Plant competitive ability and the transitivity of competitive hierarchies change with plant age

Plant competitive effect and response ability are known to change with plant age, however it remains unclear how competitive hierarchies among plant species change as plants age and transition between life stages. We examined the competitive interactions among seven species in all pairwise combinations in a greenhouse experiment. Competitive effect and response were measured as the relative yield (RY) for each target-neighbor species combination for both seedling and adult plants. Competitive hierarchies were constructed based on competitive effect and response scores, and we examined the degree of transitivity in the seedling and adult competitive hierarchies. The competitive effect hierarchy did not vary substantially with plant age, while the competitive response hierarchy was highly variable between juvenile and adult plants. Competitive effect and response ability were positively correlated at both plant stages. The seedling relative yield matrix was predominantly transitive, while there were far fewer transitive competitive relationships among the adult plants. The breakdown of the clear competitive hierarchy among seedlings as plants aged may explain why competition does not appear to be an active mechanism structuring some late-successional plant communities. In early-successional communities, interactions among seedlings with a clear competitive hierarchy may establish competitive ability—abundance relationships that persist as a legacy effect even though the breakdown of the competitive hierarchy among adult plants removes competition as an active mechanism structuring some late-successional plant communities.     

Determining the outcome of field-based competition between two Rhizopogon species using real-time PCR

Interest in the ecology of ectomycorrhizal (ECM) fungi has increased considerably, but little is known about interspecific interactions among ECM species. We examined competitive interactions between Rhizopogon occidentalis and R. salebrosus at Point Reyes National Seashore, California, USA. At three field sites, species abundances were compared in single- and two-species treatments on Pinus muricata seedlings inoculated with spores. Competition for root tips was assessed using real-time polymerase chain reaction (PCR) of internal transcribed spacer rDNA. In general, we found strong competitive exclusion of R. salebrosus by R. occidentalis, with >or= 75% of the seedlings in the two-species treatment colonized exclusively by R. occidentalis after 5 and 10 months. However, on the seedlings that were co-colonized, we observed no significant difference in the abundances of R. salebrosus and R. occidentalis, suggesting that once R. salebrosus was established, it was no longer competitively inferior. There were no significant differences in survival, growth, or percentage leaf nitrogen of seedlings colonized with either Rhizopogon species, but both growth and percentage leaf nitrogen were significantly higher for ECM than non-ECM seedlings. We also observed strong positive correlations between actual ECM root tip weight and that inferred from real-time PCR for both species, indicating that this method provided an accurate assessment of root tip occupation and hence ECM competitive dynamics. In conjunction with a previous experiment, our results indicate that competition between these two Rhizopogon species occurs similarly in both field and laboratory settings and that when colonizing from spore, timing largely determines the outcome of initial competitive interactions.     

Competitive relationships in natural and artificial algal communities

Modern data on competitive relationships and their role in the succession of natural and artificial algal communities are reviewed. The mechanisms of macroalgae competition and the factors that affect the competitive outcomes are considered. The conception of competitive interactions between seaweeds in the field and culture is suggested. (1) Competitive relationships are possible only between seaweeds which live together and are able to exchange signals. (2) Success in the competition for light is the basis for wins in the competition for space. (3) The competition for nutrients never results directly in the exclusion of the competitor from the community. It inhibits the competitor and allows the winner to overgrow, shade, act allelopathically, and to displace the inferior competitor in the community. (4) People, creating an artificial monodominant community, either increase the competitive potential of cultivated species by selection of growth conditions or exclude the competitors.     

Competition and succession among coral endosymbionts

Host species often support a genetically diverse guild of symbionts, the identity and performance of which can determine holobiont fitness under particular environmental conditions. These symbiont communities are structured by a complex set of potential interactions, both positive and negative, between the host and symbionts and among symbionts. In reef‐building corals, stable associations with specific symbiont species are common, and we hypothesize that this is partly due to ecological mechanisms, such as succession and competition, which drive patterns of symbiont winnowing in the initial colonization of new generations of coral recruits. We tested this hypothesis using the experimental framework of the de Wit replacement series and found that competitive interactions occurred among symbionts which were characterized by unique ecological strategies. Aposymbiotic octocoral recruits within high‐ and low‐light environments were inoculated with one of three Symbiodiniaceae species as monocultures or with cross‐paired mixtures, and we tracked symbiont uptake using quantitative genetic assays. Priority effects, in which early colonizers excluded competitive dominants, were evidenced under low light, but these early opportunistic species were later succeeded by competitive dominants. Under high light, a more consistent competitive hierarchy was established in which competitive dominants outgrew and limited the abundance of others. These findings provide insight into mechanisms of microbial community organization and symbiosis breakdown and recovery. Furthermore, transitions in competitive outcomes across spatial and temporal environmental variation may improve lifetime host fitness.     

Founder control and coexistence in a simple model of asymmetric competition for light

Size asymmetry in plant light acquisition complicates predictions of competitive outcomes in light-limited communities. We present a mathematically tractable model of asymmetric competition for light and discuss its implications for predicting outcomes of competition during establishment in two-, three-, and many-species communities. In contrast to the resource-reduction model of symmetric competition for a single resource, the model we present predicts that outcomes of asymmetric competition for light will sometimes depend on the timing of establishment and the consequent hierarchy among species in canopy position. Competitive outcomes in the model depend on the minimum light requirements (L(c)) and self-shading of species lower in the canopy compared to the light available (L(out)(*)) beneath species higher in the canopy. Succession progresses towards species with decreasing values for L(c), but arrested successions occur when initial dominants have relatively high values for L(c) but low values for L(out)(*), leading to founder control. A theoretically limitless number of species may coexist in competition for light when dominance is founder controlled. These model predictions have implications for an array of applied ecological questions, including methods to control invasive species in light-limited restored ecosystems.     

Can community composition be predicted from pairwise species interactions?

Plant communities are often structured by interactions among species, such as competition or facilitation. If competition is an important factor that controls the presence and absence of species within intact communities, then a competitive hierarchy, a ranked order from competitive dominant to competitive subordinate, should predict the composition of intact communities. We tested whether a competitive hierarchy derived from pairwise comparisons accurately predicts species abundances within a constructed polyculture community consisting of seven species common to old-field plant communities. We first conducted a pot experiment in field conditions wherein we grew the species in all possible combinations, then created a competitive hierarchy derived from both competitive effect and competitive response for each species. Concurrently, at the same site in native field soil, we constructed polycultures consisting of the same seven species and calculated an abundance hierarchy based on foliar cover, biomass, and an index of species performance. The competitive hierarchy was not concordant with the abundance hierarchy, indicating that simple pairwise comparisons may not account for other factors that influence the abundance of species within relatively complex communities.     

Primary succession of Bistorta vivipara (L.) Delabre (Polygonaceae) root‐associated fungi mirrors plant succession in two glacial chronosequences

Glacier chronosequences are important sites for primary succession studies and have yielded well‐defined primary succession models for plants that identify environmental resistance as an important determinant of the successional trajectory. Whether plant‐associated fungal communities follow those same successional trajectories and also respond to environmental resistance is an open question. In this study, 454 amplicon pyrosequencing was used to compare the root‐associated fungal communities of the ectomycorrhizal (ECM) herb Bistorta vivipara along two primary succession gradients with different environmental resistance (alpine versus arctic) and different successional trajectories in the vascular plant communities (directional replacement versus directional non‐replacement). At both sites, the root‐associated fungal communities were dominated by ECM basidiomycetes and community composition shifted with increasing time since deglaciation. However, the fungal community's successional trajectory mirrored the pattern observed in the surrounding plant community at both sites: the alpine site displayed a directional‐replacement successional trajectory, and the arctic site displayed a directional‐non‐replacement successional trajectory. This suggests that, like in plant communities, environmental resistance is key in determining succession patterns in root‐associated fungi. The need for further replicated study, including in other host species, is emphasized.     

Spatial analysis of ectomycorrhizal fungi reveals that root tip communities are structured by competitive interactions

Microbial ecology has made large advances over the last decade, mostly because of improvements in molecular analysis techniques that have enabled the detection and identification of progressively larger numbers of microbial species. However, determining the ecological patterns and processes taking place in communities of microbes remains a significant challenge. Are communities randomly assembled through dispersal and priority effects, or do species interact with each other leading to positive and negative associations? For mycorrhizal fungi, evidence is accumulating that stochastic and competitive interactions between species may both have a role in shaping community structure. Could the methodological approach, which is often incidence based, impact the outcomes detected? Here, we applied an incidence‐based Terminal Restriction Fragment Length Polymorphism (T‐RFLP) database approach to examine species diversity and ecological interactions within a community of ectomycorrhizal (ECM) fungi. Co‐occurrence analysis revealed that the ECM community colonizing root tips was strongly structured by competitive interactions, or ecological processes generating a similar spatial pattern, rather than neutral processes. Analysis of β‐diversity indicated that community structure was significantly more similar (spatially autocorrelated) at distances equal to or <3.41 m. The eight most frequently encountered species in the root tip community of ECM fungi displayed significant competitive interactions with at least one other species, showing that the incidence‐based approach was capable of detecting this sort of ecological information.     

Priority effects determine the outcome of ectomycorrhizal competition between two Rhizopogon species colonizing Pinus muricata seedlings

Competition is often considered a fundamental process influencing assemblage structure, yet little is known about competition among ectomycorrhizal (EM) fungi. Here, we examine competitive interactions between Rhizopogon occidentalis and Rhizopogon salebrosus in a 6-month microcosm study. Pinus muricata seedlings were grown in three EM treatments: R. occidentalis, R. salebrosus, and R. occidentalis and R. salebrosus. At 2, 4, and 6 months, five seedlings per treatment were harvested and the EM root tip biomass of each species was determined. Root tips in the two-species treatment were identified using molecular techniques. R. occidentalis had similar EM root tip biomass when grown alone or in the presence of R. salebrosus. By contrast, R. salebrosus had significantly lower EM root tip biomass when grown with R. occidentalis than when grown alone, indicating it was a competitive inferior under the conditions tested. Competition was driven by differences in timing of colonization resulting in a strong priority effect for R. occidentalis. Our results, together with two earlier studies, indicate competition may play a more important role in EM interactions than previously recognized.     

Extreme weather events and plant–plant interactions: shifts between competition and facilitation among grassland species in the face of drought and heavy rainfall

Biotic interactions play an important role in ecosystem function and structure in the face of global climate change. We tested how plant–plant interactions, namely competition and facilitation among grassland species, respond to extreme drought and heavy rainfall events. We also examined how the functional composition (grasses, forbs, legumes) of grassland communities influenced the competition intensity for grass species when facing extreme events. We exposed experimental grassland communities of different functional compositions to either an extreme single drought event or to a prolonged heavy rainfall event. Relative neighbour effect, relative crowding and interaction strength were calculated for five widespread European grassland species to quantify competition. Single climatic extremes caused species specific shifts in plant–plant interactions from facilitation to competition or vice versa but the nature of the shifts varied depending on the community composition. Facilitation by neighbouring plants was observed for Arrhenatherum elatius when subjected to drought. Contrarily, the facilitative effect of neighbours on Lotus corniculatus was transformed into competition. Heavy rainfall increased the competitive effect of neighbours on Holcus lanatus and Lotus corniculatus in communities composed of three functional groups. Competitive pressure on Geranium pratense and Plantago lanceolata was not affected by extreme weather events. Neither heavy rainfall nor extreme drought altered the overall productivity of the grassland communities. The complementary responses in competition intensity experienced by grassland species under drought suggest biotic interactions as one stabilizing mechanism for overall community performance. Understanding competitive dynamics under fluctuating resources is important for assessing plant community shifts and degree of stability of ecosystem functions.     

Phylogenetic and functional structures of succession in plant communities on mounds of Marmota himalayana in alpine regions on the northeast edge of the Qinghai–Tibet Plateau

Few studies have examined the succession of plant communities in the alpine zone. Studying the succession of plant communities is helpful to understand how species diversity is formed and maintained. In this study, we used species inventories, a molecular phylogeny, and trait data to detect patterns of phylogenetic and functional community structure in successional plant communities growing on the mounds of Himalayan marmots (Marmota himalayana) on the southeast edge of the Qinghai-Tibet Plateau. We found that phylogenetic and functional diversities of plant communities on marmot mounds tended to cluster during the early to medium stages of succession, then trended toward overdispersion from medium to late stages. Alpine species in early and late stages of succession were phylogenetically and functionally overdispersed, suggesting that such communities were assembled mainly through species interactions, especially competition. At the medium and late stages of succession, alpine communities growing on marmot mounds were phylogenetically and functionally clustered, implying that the communities were primarily structured by environmental filtering. During the medium and late stages of succession the phylogenetic and functional structures of plant communities on marmot mounds differed significantly from those on neighboring sites. Our results indicate that environmental filtering and species interactions can change plant community composition at different successional stages. Assembly of plant communities on marmot mounds was promoted by a combination of traits that may provide advantages for survival and adaptation during periods of environmental change.     

Species interactions and random dispersal rather than habitat filtering drive community assembly during early plant succession

Theory on plant succession predicts a temporal increase in the complexity of spatial community structure and of competitive interactions: initially random occurrences of early colonising species shift towards spatially and competitively structured plant associations in later successional stages. Here we use long‐term data on early plant succession in a German post mining area to disentangle the importance of random colonisation, habitat filtering, and competition on the temporal and spatial development of plant community structure. We used species co‐occurrence analysis and a recently developed method for assessing competitive strength and hierarchies (transitive versus intransitive competitive orders) in multispecies communities. We found that species turnover decreased through time within interaction neighbourhoods, but increased through time outside interaction neighbourhoods. Successional change did not lead to modular community structure. After accounting for species richness effects, the strength of competitive interactions and the proportion of transitive competitive hierarchies increased through time. Although effects of habitat filtering were weak, random colonization and subsequent competitive interactions had strong effects on community structure. Because competitive strength and transitivity were poorly correlated with soil characteristics, there was little evidence for context dependent competitive strength associated with intransitive competitive hierarchies.     

Contrasting Effects of Elevated Temperature and Invertebrate Grazing Regulate Multispecies Interactions between Decomposer Fungi

Predicting the influence of biotic and abiotic factors on species interactions and ecosystem processes is among the primary aims of community ecologists. The composition of saprotrophic fungal communities is a consequence of competitive mycelial interactions, and a major determinant of woodland decomposition and nutrient cycling rates. Elevation of atmospheric temperature is predicted to drive changes in fungal community development. Top-down regulation of mycelial growth is an important determinant of, and moderator of temperature-driven changes to, two-species interaction outcomes. This study explores the interactive effects of a 4 °C temperature increase and soil invertebrate (collembola or woodlice) grazing on multispecies interactions between cord-forming basidiomycete fungi emerging from colonised beech (Fagus sylvatica) wood blocks. The fungal dominance hierarchy at ambient temperature (16 °C; Phanerochaete velutina > Resinicium bicolor > Hypholoma fasciculare) was altered by elevated temperature (20 °C; R. bicolor > P. velutina > H. fasciculare) in ungrazed systems. Warming promoted the competitive ability of the fungal species (R. bicolor) that was preferentially grazed by all invertebrate species. As a consequence, grazing prevented the effect of temperature on fungal community development and maintained a multispecies assemblage. Decomposition of fungal-colonised wood was stimulated by warming, with implications for increased CO₂ efflux from woodland soil. Analogous to aboveground plant communities, increasing complexity of biotic and abiotic interactions appears to be important in buffering climate change effects on soil decomposers.     

When competition does not matter: grassland diversity and community composition

We examined whether the intense root competition in a rough fescue grassland plant community in central Alberta, Canada, was important in structuring plant species diversity or community composition. We measured competition intensity across gradients of species richness, evenness, and community composition, using pairs of naturally occurring plants of 12 species. One plant in each pair was isolated from neighbors to measure competition; community structure and environmental conditions were also measured at each pair. We used structural equation modeling to examine how competition influenced community structure. Competition intensity was unrelated to species richness and community composition, but increased competition intensity was associated with a slight decline in evenness. Size-symmetric root competition was probably unimportant in structuring this plant community because there are no feedback mechanisms through which size-symmetric competition can magnify small initial differences and eventually lead to competitive exclusion. In plant communities with little shoot competition, competition and community structure should be unlinked regardless of competition intensity. In more productive systems, we propose that interactions between root and shoot competition may indirectly structure communities by altering the overall asymmetry of competition.     

Interactions between functionally diverse fungal mutualists inconsistently affect plant performance and competition

Plants form mutualistic relationship with a variety of belowground fungal species. Such a mutualistic relationship can enhance plant growth and resistance to pathogens. Yet, we know little about how interactions between functionally diverse groups of fungal mutualists affect plant performance and competition. We experimentally determined the effects of interaction between two functional groups of belowground fungi that form mutualistic relationship with plants, arbuscular mycorrhizal (AM) fungi and Trichoderma, on interspecific competition between pairs of closely related plant species from four different genera. We hypothesized that the combination of two functionally diverse belowground fungal species would allow plants and fungi to partition their symbiotic relationships and relax plant–plant competition. Our results show that: 1) the AM fungal species consistently outcompeted the Trichoderma species independent of plant combinations; 2) the fungal species generally had limited effects on competitive interactions between plants; 3) however, the combination of fungal species relaxed interspecific competition in one of the four instances of plant–plant competition, despite the general competitive superiority of AM fungi over Trichoderma. We highlight that the competitive outcome between functionally diverse fungal species may show high consistency across a broad range of host plants and their combinations. However, despite this consistent competitive hierarchy, the consequences of their interaction for plant performance and competition can strongly vary among plant communities.     

Do biotic interactions shape both sides of the humped‐back model of species richness in plant communities?

A humped-back relationship between species richness and community biomass has frequently been observed in plant communities, at both local and regional scales, although often improperly called a productivity-diversity relationship. Explanations for this relationship have emphasized the role of competitive exclusion, probably because at the time when the relationship was first examined, competition was considered to be the significant biotic filter structuring plant communities. However, over the last 15 years there has been a renewed interest in facilitation and this research has shown a clear link between the role of facilitation in structuring communities and both community biomass and the severity of the environment. Although facilitation may enlarge the realized niche of species and increase community richness in stressful environments, there has only been one previous attempt to revisit the humped-back model of species richness and to include facilitative processes. However, to date, no model has explored whether biotic interactions can potentially shape both sides of the humped-back model for species richness commonly detected in plant communities. Here, we propose a revision of Grime's original model that incorporates a new understanding of the role of facilitative interactions in plant communities. In this revised model, facilitation promotes diversity at medium to high environmental severity levels, by expanding the realized niche of stress-intolerant competitive species into harsh physical conditions. However, when environmental conditions become extremely severe the positive effects of the benefactors wane (as supported by recent research on facilitative interactions in extremely severe environments) and diversity is reduced. Conversely, with decreasing stress along the biomass gradient, facilitation decreases because stress-intolerant species become able to exist away from the canopy of the stress-tolerant species (as proposed by facilitation theory). At the same time competition increases for stress-tolerant species, reducing diversity in the most benign conditions (as proposed by models of competition theory). In this way our inclusion of facilitation into the classic model of plant species diversity and community biomass generates a more powerful and richer predictive framework for understanding the role of plant interactions in changing diversity. We then use our revised model to explain both the observed discrepancies between natural patterns of species richness and community biomass and the results of experimental studies of the impact of biodiversity on the productivity of herbaceous communities. It is clear that explicit consideration of concurrent changes in stress-tolerant and competitive species enhances our capacity to explain and interpret patterns in plant community diversity with respect to environmental severity.     

Species coexistence, intransitivity, and topological variation in competitive tournaments

Competitive intransitivity occurs when species’ competitive abilities cannot be listed in a strict hierarchy, but rather form competitive loops, as in the game ‘Rock-Paper-Scissors’. Indices are useful for summarizing intransitivity in communities; however, as with most indices, a great deal of information is compressed into single number. So while recent ecological theory, experiments, and natural history observations demonstrate that competitive intransitivity can promote species coexistence, the consequence of variation in the ‘topology’ of competitive interactions that is not accounted for by intransitivity indices is much less well understood. We use a continuous analytical model and two complementary discrete lattice models (one spatially explicit, the other aspatial) to demonstrate that such variation does indeed greatly affect species coexistence. Specifically, we show that although intransitivity indices are good at capturing broad patterns of coexistence, communities with different levels of intransitivity can have equal coexistence, and communities with equal intransitivity can have different coexistence, due to underlying variation in competitive network topology.     

Experimental evidence for root competition effects on community evenness in one of two phytometer species

Aims Plant–plant interactions, being positive or negative, are recognized to be key factors in structuring plant communities. However, it is thought that root competition may be less important than shoot competition due to greater size symmetry belowground. Because direct experimental tests on the importance of root competition are scarce, we aim at elucidating whether root competition may have direct or indirect effects on community structure. Indirect effects may occur by altering the overall size asymmetry of competition through root–shoot competitive interactions.     

The effects of species pool,dispersal and competition on the diversity–productivity relationship

Aim The diversity–productivity relationship is a controversial issue in ecology. Diversity is sometimes seen to increase with productivity but a unimodal relationship has often been reported. Competitive exclusion was cited initially to account for the decrease of diversity at high productivity. Subsequently, the roles of evolutionary history (species pool size) and dispersal rate have been acknowledged. We explore how the effects of species pool, dispersal and competition combine to produce different diversity–productivity relationships. Methods We use a series of simulations with a spatially explicit, individual‐based model. Following empirical expectations, we used four scenarios to characterize species pool size along the productivity gradient (uniformly low and high, linear increase and unimodal). Similarly, the dispersal rate varied along the productivity gradient (uniformly low and high, and unimodal). We considered both neutral communities and communities with competitive exclusion. Results and main conclusions Our model predicts that competitive interactions will result in unimodal diversity–productivity relationships. The model often predicts unimodal patterns in neutral communities as well, although the decline in richness at high productivity is less than in competing communities. A positive diversity–productivity relationship is simulated for neutral communities when the species pool size increases with productivity and the dispersal rate is high. This scenario is probably more widespread in nature than the others since positive diversity–productivity relationships have been observed more frequently than previously expected, especially in the tropics and for woody species. Our simulated effects of species pool, dispersal and competition on diversity patterns can be linked to empirical observations to uncover mechanisms behind the diversity–productivity relationship.     

Permanent plots as tools for plant community ecology

Abstract. Permanent plots provide information on spatio-temporal patterns in plant communities, which could be analyzed to yield correlated changes in pairs of species and information on species replacements in space. The potential use of permanent plots to study underlying processes of interspecific interactions (namely competition) is discussed. Two examples (one simulated, using a cellular automaton model, and one from a removal experiment in grasslands) are used to demonstrate that this use is not straightforward, since (1) radically different underlying mechanisms (symmetric founder control with little competitive replacement vs. asymmetric dominance control strongly structured by competition) may produce very similar spatio-temporal patterns; and (2) topology of species replacements in space may be very different from the competitive hierarchy as determined by the removal experiment. Permanent plot data can be used to study interspecific interactions in the following ways: (1) if there is additional information to decide which of the potential underlying models is acceptable, permanent plot data may enable further articulation of the model, such as estimation of the model parameters; this approach is similar to that used in many other plant ecology fields (fine scale patterns, chronosequences, plant size distribution and dynamics); (2) data from permanent plots constrain the number of feasible underlying models; (3) permanent plot data provide verification of predictions of experimental studies; in this respect, they are much superior to non-repeated studies of pattern.