Coexistence and relative abundance in annual plant assemblages: the roles of competition and colonization

Although an interspecific trade-off between competitive and colonizing ability can permit multispecies coexistence, whether this mechanism controls the structure of natural systems remains unresolved. We used models to evaluate the hypothesized importance of this trade-off for explaining coexistence and relative abundance patterns in annual plant assemblages. In a nonspatial model, empirically derived competition-colonization trade-offs related to seed mass were insufficient to generate coexistence. This was unchanged by spatial structure or interspecific variation in the fraction of seeds dispersing globally. These results differ from those of the more generalized competition-colonization models because the latter assume completely asymmetric competition, an assumption that appears unrealistic considering existing data for annual systems. When, for heuristic purposes, completely asymmetric competition was incorporated into our models, unlimited coexistence was possible. However, in the resulting abundance patterns, the best competitors/poorest colonizers were the most abundant, the opposite of that observed in natural systems. By contrast, these natural patterns were produced by competition-colonization models where environmental heterogeneity permitted species coexistence. Thus, despite the failure of the simple competition-colonization trade-off to explain coexistence in annual plant systems, this trade-off may be essential to explaining relative abundance patterns when other processes permit coexistence.     

Restoration through reassembly: plant traits and invasion resistance

One of the greatest challenges for ecological restoration is to create or reassemble plant communities that are resistant to invasion by exotic species. We examine how concepts pertaining to the assembly of plant communities can be used to strengthen resistance to invasion in restored communities. Community ecology theory predicts that an invasive species will be unlikely to establish if there is a species with similar traits present in the resident community or if available niches are filled. Therefore, successful restoration efforts should select native species with traits similar to likely invaders and include a diversity of functional traits. The success of trait-based approaches to restoration will depend largely on the diversity of invaders, on the strength of environmental factors and on dispersal dynamics of invasive and native species.     

Disturbance affects short-term facilitation, but not long-term saturation, of exotic plant invasion in New Zealand forest

We investigate the spread of an exotic herb, Hieracium lepidulum, into a New Zealand Nothofagus forest with the aim of understanding how stand-development of tree populations, propagule pressure and invader persistence, affect invasion across the landscape and within communities. Using data repeatedly collected over 35 years, from 250 locations, we parametrize continuous-time Markov chain models and use these models to examine future projections of the invasion under a range of hypothetical scenarios. We found that the probability of invasion into a stand was relatively high following canopy disturbance and that local abundance of Hieracium was promoted by minor disturbances. However, model predictions extrapolated 45 years into the future show that neither the rate of landscape-level invasion, nor local population growth of Hieracium, was affected much by changing the frequency of canopy disturbance events. Instead, invasion levels were strongly affected by the ability of Hieracium to persist in the understorey following forest canopy closure, and by propagule supply from streams, forest edges and plants already established within the stand. Our results show that disturbance frequency has surprisingly little influence on the long-term trajectory of invasion, while invader persistence strongly determines invasion patterns.     

Abiotic and biotic resistance to grass invasion in serpentine annual plant communities

Biological invasions severely impact native plant communities, causing dramatic shifts in species composition and the restriction of native species to spatially isolated refuges. Competition from resident species and the interaction between resource limitation and competition have been overlooked as mechanisms of community resistance in refugia habitats. We examined the importance of these factors in determining the resistance of California serpentine plant communities to invasion by three common European grasses, Avena barbata, Bromus diandrus, and Hordeum murinum. We added seeds of each of these grasses to plots subjected to six levels of resource addition (N, P, Ca, H₂O, all resources together, and a no-addition control) and two levels of competition (with resident community present or removed). Resource limitation and competition had strong effects on the biomass and reproduction of the three invaders. The addition of all resources together combined with the removal of the resident community yielded individual plants that were fourfold to 20-fold larger and sixfold to 20-fold more fecund than plants from control plots. Competitor removal alone yielded invaders that were twofold to sevenfold larger and twofold to ninefold more fecund. N addition alone or in combination with other resources led to a twofold to ninefold increase in the biomass and fecundity of the invaders. No other resource alone significantly affected native or invader performance, suggesting that N was the key limiting resource during our experiment. We found a significant interaction between abiotic and biotic resistance for Bromus, which experienced increased competitive suppression in fertilized plots. The threefold increase in resident biomass with N addition was likely responsible for this result. Our results confirm that serpentine plant communities are severely N limited, which, in combination with competition from resident species, promotes the resistance of these systems to invasions. Our work suggests that better understanding the relative sensitivities of invaders and residents to the physical environment is critical to predicting how abiotic and biotic factors interact to determine community resistance.     

Plant diversity,herbivory and resistance of a plant community to invasion in Mediterranean annual communities

Several components of the diversity of plant communities, such as species richness, species composition, number of functional groups and functional composition, have been shown to directly affect the performance of exotic species. Exotics can also be affected by herbivores of the native plant community. However, these two possible mechanisms limiting invasion have never been investigated together. The aim of this study was to investigate the relationships between plant diversity, herbivory and performance of two annual exotics, Conyza bonariensis and C. canadensis, in Mediterranean annual communities. We wanted to test whether herbivory of these exotics was influenced either by species richness, functional-group richness or functional-group composition. We also studied the relationship between herbivory on the exotic species and their performance. Herbivory increased with increasing species and functional-group richness for both Conyza species. These patterns are interpreted as reflecting a greater number of available herbivore niches in a richer, more complex, plant community. The identities of functional groups also affected Conyza herbivory, which decreased in the presence of Asteraceae or Fabaceae and increased in the presence of Poaceae. Increasing herbivory had consequences for vegetative and demographic parameters of both invasive species: survival, final biomass and net fecundity decreased with increasing herbivory, leading to a loss of reproductive capacity. We conclude that communities characterised by a high number of grass species instead of Asteraceae or Fabaceae may be more resistant to invasion by the two Conyza species, in part due to predation by native herbivores.     

The importance of biological inertia in plant community resistance to invasion

Abstract. Insights into the ecology of historic invasions by introduced species can be gained by studying long‐term patterns of invasions by native species. In this paper, we review literature in palaeo‐ecology, forest‐stand simulation modelling, and historical studies of plant species invasions to illustrate the relevance of biological inertia in plant communities to invasion ecology. Resistance to invasion occurs in part because of environmental, demographic, and biotic factors influencing the arrival and establishment of invading species. We propose that biological inertia within the resident community is a fourth component of resistance to invasion, because of the lag time inherent in eliminating resident species and perhaps their traces after environmental conditions become suitable for invasion by immigrating species. Whether or not an introduced species invades can be conditioned by the presence of the pre‐existing community (and/or its legacy) in addition to the other biotic and abiotic factors.     

Exotic plant invasion alters chaparral ecosystem resistance and resilience pre- and post-wildfire

Perturbations such as wildfire and exotic plant invasion have significant impacts on soils, and the extent to which invaded soils are resistant or resilient to these disturbances varies by ecosystem type. Replacement of shrublands by herbaceous exotics pre- and post-wildfire may drastically alter soil chemical and biological properties for an unknown duration. We assessed above and belowground resistance and resilience to exotic plant invasion both before and after a chaparral wildfire. We hypothesized that exotic plant species would change chemical characteristics of chaparral soils by altering litter and microbial inputs, and that controlling exotics and seeding native species would restore chemical characteristics to pre-invaded conditions. We additionally hypothesized that exotic plant species would slow succession above- and belowground, as well as recovery of post-wildfire chaparral structure and function. Plant species composition and soil nutrient pools and cycling rates were evaluated in mature and invaded chaparral pre- and post-wildfire. Exotic plant species were weeded and native species were seeded to assess impacts of exotic competition on native species recovery. Invasion did not impact all soil characteristics before fire, but increased soil C/N ratio, pH, and N cycling rates, and reduced NO₃-N availability. After fire, invasives slowed succession above- and belowground. Removal of exotics and seeding natives facilitated succession and resulted in plant composition similar to uninvaded, post-wildfire chaparral. The chaparral ecosystem was not resistant to impacts of invasion as indicated by altered soil chemistry and C and N cycling rates; however, short-term restoration led to recovery of extractable nitrogen availability indicating resilience of chaparral soils. This suggests that the permanence of exotic plant species, once established, represents a greater ecological challenge than exotic plant impacts on soils.     

Hierarchical reliability in experimental plant assemblages

Aims: The mechanism by which species richness affects variation inecosystem functioning both within and among ecosystems remainsa key question at the interface of community and ecosystem ecology.Statistical averaging (the smoothing of average system performancevia consideration of additional components) and the insuranceeffect (reduced variation in system performance by inclusionof asynchronously varying components) predict that more diversecommunities should vary less both between replicates and internally.We experimentally tested these theories in small plant assemblages. Methods: We constructed plant assemblages modeled after old-field plantcommunities. We varied species richness, species compositionand initial densities while holding functional group richnessconstant in replicate assemblages under glasshouse conditions. Important findings: The inverse of the coefficient of variation of aboveground biomassproduction, a proxy measure of reliability, increased with higherdiversity when examined at the level of the assemblage (i.e.among-replicate assemblages) but not at the levels of functionalgroup or species. These stabilizing processes were weakest inlow-diversity, low-density assemblages. This experiment demonstratesthe utility of hierarchical analysis of ecosystem reliabilityat the assemblage, functional group and species level.     

Intraspecific variation in plant size,secondary plant compounds,herbivory and parasitoid assemblages during secondary succession

During secondary succession on abandoned agricultural fields the diversity and abundance of insect communities often increases, whereas the performance and nutritional quality of early successional plants often declines. As the diversity and abundance of insects on a single plant are determined by characteristics of the environment as well as of the host plant, it is difficult to predict how insects associated with a single plant species will change during succession. We examined how plant characteristics of the early successional plant species ragwort (Jacobaea vulgaris), and the herbivores and parasitoids associated with these plants change during secondary succession. In ten grasslands that differed in time since abandonment (3–26 years), we measured the size and primary and secondary chemistry of individual ragwort plants. For each plant we also recorded the presence of herbivores in flowers, leaves and stems, and reared parasitoids from these plant parts. Ragwort plants were significantly larger but had lower nitrogen concentrations in recently abandoned sites than in older sites. Pyrrolizidine alkaloid (PA) composition varied among plants within sites but also differed significantly among sites. However, there was no relationship between the age of a site and PA composition. Even though plant size decreased with time since abandonment, the abundance of stem-boring insects and parasitoids emerging from stems significantly increased with site age. The proportion of plants with flower and leaf herbivory and the number of parasitoids emerging from flowers and leaves was not related to site age. Parasitoid diversity significantly increased with site age. The results of our study show that ragwort and insect characteristics both change during secondary succession, but that insect herbivore and parasitoid abundances are not directly related to plant size or nutritional quality.     

Evidence of biotic resistance to exotic plant invasion in degraded Bornean forests

Intact tropical forests are generally considered to be resistant to invasions by exotic species, although the shrub Clidemia hirta (Melastomataceae) is highly invasive in tropical forests outside its native range. Release from natural enemies (e.g., herbivores and pathogens) contributes to C. hirta invasion success where native melastomes are absent, and here we examine the role of enemies when C. hirta co-occurs with native Melastomataceae species and associated herbivores and pathogens. We study 21 forest sites within agricultural landscapes in Sabah, Malaysian Borneo, recording herbivory rates in C. hirta and related native Melastoma spp. plants along two 100-m transects per site that varied in canopy cover. Overall, we found evidence of enemy release; C. hirta had significantly lower herbivory (median occurrence of herbivory per plant = 79% of leaves per plant; median intensity of herbivory per leaf = 6% of leaf area) than native melastomes (93% and 20%, respectively). Herbivory on C. hirta increased when closer to native Melastoma plants with high herbivory damage, and in more shaded locations, and was associated with fewer reproductive organs on C. hirta. This suggests host-sharing by specialist Melastomataceae herbivores is occurring and may explain why invasion success of C. hirta is lower on Borneo than at locations without related native species present. Thus, natural enemy populations may provide a “biological control service” to suppress invasions of exotic species (i.e., biotic resistance). However, lower herbivory pressures in more open canopy locations may make highly degraded forests within these landscapes more susceptible to invasion.     

Foliar resistance to simulated extreme temperature events in contrasting plant functional and chorological types

We analysed leaf resistance of 41 Angiosperms belonging to a wide range of plant functional (PFTs) and chorological types (PCTs) to simulated frost and high‐temperature extreme events (EE). Leaf resistance was estimated as percentage of membrane electrolyte leakage under heating and freezing treatments in the lab. Leaf resistance to heating or freezing was not significantly correlated with the main resource‐use characteristics that defined PFTs, such as leaf specific area, toughness, N concentration or thickness. Leaf resistance to heating differed among PFTs (graminoids and bromeliads were the more resistant groups), but not among PCTs. In contrast, leaf resistance to freezing significantly differed among PCTs. Along a steep regional climatic gradient, climate variables (annual mean temperature, mean minimum temperature, mean maximum temperature and number of frost‐free months) at the locations where the given species were most abundant were also significantly correlated with freezing resistance. Species from colder habitats both at the sub‐continental and regional scales showed the highest leaf resistance to freezing. Our work indicates that leaf resistance to climatic EE and resource‐use strategy (assessed in previous studies) represent two different, partially decoupled axes of plant specialisation. It also suggests that changes in the frequency of very low temperature events might have regional‐scale impacts on vegetation, whereas changes in the frequency of very high temperature events might have more influence at the local scale.     

Rhizosphere microbiome functional diversity and pathogen invasion resistance build up during plant development

The rhizosphere microbiome is essential for plant growth and health, and numerous studies have attempted to link microbiome functionality to species and trait composition. However, to date little is known about the actual ecological processes shaping community composition, complicating attempts to steer microbiome functionality. Here, we assess the development of microbial life history and community-level species interaction patterns that emerge during plant development. We use microbial phenotyping to experimentally test the development of niche complementarity and life history traits linked to microbiome performance. We show that the rhizosphere microbiome assembles from pioneer assemblages of species with random resource overlap into high-density, functionally complementary climax communities at later stages. During plant growth, fast-growing species were further replaced by antagonistic and stress-tolerant ones. Using synthetic consortia isolated from different plant growth stages, we demonstrate that the high functional diversity of ‘climax’ microbiomes leads to a better resistance to bacterial pathogen invasion. By demonstrating that different life-history strategies prevail at different plant growth stages and that community-level processes may supersede the importance of single species, we provide a new toolbox to understand microbiome assembly and steer its functionality at a community level.     

Ecological application of biotic resistance to control the invasion of an invasive plant,Ageratina altissima

Biotic resistance is the ability of species in a community to limit the invasion of other species. However, biotic resistance is not widely used to control invasive plants. Experimental, functional, and modeling approaches were combined to investigate the processes of invasion by Ageratina altissima (white snakeroot), a model invasive species in South Korea. We hypothesized that (1) functional group identity would be a good predictor of biotic resistance to A. altissima, whereas a species identity effect would be redundant within a functional group, and (2) mixtures of species would be more resistant to invasion than monocultures. We classified 37 species of native plants into three functional groups based on seven functional traits. The classification of functional groups was based primarily on differences in life longevity and woodiness. A competition experiment was conducted based on an additive competition design with A. altissima and monocultures or mixtures of resident plants. As an indicator of biotic resistance, we calculated a relative competition index (RCI_avg) based on the average performance of A. altissima in a competition treatment compared with that of the control where only seeds of A. altissima were sown. To further explain the effect of diversity, we tested several diversity–interaction models. In monoculture treatments, RCI_avg of resident plants was significantly different among functional groups but not within each functional group. Fast‐growing annuals (FG1) had the highest RCI_avg, suggesting priority effects (niche pre‐emption). RCI_avg of resident plants was significantly greater in a mixture than in a monoculture. According to the diversity–interaction models, species interaction patterns in mixtures were best described by interactions between functional groups, which implied niche partitioning. Functional group identity and diversity of resident plant communities were good indicators of biotic resistance to invasion by introduced A. altissima, with the underlying mechanisms likely niche pre‐emption and niche partitioning. This method has most potential in assisted restoration contexts, where there is a desire to reintroduce natives or boost their population size due to some previous level of degradation.     

Allelopathy and exotic plant invasion

The primary hypothesis for the astonishing success of many exotics as community invaders relative to their importance in their native communities is that they have escaped the natural enemies that control their population growth – the `natural enemies hypothesis'. However, the frequent failure of introduced biocontrols, weak consumer effects on the growth and reproduction of some invaders, and the lack of consistent strong top-down regulation in many natural ecological systems indicate that other mechanisms must be involved in the success of some exotic plants. One mechanism may be the release by the invader of chemical compounds that have harmful effects on the members of the recipient plant community (i.e., allelopathy). Here, we provide an abbreviated compilation of evidence for allelopathy in general, present a detailed case study for Centaurea diffusa, an invasive Eurasian forb in western North America, and review general evidence for allelopathic effects of invasive plants in native communities. The primary rationale for considering allelopathy as a mechanism for the success of invaders is based on two premises. First, invaders often establish virtual monocultures where diverse communities once flourished, a phenomenon unusual in natural communities. Second, allelopathy may be more important in recipient than in origin communities because the former are more likely to be naïve to the chemicals possessed by newly arrived species. Indeed, results from experiments on C. diffusa suggest that this invader produces chemicals that long-term and familiar Eurasian neighbors have adapted to, but that C. diffusa's new North American neighbors have not. A large number of early studies demonstrated strong potential allelopathic effects of exotic invasive plants; however, most of this work rests on controversial methodology. Nevertheless, during the last 15 years, methodological approaches have improved. Allelopathic effects have been tested on native species, allelochemicals have been tested in varying resource conditions, models have been used to estimate comparisons of resource and allelopathic effects, and experimental techniques have been used to ameliorate chemical effects. We do not recommend allelopathy as a `unifying theory' for plant interactions, nor do we espouse the view that allelopathy is the dominant way that plants interact, but we argue that non-resource mechanisms should be returned to the discussion table as a potential mechanism for explaining the remarkable success of some invasive species. Ecologists should consider the possibility that resource and non-resource mechanisms may work simultaneously, but vary in their relative importance depending on the ecological context in which they are studied. One such context might be exotic plant invasion.     

Genomes, free radicals and plant cell invasion: recent developments in plant pathogenic fungi

This review describes current advances in our understanding of fungal-plant interactions. The widespread application of whole genome sequencing to a diverse range of fungal species has allowed new insight into the evolution of fungal pathogenesis and the definition of the gene inventories associated with important plant pathogens. This has also led to functional genomic approaches to carry out large-scale gene functional analysis. There has also been significant progress in understanding appressorium-mediated plant infection by fungi and its underlying genetic basis. The nature of biotrophic proliferation of fungal pathogens in host tissue has recently revealed new potential mechanisms for cell-to-cell movement by invading pathogens.     

Cyclic dynamics in simulated plant populations

Despite the general interest in nonlinear dynamics in animal populations, plant populations are supposed to show a stable equilibrium that is attributed to fundamental differences compared with animals. Some studies find more complex dynamics, but empirical studies usually are too short and most modelling studies ignore important spatial aspects of local competition and establishment. Therefore, we used a spatially explicit individual-based model of a hypothetical, non-clonal perennial to explore which mechanisms might generate complex dynamics, i.e. cycles. The model is based on the field-of-neighbourhood approach that describes local competition and establishment in a phenomenological manner. We found cyclic population dynamics for a wide spectrum of model variants, provided that mortality is determined by local competition and recruitment is virtually completely suppressed within the zone of influence of established plants. This destabilizing effect of local processes within plant populations might have wide-ranging implications for the understanding of plant community dynamics and coexistence.