i.e., the plant species beneath which the soil was collected) affected biomass of individual plant species in the feedback experiment, species-level plant–soil feedbacks were ‘neutral’. Soil site origin (old-field, steppe) significantly affected biomass of old-field and steppe species. For example, old-field species had higher biomass in old-field soils than in steppe soils, indicating a positive land-use legacy. However, soil site origin effects depended on the plant species beneath which the soils were collected. The predictive value of abiotic and biotic soil properties in explaining plant biomass differed between and within groups of old-field and steppe species. We conclude that the occurrence of positive land-use legacies for old-field species may retard successional replacement of old-field species by steppe species. However, high levels of idiosyncrasy in responses of old-field and steppe plant species to con- and heterospecific soils indicate interspecific variation in the extent to which soil legacies and plant–soil feedbacks control successional species replacements in Chinese steppe ecosystems.     

Native and non‐native grasses generate common types of plant–soil feedbacks by altering soil nutrients and microbial communities

Soil conditioning occurs when plants alter features of their soil environment. When these alterations affect subsequent plant growth, it is a plant soil feedback. Plant–soil feedbacks are an important and understudied aspect of aboveground–belowground linkages in plant ecology that influence plant coexistence, invasion and restoration. Here, we examine plant–soil feedback dynamics of seven co‐occurring native and non‐native grass species to address the questions of how plants modify their soil environment, do those modifications inhibit or favor their own species relative to other species, and do non‐natives exhibit different plant–soil feedback dynamics than natives. We used a two‐phase design, wherein a first generation of plants was grown to induce species‐specific changes in the soil and a second generation of plants was used as a bioassay to determine the effects of those changes. We also used path‐analysis to examine the potential chain of effects of the first generation on soil nutrients and soil microbial composition and on bioassay plant performance. Our findings show species‐specific (rather than consistent within groups of natives and non‐natives) soil conditioning effects on both soil nutrients and the soil microbial community by plants. Additionally, native species produced plant–soil feedback types that benefit other species more than themselves and non‐native invasive species tended to produce plant–soil feedback types that benefit themselves more than other species. These results, coupled with previous field observations, support hypotheses that plant–soil feedbacks may be a mechanism by which some non‐native species increase their invasive potential and plant–soil feedbacks may influence the vulnerability of a site to invasion.     

Disentangling the relative importance of species occurrence,abundance and intraspecific variability in community assembly: a trait‐based approach at the whole‐plant level in Mediterranean forests

Understanding which factors and rules govern the process of assembly in communities constitutes one of the main challenges of plant community ecology. The presence of certain functional strategies along broad environmental gradients can help to understand the patterns observed in community assembly and the filtering mechanisms that take place. We used a trait‐based approach, quantifying variations in aboveground (leaf and stem) and belowground (root) functional traits along environmental gradients in Mediterranean forest communities (south Spain). We proposed a new practical method to quantify the relative importance of species turnover (distinguishing between species occurrence and abundance) versus intraspecific variation, which allowed us to better understand the assemblage rules of these plant communities along environmental gradients. Our results showed that the functional structure of the studied plant communities was highly determined by soil environment. Results from our modelling approach based on maximum likelihood estimators showed a predominant influence of soil water storage on most of the community functional traits. We found that changes in community functional structure along environmental gradients were mainly promoted by species turnover rather than by intraspecific variability. Specifically, our new method of variance decomposition demonstrated that between‐site trait variation was the result of changes in species occurrence rather than in the abundance of certain dominant species. In conclusion, this study showed that water availability promoted the predominance of specific trait values (both in above and belowground fractions) associated to a resource acquisition or conservation strategy. In addition, we provided evidence that changes on community functional structure along the environmental gradient were mainly promoted by a process of species replacement, which represent a crucial step towards a more general understanding of the relative importance of intraspecific versus interspecific trait variation in these woody Mediterranean communities.     

Forest fire may disrupt plant–microbial feedbacks

Plant–microbial feedbacks are important drivers of plant community structure and dynamics. These feedbacks are driven by the variable modification of soil microbial communities by different plant species. However, other factors besides plant species can influence soil communities and potentially interact with plant–microbial feedbacks. We tested for plant–microbial feedbacks in two Eucalyptus species, E. globulus and E. obliqua, and the influence of forest fire on these feedbacks. We collected soils from beneath mature trees of both species within native forest stands on the Forestier Peninsula, Tasmania, Australia, that had or had not been burnt by a recent forest fire. These soils were subsequently used to inoculate seedlings of both species in a glasshouse experiment. We hypothesized that (i) eucalypt seedlings would respond differently to inoculation with conspecific versus heterospecific soils (i.e., exhibit plant–microbial feedbacks) and (ii) these feedbacks would be removed by forest fire. For each species, linear mixed effects models tested for differences in seedling survival and biomass in response to inoculation with conspecific versus heterospecific soils that had been collected from either unburnt or burnt stands. Eucalyptus globulus displayed a response consistent with a positive plant–microbial feedback, where seedlings performed better when inoculated with conspecific versus heterospecific soils. However, this effect was only present when seedlings were inoculated with unburnt soils, suggesting that fire removed the positive effect of E. globulus inoculum. These findings show that external environmental factors can interact with plant–microbial feedbacks, with possible implications for plant community structure and dynamics.     

The interplay between soil structure,roots, and microbiota as a determinant of plant–soil feedback

Plant–soil feedback (PSF) can influence plant community structure via changes in the soil microbiome. However, how these feedbacks depend on the soil environment remains poorly understood. We hypothesized that disintegrating a naturally aggregated soil may influence the outcome of PSF by affecting microbial communities. Furthermore, we expected plants to differentially interact with soil structure and the microbial communities due to varying root morphology. We carried out a feedback experiment with nine plant species (five forbs and four grasses) where the “training phase” consisted of aggregated versus disintegrated soil. In the feedback phase, a uniform soil was inoculated in a fully factorial design with soil washings from conspecific‐ versus heterospecific‐trained soil that had been either disintegrated or aggregated. This way, the effects of prior soil structure on plant performance in terms of biomass production and allocation were examined. In the training phase, soil structure did not affect plant biomass. But on disintegrated soil, plants with lower specific root length (SRL) allocated more biomass aboveground. PSF in the feedback phase was negative overall. With training on disintegrated soil, conspecific feedback was positively correlated with SRL and significantly differed between grasses and forbs. Plants with higher SRL were likely able to easily explore the disintegrated soil with smaller pores, while plants with lower SRL invested in belowground biomass for soil exploration and seemed to be more susceptible to fungal pathogens. This suggests that plants with low SRL could be more limited by PSF on disintegrated soils of early successional stages. This study is the first to examine the influence of soil structure on PSF. Our results suggest that soil structure determines the outcome of PSF mediated by SRL. We recommend to further explore the effects of soil structure and propose to include root performance when working with PSF.     

Intraspecific and intergenerational differences in plant–soil feedbacks

Interactions between plant and soil communities are known to play an integral role in shaping ecosystems. Plants influence the composition of soil communities and soil communities in turn influence plant performance. Such a plant–soil feedback may incur selection pressure on plants and the associating soil community. However, the evolutionary consequences of these above–belowground feedback interactions remain largely speculative. Here we assess whether plant–soil feedback effects differ between intraspecific plant populations and between generations within the same plant population. We used two populations of Trifolium pratense and assessed their performance when grown in association with their home versus away soil biota. Both populations were colonized by distinct microbial communities and performed better with their own home soil communities than with the soil community from the other intraspecific population, demonstrating intraspecific positive feedback effects of home soil. In one of the two populations, we found that plant performance and the root associated microbiota community differed between parental and progeny plants when inoculated with their own home soil. Differences in root associated community characteristics could explain more than 80% of the variation in performance among the progeny and parental plants. Our results highlight that intraspecific differences in both plant and associated soil communities shape plant–soil feedback effects, and consequently indicate that plant–soil feedback can influence the direction of selection between intraspecific plant populations.     

Competition overwhelms the positive plant–soil feedback generated by an invasive plant

Invasive plant species can modify soils in a way that benefits their fitness more than the fitness of native species. However, it is unclear how competition among plant species alters the strength and direction of plant–soil feedbacks. We tested how community context altered plant–soil feedback between the non-native invasive forb Lespedeza cuneata and nine co-occurring native prairie species. In a series of greenhouse experiments, we grew plants individually and in communities with soils that differed in soil origin (invaded or uninvaded by L. cuneata) and in soils that were live vs. sterilized. In the absence of competition, L. cuneata produced over 60% more biomass in invaded than uninvaded soils, while native species performance was unaffected. The absence of a soil origin effect in sterile soil suggests that the positive plant–soil feedback was caused by differences in the soil biota. However, in the presence of competition, the positive effect of soil origin on L. cuneata growth disappeared. These results suggest that L. cuneata may benefit from positive plant–soil feedback when establishing populations in disturbed landscapes with few interspecific competitors, but does not support the hypothesis that plant–soil feedbacks influence competitive outcomes between L. cuneata and native plant species. These results highlight the importance of considering whether competition influences the outcome of interactions between plants and soils.     

Plant–soil feedbacks and the coexistence of competing plants

Plant–soil feedbacks can have important implications for the interactions among plants. Understanding these effects is a major challenge since it is inherently difficult to measure and manipulate highly diverse soil communities. Mathematical models may advance this understanding by making the interplay of the various processes affecting plant–soil interaction explicit and by quantifying the relative importance of the factors involved. The aim of this paper is to provide a complete analysis of a pioneering plant–soil feedback model developed by Bever and colleagues (J Ecol 85: 561–573, 1997; Ecol Lett 2: 52–62, 1999; New Phytol 157: 465–473, 2003) to fully understand the range of possible impacts of plant–soil feedbacks on plant communities within this framework. We analyze this model by means of a new graphical method that provides a complete classification of the potential effects of soil communities on plant competition. Due to the graphical character of the method, the results are relatively easy to obtain and understand. We show that plant diversity depends crucially on two key parameters that may be viewed as measures of the intensity of plant competition and the direction and strength of plant–soil feedback, respectively. Our analysis provides a formal underpinning of earlier claims that plant–soil feedbacks, especially when they are negative, may enhance the diversity of plant communities. In particular, negative plant–soil feedbacks can enhance the range of plant coexistence by inducing competitive oscillations. However, these oscillations can also destabilize plant coexistence, leading to low population densities and extinctions. In addition, positive feedbacks can allow locally stable forms of plant coexistence by inducing alternative stable states. Our findings highlight that the inclusion of plant–soil interactions may fundamentally alter the predictions on the structure and functioning of above-ground ecosystems. The scenarios presented in this study can be used to formulate hypotheses about the ways soil community effects may influence plant competition that can be tested with empirical studies. This will advance our understanding of the role of plant–soil feedback in ecological communities.     

Grazing‐induced changes in plant–soil feedback alter plant biomass allocation

Large vertebrate herbivores, as well as plant–soil feedback interactions are important drivers of plant performance, plant community composition and vegetation dynamics in terrestrial ecosystems. However, it is poorly understood whether and how large vertebrate herbivores and plant–soil feedback effects interact. Here, we study the response of grassland plant species to grazing‐induced legacy effects in the soil and we explore whether these plant responses can help us to understand long‐term vegetation dynamics in the field. In a greenhouse experiment we tested the response of four grassland plant species, Agrostis capillaris, Festuca rubra, Holcus lanatus and Rumex acetosa, to field‐conditioned soils from grazed and ungrazed grassland. We relate these responses to long‐term vegetation data from a grassland exclosure experiment in the field. In the greenhouse experiment, we found that total biomass production and biomass allocation to roots was higher in soils from grazed than from ungrazed plots. There were only few relationships between plant production in the greenhouse and the abundance of conspecifics in the field. Spatiotemporal patterns in plant community composition were more stable in grazed than ungrazed grassland plots, but were not related to plant–soil feedbacks effects and biomass allocation patterns. We conclude that grazing‐induced soil legacy effects mainly influenced plant biomass allocation patterns, but could not explain altered vegetation dynamics in grazed grasslands. Consequently, the direct effects of grazing on plant community composition (e.g. through modifying light competition or differences in grazing tolerance) appear to overrule indirect effects through changes in plant–soil feedback.     

Plant–mycorrhizal fungus co‐occurrence network lacks substantial structure

The interactions between plants and arbuscular mycorrhizal fungi (AMF) maintain a crucial link between macroscopic organisms and the soil microbial world. These interactions are of extreme importance for the diversity of plant communities and ecosystem functioning. Despite this importance, only recently has the structure of plant–AMF interaction networks been studied. These recent studies, which used genetic data, suggest that these networks are highly structured, very similar to plant–animal mutualistic networks. However, the assembly process of plant–AMF communities is still largely unknown, and an important feature of plant–AMF interactions has not been incorporated: they occur at an extremely localized scale. Studying plant–AMF networks in a spatial context seems therefore a crucial step. This paper studies a plant–AMF spatial co‐occurrence network using novel methodology based on information theory and a unique set of spatially explicit species‐level data. We apply three null models of which only one accounts for spatial effects. We find that the data show substantial departures from null expectations for the two non‐spatial null models. However, for the null model considering spatial effects, there are few significant co‐occurrences compared with the other two null models. Thus, plant–AMF spatial co‐occurrences seem to be mostly explained by stochasticity, with a small role for other factors related to plant–AMF specialization. Furthermore, we find that the network is not significantly nested or modular. We conclude that this plant–AMF spatial co‐occurrence network lacks substantial structure and, therefore, plants and AMF species do not track each other over space. Thus, random encounters seem more important in the first step of the assembly of plant–AMF communities. Synthesis The symbiotic interaction between plants and arbuscular mycorrhizal fungi (AMF) is crucial for ecosystem functioning. However, the factors affecting the assembly of plant‐AMF communities are poorly understood. An important factor of the assembly of plant‐AMF communities has been overlooked: plant‐AMF interactions occur at a localized spatial scale. Our study investigated the importance of space in the structure of plant‐AMF communities. We studied a plant‐AMF spatial co‐occurrence network using a unique set of spatially explicit data and applied three null models. We found that plant‐AMF spatial co‐occurrences seem to be mostly explained by stochasticity. In particular, our study shows that this plant‐AMF spatial co‐occurrence network lacks substantial structure and, therefore, plants and AMF species do not track each other over space. Thus, random encounters seem to drive the assembly of plant‐AMF communities.     

Species with greater seed mass are more tolerant of conspecific neighbours: a key driver of early survival and future abundances in a tropical forest

Multiple niche‐based processes including conspecific negative density dependence (CNDD) determine plant regeneration and community structure. We ask how interspecific and intraspecific density‐dependent interactions relate to plant life histories and associated functional traits. Using hierarchical models, we analysed how such interactions affected first‐year survival of seedling recruits of 175 species in a tropical forest, and how species abundances and functional traits are related to interspecific variation in density‐dependent effects. Conspecific seedling neighbour effects prevailed over the effects of larger conspecific and all heterospecific neighbours. Tolerance of seedling CNDD enhanced recruit survival and subsequent abundance, all of which were greater among larger seeded, slow‐growing and well‐defended species. Niche differentiation along the growth–survival trade‐off and tolerance of seedling CNDD strongly correlated with regeneration success, with manifest consequences for community structure. The ability of larger seeded species to better tolerate CNDD suggests a novel mechanism for CNDD to contribute to seed‐size variation and promote species coexistence through a tolerance–fecundity trade‐off.     

Individual species–area relationships and spatial patterns of species diversity in a Great Basin,semi‐arid shrubland

Traditional biodiversity metrics operate at the level of a plant community but do not capture spatial variation in diversity from a ‘plant's‐eye view’ of a community. Recently‐developed statistics consider the spatial patterns of plants as well as the number and distribution of species in local plant neighborhoods to quantitatively assess multispecies spatial patterns from a ‘plant's‐eye view’. We used one such statistic, the individual species area relationship (ISAR), to assess spatial patterns of species diversity in a Great Basin (USA) semi‐arid shrubland through an analysis of a spatial dataset on shrub species and locations. In conjunction with appropriate null models, the ISAR blends species area relationships with second‐order spatial statistics to measure the expected species richness in local neighborhoods of variable size around the individuals of a focal species within a community. We found that, contrary to a previous analysis using more traditional methods, the community was well‐mixed with a typical shrub surrounded on average by 4.9 shrub neighbors of 2.1 species at a neighborhood scale of 1.0 m. We also found statistically significant fine‐scale variation in diversity patterns, such that neighborhoods of two species were more diverse than expected by a heterogeneous Poisson null model that accounted for larger‐scale habitat heterogeneity. However, this effect was caused by intraspecific aggregation of these species and was not due to positive interspecific association. Contrary to previous findings in other semi‐arid shrublands, our analysis suggests that the spatial pattern of the shrub community was not significantly structured by interspecific facilitation. This result supports growing evidence for balanced species patterns of adult plants in multispecies communities. Our approach may be used in other communities to describe complex multispecies spatial patterns, quantify species‐specific associations with diversity patterns, and to generate hypotheses regarding relationships between patterns and community‐structuring processes.     

Effects of exotic annual grass litter and local environmental gradients on annual plant community structure

Exotic annual grasses have been introduced into many semi-arid ecosystems worldwide, often to the detriment of native plant communities. The accumulation of litter from these grasses (i.e. residual dry biomass) has been demonstrated to negatively impact native plant communities and promote positive feedbacks to exotic grass persistence. More targeted experiments are needed, however, to determine the relative impact of exotic grass litter on plant community structure across local environmental gradients. We experimentally added exotic grass litter to annual forb-dominated open woodland communities positioned along natural canopy cover gradients in southwest Western Australia. These communities are an important component of this region’s plant biodiversity hotspot and are documented to be under threat from exotic annual grasses. After a one-year treatment period, we measured the effects of exotic grass litter, soil properties, and canopy cover on native and exotic species richness and abundance, as well as common species’ biomass and abundances. Plant community structure was more strongly influenced by soil properties and canopy cover than by grass litter. Total plant abundances per plot, however, were significantly lower in litter addition plots than control plots, a trend driven by native species. Exotic grass litter was also associated with lower abundances of one very common native species: Waitzia acuminata. Our results suggest that exotic grass litter limits the establishment of some native species in this system. Over multiple years, these subtle impacts may contribute substantially to the successful advancement of exotic species into this system, particularly in certain microenvironments.     

Species interactions and thermal constraints on ant community structure

Patterns of species occurrence and abundance are influenced by abiotic factors and biotic interactions, but these factors are difficult to disentangle without experimental manipulations. In this study, we used observational and experimental approaches to investigate the role of temperature and interspecific competition in controlling the structure of ground‐foraging ant communities in forests of the Siskiyou Mountains of southwestern Oregon. To assess the potential role of competition, we first used null model analyses to ask whether species partition temporal and/or spatial environments. To understand how thermal tolerances influence the structure of communities, we conducted a laboratory experiment to estimate the maximum thermal tolerance of workers and a field experiment in which we added shaded microhabitats and monitored the response of foragers. Finally, to evaluate the roles of temperature and interspecific competition in the field, we simultaneously manipulated shading and the presence of a dominant competitor (Formica moki). The foraging activity of species broadly overlapped during the diurnal range of temperatures. Species co‐occurrence patterns varied across the diurnal temperature range: species were spatially segregated at bait stations at low temperatures, but co‐occurred randomly at high temperatures. The decreased abundance of the co‐occurring thermophilic Temnothorax nevadensis in shaded plots was a direct effect of shading and not an indirect effect of competitive interactions. Thermal tolerance predicted the response of ant species to the shading experiment: species with the lowest tolerances to high temperatures showed the greatest increase in abundance in the shaded plots. Moreover, species with more similar thermal tolerance values segregated more frequently on baits than did species that differed in their thermal tolerances. Collectively, our results suggest that thermal tolerances of ants may mediate competitive effects in habitats that experience strong diurnal temperature fluctuations.