Strength Of Competitive Interactions Research Articles

Intraspecific interference retards growth and development of cane toad tadpoles, but those effects disappear by the time of metamorphosis.

Competition among larval anurans can occur via interference as well as via a reduction in per-capita food supply. Previous research on intraspecific interference competition in cane toad (Rhinella marina) tadpoles found conflicting results, with one study detecting strong effects on tadpoles and another detecting no effects on metamorphs. A capacity to recover from competitive suppression by the time of metamorphosis might explain those contrasting impacts. In a laboratory experiment, we found that nine days of exposure to intraspecific interference competition strongly reduced tadpole growth and development, especially when the competing tadpoles were young (early-stage) individuals. Those competitive effects disappeared by the time of metamorphosis, with no significant effect of competition on metamorph body condition, size, larval period or survival. Temporal changes in the impact of competition were not related to tadpole density or to variation in water quality. The ability of larval cane toads to recover from intraspecific interference competition may enhance the invasive success of this species, because size at metamorphosis is a significant predictor of future fitness. Our study also demonstrates a cautionary tale: conclusions about the existence and strength of competitive interactions among anuran larvae may depend on which developmental stages are measured.

Metabolic phenotype mediates the outcome of competitive interactions in a response-surface field experiment.

Competition and metabolism should be linked. Intraspecific variation in metabolic rates and, hence, resource demands covary with competitive ability. The effects of metabolism on conspecific interactions, however, have mostly been studied under laboratory conditions. We used a trait‐specific response‐surface design to test for the effects of metabolism on pairwise interactions of the marine colonial invertebrate, Bugula neritina in the field. Specifically, we compared the performance (survival, growth, and reproduction) of focal individuals, both in the presence and absence of a neighbor colony, both of which had their metabolic phenotype characterized. Survival of focal colonies depended on the metabolic phenotype of the neighboring individual, and on the combination of both the focal and neighbor colony metabolic phenotypes that were present. Surprisingly, we found pervasive effects of neighbor metabolic phenotypes on focal colony growth and reproduction, although the sign and strength of these effects showed strong microenvironmental variability. Overall, we find that the metabolic phenotype changes the strength of competitive interactions, but these effects are highly contingent on local conditions. We suggest future studies explore how variation in metabolic rate affects organisms beyond the focal organism alone, particularly under field conditions.

Benthic invertebrate assemblages and leaf‐litter breakdown along the eucrenal–hypocrenal ecotone of a rheocrene spring in Central Italy: Are there spatial and seasonal differences?

AbstractTo better understand the pattern of variation in biocomplexity of springs and first‐order streams, it is essential to explore how the characteristics of spring communities may vary from the spring source (eucrenal) to the first reach of the originating springbrook (hypocrenal). For these reasons, we assessed spatial and temporal variation of crenic assemblages and leaf‐litter breakdown along a 45‐m reach of a spring–springbrook system in Central Italy. We found that physicochemical and hydrological parameters of Vera Spring did not vary significantly along the 45‐m reach investigated. In contrast, total richness of assemblages was significantly higher at 20 and 45 m from the spring source, while abundances were more variable and did not show significant differences between seasons and among sites. The composition of assemblages did not show significant temporal trends in all sites investigated, while spatial differences along the eucrenal–hypocrenal ecotone were more prominent. The functional organization of assemblages was relatively stable in both space and time, and no spatial differences were observed in the rate of leaf‐litter breakdown, though the dry mass loss of poplar leaves was slightly higher at the most downstream site. We conclude that immediately after groundwater emergence, spring communities are less stable and more variable while they tend to be better organized at short distance downstream. Small and gradual spatial changes in the strength of competitive interactions and predation pressures may have a fundamental role in structuring spring communities more than variation in abiotic parameters and/or differences in quantity and quality of food resources.

Structured environments foster competitor coexistence by manipulating interspecies interfaces.

Natural environments, like soils or the mammalian gut, frequently contain microbial consortia competing within a niche, wherein many species contain genetically encoded mechanisms of interspecies competition. Recent computational work suggests that physical structures in the environment can stabilize local competition between species that would otherwise be subject to competitive exclusion under isotropic conditions. Here we employ Lotka-Volterra models to show that interfacial competition localizes to physical structures, stabilizing competitive ecological networks of many species, even with significant differences in the strength of competitive interactions between species. Within a limited range of parameter space, we show that for stable communities the length-scale of physical structure inversely correlates with the width of the distribution of competitive fitness, such that physical environments with finer structure can sustain a broader spectrum of interspecific competition. These results highlight the potentially stabilizing effects of physical structure on microbial communities and lay groundwork for engineering structures that stabilize and/or select for diverse communities of ecological, medical, or industrial utility.

Soil biota increase the likelihood for coexistence among competing plant species.

Theory predicts that stable species coexistence will occur when population growth rates of competitively dominant species are suppressed when at high conspecific density. Although there is now compelling evidence that plant communities exhibit negative density dependence, the relative importance of the underlying processes leading to these patterns is rarely tested. We coupled reciprocal greenhouse and field experiments with community dynamics modeling to untangle the relative importance of soil biota from competition as stabilizing forces to coexistence. We found that (1) plant-soil biotic interactions compared to competitive interactions were stronger stabilizing forces, (2) only the strength of plant-soil biotic interactions was dependent on plant evolutionary history, and (3) the variation in the strength of plant-soil biotic interactions was correlated with relative abundance patterns in an opposite way than was the variation in the strength of competitive interactions. Collectively, our results demonstrate the fundamental role soil biota have in maintaining plant community diversity.

Species richness and vertical distribution of ferns and lycophytes along an elevational gradient in Los Tuxtlas, Veracruz, Mexico

We studied species richness and community composition as well as the vertical distribution of ferns and lycophytes in three forest types along an elevational gradient in the region of Los Tuxtlas, Mexico. We established 23 plots of 20m x 20m at 150–1650m, recording presence-absence of all species in eight vertical distribution zones (terrestrial and seven epiphytic zones). We found 84 species, including 79 ferns and five lycophytes. Species richness patterns differed markedly between different vertical zones, with the forest understory having a high diversity of epiphytes, many of which were specialized to this habitat. Overall species richness increased with elevation, but the richness of ground-living, terrestrial species showed the lowest elevational increase by only 50%, whereas epiphytic richness increased fourfold to tenfold. These differences presumably reflect the stronger influence of increasing environmental humidity on the water-limited epiphytic assemblages. We also found that the degree of habitat specialization of epiphytes decreased with elevation. This probably is due to decreasing tree height and increasing overall humidity, reducing the microclimatic differences between the forest interior and canopy. This elevational decrease in habitat specialization raises important questions as to, e.g., the strength of competitive interactions or the possibility that the evolutionary transition from terrestrial to epiphytic growth forms is more easily achieved at higher elevations.

Jointly modeling niche width and phylogenetic distance to explain species co‐occurrence

AbstractCompetitive exclusion is most likely when there are large differences in competitive ability and the strength of competitive interactions between species is high, but predicting competitive outcomes is not straightforward. Assuming a trade‐off between competitive ability and ecological generalism, we would predict larger competitive differences between species with different niche widths. Community phylogenetic theory predicts that competition will be stronger among more closely related species, assuming that phylogenetic distance reflects ecological similarity. We would therefore expect the probability of competitive exclusion to be highest among closely related species with different niche widths. Here, we assess how well differences in niche width and phylogenetic distance correlate with co‐occurrences among 34 species of Cyperaceae (sedges) in the eastern Canadian subarctic. The Cyperaceae is a species‐rich family, with many species sharing similar niches and environmental tolerances, making it a model clade for evaluating the importance of niche width differences and phylogenetic distances on co‐occurrence. Consistent with both hypotheses, we found that higher co‐occurrence scores correlated with species pairs that were distantly or only intermediately related and that had similar niche widths. Furthermore, we show that this correlation is stronger when considering only more recently diverged species pairs and that there is a triangular relationship between phylogenetic distance and species co‐occurrence, suggesting that distantly related species might have both strong and weak competitive interactions. Using co‐occurrence as a proxy for competitive outcomes, our results support both a negative correlation between phylogenetic distance and strength of competitive interactions, and a trade‐off between niche width and competitive ability.

Foraging modality and plasticity in foraging traits determine the strength of competitive interactions among carnivorous plants, spiders and toads.

Foraging modalities (e.g. passive, sit-and-wait, active) and traits are plastic in some species, but the extent to which this plasticity affects interspecific competition remains unclear. Using a long-term laboratory mesocosm experiment, we quantified competition strength and the plasticity of foraging traits in a guild of generalist predators of arthropods with a range of foraging modalities. Each mesocosm contained eight passively foraging pink sundews, and we employed an experimental design where treatments were the presence or absence of a sit-and-wait foraging spider and actively foraging toad crossed with five levels of prey abundance. We hypothesized that actively foraging toads would outcompete the other species at low prey abundance, but that spiders and sundews would exhibit plasticity in foraging traits to compensate for strong competition when prey were limited. Results generally supported our hypotheses. Toads had a greater effect on sundews at low prey abundances, and toad presence caused spiders to locate webs higher above the ground. Additionally, the closer large spider webs were to the ground, the greater the trichome densities produced by sundews. Also, spider webs were larger with than without toads and as sundew numbers increased, and these effects were more prominent as resources became limited. Finally, spiders negatively affected toad growth only at low prey abundance. These findings highlight the long-term importance of foraging modality and plasticity of foraging traits in determining the strength of competition within and across taxonomic kingdoms. Future research should assess whether plasticity in foraging traits helps to maintain coexistence within this guild and whether foraging modality can be used as a trait to reliably predict the strength of competitive interactions.

Determinants of growth of the flammable grass, Triodia scariosa: Consequences for fuel dynamics under climate change in the Mediterranean region of South Eastern Australia

AbstractEnvironmental conditions may influence the presence and strength of competitive interactions between different life forms, thereby shaping community composition and structure, and corresponding fuel dynamics. Woodland and shrubland communities of the Mediterranean climate region of South Eastern Australia contain a varied mixture of herbaceous and woody plants. The ratio of herbaceous to woody plants changes along gradients of temperature, moisture and soil fertility. This study aimed to experimentally examine the relative importance of, and interactions between environmental controls (moisture and soil fertility) on the balance of dominant herbaceous (Triodia scariosa) and woody plants (e.g. Acacia ligulata and Leptospermum coriaceum) and their ultimate effects on fuel and fire regimes. The results suggest that environmental determinants of the growth of T. scariosa are likely to be more important than interactions with shrubs in controlling the distribution of T. scariosa. The growth of T. scariosa was consistently higher under hot temperatures and on the less fertile yellow sands, which dominate the south of the region. The results suggest that there is strong potential for the distribution and abundance of T. scariosa to be altered in the future with changes in temperature associated with climate change. The distribution of soil types across the Mediterranean climate region of South Eastern Australia may be predisposed to favour the southerly expansion of T. scariosa‐dominated communities in the future under a warmer climate.

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.

Phylogenetic distance does not predict competition in green algal communities

Biologists have held the tenet that closely related species compete more strongly with each other than with distant relatives since 1859, when Darwin observed that close relatives seldom co‐occur in nature and suggested it was because they competitively exclude one another. The expectation that close relatives experience greater competition than distant relatives has become known as the “competition‐relatedness hypothesis (CRH).” The CRH is predicated on the assumption that closely related species are more likely to have similar resource requirements than distant relatives, and thus, compete more strongly for limited resources. While this assumption has been popular because it is intuitive, it has also been subject to relatively little experimentation. Over the past decade, a growing number of CRH studies have arrived at divergent conclusions showing that the strength of competitive interactions can increase, decrease, or be independent of evolutionary relatedness. Most of these studies have focused on measuring competition among species pairs as opposed to competition experienced by species when part of whole communities. We tested whether the CRH holds in communities where individual species experience interactions with a variety of other taxa, which we call the ‘resident community'. We performed a laboratory mesocosm study using communities of eight species of freshwater green algae whose evolutionary relationships were quantified using a recently developed multi‐gene molecular phylogeny of 59 North American green algae. We grew species alone and in various combinations in polyculture so that we could measure each species' sensitivity to competition (reduction in intrinsic growth rate when grown alone vs. with a resident community), relative yield, and competitive release (proportional change in biomass of a species when grown in a resident community missing one competitor vs. in a community with all possible competitors). While each of these metrics consistently revealed a prevalence of competitive interactions among the algal species, none were predicted by the relatedness of a species to a resident community. This suggests that the results of prior pairwise studies refuting the competition‐relatedness hypothesis for green algae can be extended to larger resident communities in which more complex ecological interactions possibly occur.

Neglected role of fungal community composition in explaining variation in wood decay rates.

Decomposition of wood is an important component of global carbon cycling. Most wood decomposition models are based on tree characteristics and environmental conditions; however, they do not include community dynamics of fungi that are the major wood decomposers. We examined the factors explaining variation in sapwood decay in oak tree stumps two and five years after cutting. Wood moisture content was significantly correlated with sapwood decay in younger stumps, whereas ITS-based composition and species richness of the fungal community were the best predictors for mass loss in the older stumps. Co-occurrence analysis showed that, in freshly cut trees and in younger stumps, fungal communities were nonrandomly structured, whereas fungal communities in old stumps could not be separated from a randomly assembled community. These results indicate that the most important factors explaining variation in wood decay rates can change over time and that the strength of competitive interactions between fungi in decaying tree stumps may level off with increased wood decay. Our field analysis further suggests that ascomycetes may have a prominent role in wood decay, but their wood-degrading abilities need to be further tested under controlled conditions. The next challenging step will be to integrate fungal community assembly processes in wood decay models to improve carbon sequestration estimates of forests.

Testing experimentally the effect of soil resource mobility on plant competition

Aims The volume of soil beyond a plant’s roots from which that plant is able to acquire a particular nutrient depends upon the mobility of that nutri ent in the soil. For this reason it has been hypothesized that the strength of competitive interactions between plants vary with soil nutrient mobility. We aimed to provide an experimental test of this hypothesis. Methods We devised two experimental systems to investigate specifically the effect of nutrient transport rates upon intraspecific competi tion. In the first, the exchange of rhizosphere water and dissolved nutrients between two connected pots, each containing one plant, was manipulated by alternately raising and lowering the pots. In the second experiment, the roots systems of two competing plants were separated by partitions of differing porosity, thereby varying

Disturbance, resource pulses and invasion: short-term shifts in competitive effects, not growth responses, favour exotic annuals

1. Increased resource availability and resource pulses often promote invasion by exotic invasive plants, but the relative importance of increased resource supply for invaders with different life histories is likely to vary. It is also unclear whether increased resources allow invaders to outgrow their native neighbours or alter the outcome of competition. Understanding mechanisms by which different invasive life histories respond to resource supply will help to target the most important species to control. 2. We found that aerially dropped wildfire retardant on intermountain grassland provided a very high pulse of N and P and caused dramatic increases in the abundance of annual exotic invaders already present in the system. Field experiments with retardant stimulated similar shifts to dominance by exotic annuals and decreases in native perennial grasses, but elicited no responses by native annual species. 3. In a greenhouse experiment, the growth response of both exotic and native species to retardant was not related to the intensity of mean competitive effects or responses. Concomitantly, gram per gram competitive effects differed substantially among species and treatments. N-caused shifts in the strength of competitive interactions were far better predictors than growth responses of the effects of retardant in the field. For example, Centaurea stoebe, an invasive perennial forb, grew disproportionally more than any other species in response to retardant when grown alone, but retardant significantly shifted competitive outcomes with this invader in favour of exotic annuals and the native bunch grass Pseudoroegneria spicata. 4. Synthesis and applications. Our results indicate that an extreme nutrient pulse did not create conditions whereby invaders outgrew native neighbours or other exotics, but instead altered the competitive playing field in ways that resulted in considerable changes in community composition. Given the minuscule scale of retardant drops on the landscape, our results do not suggest changing the way retardant is used to combat wildfire but do suggest that localized post-retardant weed control may be important so that retardant drops do not become point sources for exotic invasion. Furthermore, our results suggest that under high N conditions, invaders with annual life histories should be prioritized for control.

Testing Spatial Theories of Plant Coexistence: No Consistent Differences in Intra‐ and Interspecific Interaction Distances

Plants stand still and interact with their immediate neighbors. Theory has shown that the distances over which these interactions occur may have important consequences for population and community dynamics. In particular, if intraspecific competition occurs over longer distances than interspecific competition (heteromyopia), coexistence can be promoted. We examined how intraspecific and interspecific competition scales with neighbor distance in a target-neighbor greenhouse competition experiment. Individuals from co-occurring forbs from calcareous grasslands were grown in isolation and with single conspecific or heterospecific neighbors at distances of 5, 10, or 15 cm (Plantago lanceolata vs. Plantago media and Hieracium pilosella vs. Prunella grandiflora). Neighbor effects were strong and declined with distance. Interaction distances varied greatly within and between species, but we found no evidence for heteromyopia. Instead, neighbor identity effects were mostly explained by relative size differences between target and neighbor. We found a complex interaction between final neighbor size and identity such that neighbor identity may become important only as the neighbor becomes very large compared with the target individual. Our results suggest that species-specific size differences between neighboring individuals determine both the strength of competitive interactions and the distance over which these interactions occur.

Order of arrival affects competition in two reef fishes

Many communities experience repeated periods of colonization due to seasonally regenerating habitats or pulsed arrival of young-of-year. When an individual's persistence in a community depends upon the strength of competitive interactions, changes in the timing of arrival relative to the arrival of a competitor can modify competitive strength and, ultimately, establishment in the community. We investigated whether the strength of intracohort competitive interactions between recent settlers of the reef fishes Thalassoma hardwicke and T. quinquevittatum are dependent on the sequence and temporal separation of their arrival into communities. To achieve this, we manipulated the sequence and timing of arrival of each species onto experimental patch reefs by simulating settlement pulses and monitoring survival and aggressive interactions. Both species survived best in the absence of competitors, but when competitors were present, they did best when they arrived at the same time. Survival declined as each species entered the community progressively later than its competitor and as aggression by its competitor increased. Intraspecific effects of resident T. hardwicke were similar to interspecific effects. This study shows that the strength of competition depends not only on the identity of competitors, but also on the sequence and timing of their interactions, suggesting that when examining interaction strengths, it is important to identify temporal variability in the direction and magnitude of their effects. Furthermore, our findings provide empirical evidence for the importance of competitive lotteries in the maintenance of species diversity in demographically open marine systems.

Morphological convergence and coexistence in three sympatric North American species of Microtus (Rodentia: Arvicolinae)

AbstractAim Niche theory predicts that ecologically identical species cannot stably coexist in local communities. My aim was to investigate morphological diversity as a possible factor enabling the coexistence of a species‐rich Microtus (Rodentia: Arvicolinae) fauna in a hotspot of North American mammalian diversity, the Greater Yellowstone Ecosystem (GYE).Location The Greater Yellowstone Ecosystem, North America.Methods Using in‐hand morphological measurements of size and shape, I compared the morphologies of three North American vole species (Microtus spp.), in sympatry in the GYE and in allopatry across their ranges, in order to examine: (1) whether morphologies are fixed or plastic and (2) the degree of morphological character displacement or convergence in sympatric species.Results Support was found for plasticity of morphology for all three vole species: M. longicaudus, M. montanus and M. pennsylvanicus. However, Microtus individuals of all species from the GYE area of sympatry were more similar to each other than to allopatric individuals of the same species.Main conclusions Competition among these congeners is not manifested in morphological overdispersion. The response of these congeneric species to the same local ecological conditions is convergent. The relative strength of environmental conditions appears to be stronger than the strength of competitive interactions among the study species.

Uncertainty and the measurement of terrestrial biodiversity gradients

Uncertainty and the measurement of terrestrial biodiversity gradients

LIMITING RESOURCES AND THE REGULATION OF DIVERSITY IN PHYTOPLANKTON COMMUNITIES

Species diversity is a key concept in ecology, yet the mechanisms regulating diversity in most systems are not completely understood. To address this issue, we analyzed the relationship between phytoplankton diversity and limiting resources (N, P, Si, and light) over two summers in three lakes in the Yellowstone (Wyoming, USA) region. Diversity was highly variable along temporal and spatial axes within lakes. We discovered a strong positive correlation between diversity and the number of resources at physiologically limiting levels. Consistent with resource-competition theory, we found the highest diversity to occur when many resources were limiting. Conversely, the lowest diversity occurred when few resources were measured at limiting levels. Sensitivity analyses demonstrated that threshold levels of resources (below which growth is limited and above which there is saturation for the resource) appear to exist in the natural environment, and that diversity is regulated in part by absolute levels of resources available to phytoplankton in aquatic systems. Threshold levels are generally in agreement with those determined to be limiting in prior physiological experiments. Consistent with previous observations, among-lake comparisons yielded a significant negative relationship between species diversity and system productivity. This result tentatively supports the hypothesis that the proximity of individual phytoplankton plays a role in the strength of competitive interactions. Our results generally suggest that, even in dynamic environments, where equilibrium conditions are rare, resource competition among phytoplankton is a mechanism by which communities are continually structured.

Limiting Resources and the Regulation of Diversity in Phytoplankton Communities

Species diversity is a key concept in ecology, yet the mechanisms regulating diversity in most systems are not completely understood. To address this issue, we analyzed the relationship between phytoplankton diversity and limiting resources (N, P, Si, and light) over two summers in three lakes in the Yellowstone (Wyoming, USA) region. Diversity was highly variable along temporal and spatial axes within lakes. We discovered a strong positive correlation between diversity and the number of resources at physiologically limiting levels. Consistent with resource-competition theory, we found the highest diversity to occur when many resources were limiting. Conversely, the lowest diversity occurred when few resources were measured at limiting levels. Sensitivity analyses demonstrated that threshold levels of resources (below which growth is limited and above which there is saturation for the resource) appear to exist in the natural environment, and that diversity is regulated in part by absolute levels of resources available to phytoplankton in aquatic systems. Threshold levels are generally in agreement with those determined to be limiting in prior physiological experiments. Consistent with previous observations, among-lake comparisons yielded a significant negative relationship between species diversity and system productivity. This result tentatively supports the hypothesis that the proximity of individual phytoplankton plays a role in the strength of competitive interactions. Our results generally suggest that, even in dynamic environments, where equilibrium conditions are rare, resource competition among phytoplankton is a mechanism by which communities are continually structured.