A meta-analysis of biotic resistance to exotic plant invasions

Biotic resistance describes the ability of resident species in a community to reduce the success of exotic invasions. Although resistance is a well‐accepted phenomenon, less clear are the processes that contribute most to it, and whether those processes are strong enough to completely repel invaders. Current perceptions of strong, competition‐driven biotic resistance stem from classic ecological theory, Elton's formulation of ecological resistance, and the general acceptance of the enemies‐release hypothesis. We conducted a meta‐analysis of the plant invasions literature to quantify the contribution of resident competitors, diversity, herbivores and soil fungal communities to biotic resistance. Results indicated large negative effects of all factors except fungal communities on invader establishment and performance. Contrary to predictions derived from the natural enemies hypothesis, resident herbivores reduced invasion success as effectively as resident competitors. Although biotic resistance significantly reduced the establishment of individual invaders, we found little evidence that species interactions completely repelled invasions. We conclude that ecological interactions rarely enable communities to resist invasion, but instead constrain the abundance of invasive species once they have successfully established.     

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.     

Abiotic and biotic contributions to invasion resistance for ornamental fish in west-central Florida,USA

Hydrobiologia - Explaining the varying success of invaders is a central question in invasion biology. Florida is a good region to test hypotheses related to invasion because of variation in...     

The physical resistance of grass patches to invasion

One of the important factors determining success in plant competition is the ability of a plant to extend laminae in order to capture resources.To do this in mixed swards the laminae of one plant must first grow into the volume that contains laminae of another. The ability of laminae to overcome the resistance presented by a neighbour, and the ability to resist this ingress, was examined for the grasses Agrostis capillaris, Festuca rubra, Holcus lanatus, Lolium perenne and Poa trivialis that were subject to 3 cm and 6 cm cutting treatments. These abilities were inferred from the behaviour of ‘indicator ’ leaves as they were pushed into monoculture target patches of each species. The 3 cm treatment resisted ingress significantly more than the 6 cm. Species patches differed significantly both in their ability to resist the ingress by the indicator species and in the ability of different indicator species to penetrate the target swards. These effects were still present when differences in leaf density (leaves cm^–2) had been taken into account. The results suggest that grasses can vary in the physical resistance that they present to the leaves of an invading neighbour.     

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.     

The role of reproductive plant traits and biotic interactions in the dynamics of semi-arid plant communities

The dynamics of semi-arid plant communities are determined by the interplay between competition and facilitation among plants. The sign and strength of these biotic interactions depend on plant traits. However, the relationships between plant traits and biotic interactions, and the consequences for plant communities are still poorly understood. Our objective here was to investigate, with a modelling approach, the role of plant reproductive traits on biotic interactions, and the consequences for processes such as plant succession and invasion. The dynamics of two plant types were modelled with a spatially-explicit integrodifferential model: (1) a plant with seed dispersal (colonizer of bare soil) and (2) a plant with local vegetative propagation (local competitor). Both plant types were involved in facilitation due to a local positive feedback between vegetation biomass and soil water availability, which promoted establishment and growth. Plants in the system also competed for limited water. The efficiency in water acquisition (dependent on reproductive and growth plant traits) determined which plant type dominated the community at the steady state. Facilitative interactions between plant types also played an important role in the community dynamics, promoting establishment in the driest conditions and recovery from low biomass. Plants with vegetative propagation took advantage of the ability of seed dispersers to establish on bare soil from a low initial biomass. Seed dispersers were good invaders, maintained high biomass at intermediate and high rainfall and showed a high ability in taking profit from the positive feedback originated by plants with vegetative propagation under the driest conditions. However, seed dispersers lost competitiveness with an increasing investment in fecundity. All together, our results showed that reproductive plant traits can affect the balance between facilitative and competitive interactions. Understanding this effect of plant traits on biotic interactions provides insights in processes such as plant succession and shrub encroachment.     

Positive interactions, discontinuous transitions and species coexistence in plant communities

The population and community level consequences of positive interactions between plants remain poorly explored. In this study we incorporate positive resource-mediated interactions in classic resource competition theory and investigate the main consequences for plant population dynamics and species coexistence. We focus on plant communities for which water infiltration rates exhibit positive dependency on plant biomass and where plant responses can be improved by shading, particularly under water limiting conditions. We show that the effects of these two resource-mediated positive interactions are similar and additive. We predict that positive interactions shift the transition points between different species compositions along environmental gradients and that realized niche widths will expand or shrink. Furthermore, continuous transitions between different community compositions can become discontinuous and bistability or tristability can occur. Moreover, increased infiltration rates may give rise to a new potential coexistence mechanism that we call controlled facilitation.     

Species richness and evenness respond in a different manner to propagule density in developing prairie microcosm communities

Diversity has two basic components: richness, or number of species in a given area, and evenness, or how relative abundance or biomass is distributed among species. Previously, we found that richness and evenness can be negatively related across plant communities and that evenness can account for more variation in Shannon’s diversity index (H′) than richness, which suggests that relationships among diversity components can be complex. Non-positive relationships between evenness and richness could arise due to the effects of migration rate or local species interactions, and relationships could vary depending on how these two processes structure local communities. Here we test whether diversity components are equally or differentially affected over time by changes in seed density (and associated effects on established plant density and competition) in greenhouse communities during the very early stages of community establishment. In our greenhouse experiment, we seeded prairie microcosms filled with bare field soil at three densities with draws from a mix of 22 grass and forb species to test if increased competition intensity or seedling density would affect the relationships among diversity components during early community establishment. Increased seed density treatments caused diversity components to respond in a different manner and to have different relationships with time. Richness increased linearly with seed density early in the experiment when seedling emergence was high, but was unrelated to density later in the experiment. Evenness decreased log-linearly with seed densities on all sampling dates due to a greater dominance by Rudbeckia hirta with higher densities. Early in the experiment, diversity indices weakly reflected differences in richness, but later, after the competitive effects of Rudbeckia hirta became more intense, diversity indices more strongly reflected differences in evenness. This suggests that species evenness and diversity indices do not always positively covary with richness. Based on these results, we suggest that evenness and richness can be influenced by different processes, with richness being more influenced by the number of emerging seedlings and evenness more by species interactions like competition. These results suggest that both diversity components should be measured in plant diversity studies whenever it is possible.     

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.     

Response of annual plants and communities to tilling in a semi-arid temperate savanna

Abstract. Despite the obvious importance of savannas, the response of semi-arid temperate savannas to disturbance has hardly been studied. The objective of this study was to determine the response of annual plants and vegetation to severe disturbance (tilling) in the grassland phase of a semi-arid savanna characterized by perennial bunchgrasses. Most species of annual plants responded positively to tilling; however, 25% of the species did not respond or responded negatively. Response of plants and vegetation was more strongly influenced by precipitation than disturbance, even for the severe disturbance used in this study. Tilling-induced differences were negligible during a relatively dry year. Response of annuals to disturbance in semi-arid temperate savannas may not be consistent with response in other systems.     

A theory of the spatial and temporal dynamics of plant communities

An individual-based model of plant competition for light that uses a definition of plant functional types based on adaptations for the simultaneous use of water and light can reproduce the fundamental spatial and temporal patterns of plant communities. This model shows that succession and zonation result from the same basic processes. Succession is interpreted as a temporal shift in species dominance, primarily in response to autogenic changes in light availability. Zonation is interpreted as a spatial shift in species dominance, primarily in response to the effect of allogenic changes in water availability on the dynamics of competition for light. Patterns of succession at different points along a moisture gradient can be used to examine changes in the ecological roles of various functional types, as well as to address questions of shifts in patterns of resource use through time.Our model is based on the cost-benefit concept that plant adaptations for the simultaneous use of two or more resources are limited by physiological and life history constraints. Three general sets of adaptive constraints produce inverse correlations in the ability of plants to efficiently use (1) light at both high and low availability, (2) water at both high and low availability, and (3) both water and light at low availabilities.The results of this type of individual-based model can be aggregated to examine phenomena at several levels of system organization (i.e., subdisciplines of ecology), including (1) plant growth responses over a range of environmental conditions, (2) population dynamics and size structure, (3) experimental and field observations on the distribution of species across environmental gradients, (4) studies of successional pattern, (5) plant physiognomy and community structure across environmental gradients, and (6) nutrient cycling.     

Responses of plant and bird communities to prescribed burning in tallgrass prairies

Historic losses and fragmentation of tallgrass prairie habitat to agriculture and urban development have led to declines in diversity and abundance of plants and birds associated with such habitat. Prescribed burning is a management strategy that has potential for restoring and rejuvenating prairies in fragmented landscapes, and through such restoration, might create habitat for birds dependent upon prairies. To provide improved data for management decision-making regarding the use of prescribed fire in tallgrass prairies, we compared responses of plant and bird communities on five burned and five unburned tallgrass prairie fragments at the DeSoto National Wildlife Refuge, Iowa, USA, from 1995 to 1997. Overall species richness and diversity were unaffected by burning, but individual species of plants and birds were affected by year-treatment interactions, including northern bobwhite (Colinus virginianus) and ring-necked pheasant (Phasianus colchicus), which showed time-delayed increases in density on burned sites. Analyses of species/area relationships indicated that, collectively, many small sites did make significant contributions to plant biodiversity at landscape levels, supporting the overall conservation value of prairie fragments. In contrast, most birds species were present on larger sites. Thus, higher biodiversity in bird communities which contain area-sensitive species might require larger sites able to support larger, more stable populations, greater habitat heterogeneity, and greater opportunity for niche separation.     

Community‐level consequences of species interactions in an annual plant community

Question: How does the intensity of species interactions affect species and functional group composition of an annual plant community? Location: Sede Boqer, Negev Desert, Israel. Methods: The potential for competitive interactions in two annual plant communities (desert and coastal) from semi‐stabilized sand dunes was manipulated by varying seed bank density and therefore the number of potentially interacting individuals. Communities were exposed to three different irrigation regimes, mimicking precipitation at the desert site, the coastal site, and an intermediate precipitation level. Plots were maintained for 3 years, and percentage cover of each species in the plots was recorded at the end of each growing season. We used redundancy analysis to test for effects of initial density, irrigation, and year on the species and functional group composition of the communities. Results: Initial density had significant effects on species composition, and these effects remained significant over 3 years, even as total community percentage cover became more similar among treatments over time. Density effects did not depend on resource availability (irrigation level). Functional group identity or individual plant size did not predict which species would be good competitors, and a species' competitive ability did not predict its abundance in the field. Conclusions: Species interactions strongly affect community composition, and those effects carry over into subsequent years such that competition does not lead to convergence in community structure over time. However, the particular changes in composition observed were not predictable by some of the traits that have been found important in individual‐level experiments. We speculate that the outcome of competition in diverse communities will depend on multiple traits, in contrast to the outcome of individual‐level pairwise experiments. We also speculate that the shift in composition with density could mean that local variation in density may contribute to maintenance of diversity in this system.     

The role of mycorrhiza in plant populations and communities

Summary There is a large body of literature concerning the value of mycorrhiza to plant growth. Recent emphasis on the potential benefits of the fungi to natural ecosystems conservation and productive agriculture has focused attention on the roles and underlying mechanisms of the association. In parallel, recognition that isolates/ species of vesicular arbuscular mycorrhizal fungi have variable life-history traits has resulted in investigations focused on the symbiotic fungi rather than the host, a perspective which may reflect the fungal evolutionary strategies. This paper discusses progress in understanding interactions amongst hyphae, which in one form or another are a major component of mycorrhiza and each phase of the life-history.     

Factors affecting the invasion success of Senecio inaequidens and S. pterophorus in Mediterranean plant communities

Question: Plant invasions result from complex interactions between species traits, community characteristics and environmental variations. We examined the effect of these interactions on the invasion potential of two invasive Senecio species, S. inaequidens and S. pterophorus, across three Mediterranean plant communities in a natural park. Location: Catalonia, NE Spain. Methods: We carried out two series of experimental seedling transplantations, in the spring and fall of 2003, in grassland, shrubland and Quercus ilex forest. Competition with neighbouring plants and water availability were manipulated. We evaluated the survival, growth and reproduction with respect to each treatment combination. Results: Any habitat can be colonised if disturbance occurs. In the absence of disturbance, shrubland enhanced the survival of seedlings. Competition with resident vegetation dramatically reduced survival in grassland and forest when establishment occurred in the spring. However, establishment in the fall promoted invasion in grassland and shrubland, even in the undisturbed treatment. Grassland allowed the highest growth and reproductive performance of both species while forest was the most resistant habitat to invasion. S. inaequidens had a higher growth rate and a shorter pre‐reproductive period than S. pterophorus. S. pterophorus produced more biomass and was more dependent on water availability than S. inaequidens. Conclusions: In the light of our results, we recommend surveying open shrublands and grasslands after periods of rainfall. Special attention should be paid to S. pterophorus, which is currently spreading. A preliminary assessment of the invasive‐ness of this plant is given in this study.     

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.     

Competition and resource availability in an annual plant community dominated by an invasive species,Carrichtera annua (L. Aschers.), in South Australia

The last decade has seen spirited debates about how resource availability affect the intensity of competition. This paper examines the effect that a dominant introduced species, Carrichtera annua, has upon the winter annual community in the arid chenopod shrublands of South Australia. Manipulative field experiments were conducted to assess plant community response to changing below-ground resource levels and to the manipulation of the density of C. annua. Changes in the density of C. annua had little effect on the abundance of all other species in the guild. Nutrient addition produced an increase in the biomass of the most abundant native species, Crassula colorata. An analysis of the root distribution of the main species suggested that the areas of soil resource capture of C. annua and C. colorata are largely segregated. Our results suggest that intraspecific competition may be stronger than interspecific competition, controlling the species responses to increased resource availability. The results are consistent with a two-phase resource dynamics systems, with pulses of high resource availability triggering growth, followed by pulses of stress. Smaller plants were nutrient limited under natural field conditions, suggesting that stress experienced during long interpulse phases may override competitive effects after short pulse phases. The observed differences in root system structure will determine when plants of a different species are experiencing a pulse or an interpulse phase. We suggest that the limitations to plant recruitment and growth are the product of a complex interplay between the length and intensity of the pulse of resource availability, the duration and severity of the interpulse periods, and biological characters of the species.     

Phospholipase C: Diverse functions in plant biotic stress resistance and fungal pathogenicity

Phospholipase C (PLC) generates various second messenger molecules and mediates phospholipid hydrolysis. In recent years, the important roles of plant and fungal PLC in disease resistance and pathogenicity, respectively, have been determined. However, the roles of PLC in plants and fungi are unintegrated and relevant literature is disorganized. This makes it difficult for researchers to implement PLC-based strategies to improve disease resistance in plants. In this comprehensive review, we summarize the structure, classification, and phylogeny of the PLCs involved in plant biotic stress resistance and fungal pathogenicity. PLCs can be divided into two groups, nonspecific PLC (NPC) and phosphatidylinositol-specific PLC (PI-PLC), which present marked differences in phylogenetic evolution. The products of PLC genes in fungi play significant roles in physiological activity and pathogenesis, whereas those encoded by plant PLC genes mediate the immune response to fungi. This review provides a perspective for the future control of plant fungal diseases.     

Tree invasion of a montane meadow complex: temporal trends,spatial patterns,and biotic interactions

Questions: Do spatial and temporal patterns of encroachment of Pinus contorta and Abies grandis in a montane meadow suggest strong biotic controls on the invasion process? Location: Forest–meadow mosaic, 1350 m a.s.l., Cascade Range, Oregon, US. Methods: We combined spatial point pattern analysis, population age structures, and a time‐series of stem maps to quantify spatial and temporal patterns of conifer invasion over a 200‐yr period in three plots totaling 4 ha. Results: Trees established during two broad, but distinct periods (late 1800s, then at much greater density in the mid‐1900s). Recent invasion was not correlated with climatic variation. Abies grandis dominated both periods; P. contorta established at lower density, peaking before A. grandis. Spatially, older (≥90 yr) P. contorta were randomly distributed, but older A. grandis were strongly clumped (0.2‐20 m). Younger (<90 yr) stems were positively associated at small distances (both within and between species), but were spatially displaced from older A. grandis, suggesting a temporal shift from facilitation to competition. Establishment during the 1800s resulted in widely scattered P. contorta and clumps of A. grandis that placed most areas of meadow close to seed sources permitting more rapid invasion during the mid‐1900s. Rapid conversion to forest occurred via colonization of larger meadow openings – first by shade‐intolerant P. contorta, then by shade‐tolerant A. grandis– and by direct infilling of smaller openings by A. grandis. Conclusions: In combination, spatial and temporal patterns of establishment suggest an invasion process shaped by biotic interactions, with facilitation promoting expansion of trees into meadows and competition influencing subsequent forest development. Once invasion is initiated, tree species with different life histories and functional traits can interact synergistically to promote rapid conversion of meadow to forest under a broad range of climatic conditions.