Competition-colonization Trade-off Research Articles

Solitary foundation or colony fission in ants: an intraspecific study shows that worker presence and number increase colony foundation success.

Dispersal and establishment strategies are highly variable. Each strategy is associated with specific costs and benefits, and understanding which factors favour or disfavour a strategy is a key issue in ecology and evolution. Ants exhibit several strategies of establishment, i.e. of colony foundation. Some species rely on winged queens that found new colonies alone when others found with accompanying workers (colony fission). The benefits conferred by these workers have been little studied and quantified, because comparing the costs and benefits of solitary foundation vs. colony fission is difficult when comparing different species. We investigated this using the ant Myrmecina graminicola, one of the few species that use both strategies. Young mated queens were allowed to found new colonies in the laboratory, with either zero (solitarily), two or four workers (colony fission). The presence of workers increased both survival and growth of the foundations over the first year, with more workers yielding higher growth. Few workers (as little as two workers) were sufficient to provide benefits, suggesting that in M. graminicola the strategy of colony fission may not dramatically decrease the number of new colonies produced compared to solitary foundation. Because queens performing solitary foundation or colony fission differ in dispersal (by flight vs. on foot), our results support the hypothesis that these two strategies of foundation coexist along a competition-colonization trade-off, where solitary foundation offers a colonization advantage, while colony fission has a competitive advantage.

Coexistence of many species under a random competition–colonization trade-off

The competition-colonization (CC) trade-off is a well-studied coexistence mechanism for metacommunities. In this setting, it is believed that the coexistence of all species requires their traits to satisfy restrictive conditions limiting their similarity. To investigate whether diverse metacommunities can assemble in a CC trade-off model, we study their assembly from a probabilistic perspective. From a pool of species with parameters (corresponding to traits) sampled at random, we compute the probability that any number of species coexist and characterize the set of species that emerges through assembly. Remarkably, almost exactly half of the species in a large pool typically coexist, with no saturation as the size of the pool grows, and with little dependence on the underlying distribution of traits. Through a mix of analytical results and simulations, we show that this unlimited niche packing emerges as assembly actively moves communities toward overdispersed configurations in niche space. Our findings also apply to a realistic assembly scenario where species invade one at a time from a fixed regional pool. When diversity arises de novo in the metacommunity, richness still grows without bound, but more slowly. Together, our results suggest that the CC trade-off can support the robust emergence of diverse communities, even when coexistence of the full species pool is exceedingly unlikely.

Functional diversity and spatial association analyses at different spatial scales reveal no changes in community assembly processes along an aridity gradient in the Atacama Desert

The structuring of plant assemblages along environmental gradients is typically explained by shifts from competition (limiting similarity) to environmental filtering as the environment becomes more stressful. However, facilitation, weaker-competitor exclusion, environmental heterogeneity, and the colonization-competition tradeoff can also structure plant assemblages along gradients. These assembly processes act on different plant traits and organs, and their prevalence varies with respect to the spatial scale. Using patterns of functional diversity, coupled with patterns of species association at two spatial scales, here we discern the assembly processes that structure shrub communities in four localities along an aridity gradient of the Atacama Desert. At each site, we calculated functional dispersion indexes for above- and below-ground traits, and patterns of species association at a patch and neighborhood scale. Our results revealed that at the patch scale in intermediate levels of aridity, the dominant assembly process was within-site environmental heterogeneity. At the neighborhood scale, communities are assembled mainly through random processes. Nonetheless, in some communities, the dominant assembly process was competition via limiting similarity or exclusion of the weaker competitor, and these did not change along the gradient. Together, these results reveal that environmental heterogeneity and competition are the main drivers of plant community assembly in a hyper-arid environment.

Complex patch geometry promotes species coexistence through a reverse competition-colonization trade-off.

Explaining the maintenance of diverse species assemblages is a central goal of ecology and conservation. Recent coexistence mechanisms highlight the role of dispersal as a source of the differences that allow similar species to coexist. Here, we propose a new mechanism for species coexistence that is based on dispersal differences, and on the geometry of the habitat patch. In a finite habitat patch with complex boundaries, species with different dispersal abilities will arrange themselves in stable, concentric patterns of dominance. Species with superior competitive and dispersal abilities will dominate the interior of the patch, with inferior species at the periphery. We demonstrate and explain the mechanism on a simple one-dimensional domain, and then on two-dimensional habitat patches with realistic geometries. Finally, we use metrics from landscape ecology to demonstrate that habitat patches with more complex geometries can more easily support coexistence. The factors that underpin this new coexistence mechanism-different dispersal abilities and habitat patches with complex geometries-are common to many marine and terrestrial ecosystems, and it is therefore possible that the mechanism is a common factor supporting diverse species assemblages.

Complex nonmonotonic responses of biodiversity to habitat destruction.

It has typically been assumed that habitat destruction, characterized by habitat loss and fragmentation, has consistently negative effects on biodiversity. While numerous empirical studies have shown the detrimental effects of habitat loss, debate continues as to whether habitat fragmentation has universally negative effects. To explore the effects of habitat fragmentation, we developed a simple model for site-occupancy dynamics in fragmented landscapes. With the model, we demonstrate that a competition-colonization trade-off can result in nonlinear oscillatory responses in biodiversity to both habitat loss and fragmentation. However, the overall pattern of habitat loss reducing species richness is still established, in line with empirical observations. Interestingly, the existence of localized oscillations in biodiversity can explain the mixed responses of species richness to habitat fragmentation per se observed in nature, thereby reconciling the debate on the fragmentation-diversity relationship. Therefore, this study offers a parsimonious mechanistic explanation for empirically observed biodiversity patterns in response to habitat destruction.

Assembly in metacommunities driven by a competition-colonization tradeoff: more species in, more species out

Assembly in metacommunities driven by a competition-colonization tradeoff: more species in, more species out

Coevolution of species colonisation rates controls food-chain length in spatially structured food webs.

How the complexity of food webs depends on environmental variables is a long-standing ecological question. It is unclear though how food-chain length should vary with adaptive evolution of the constitutive species. Here we model the evolution of species colonisation rates and its consequences on occupancies and food-chain length in metacommunities. When colonisation rates can evolve, longer food-chains can persist. Extinction, perturbation and habitat loss all affect evolutionarily stable colonisation rates, but the strength of the competition-colonisation trade-off has a major role: weaker trade-offs yield longer chains. Although such eco-evo dynamics partly alleviates the spatial constraint on food-chain length, it is no magic bullet: the highest, most vulnerable, trophic levels are also those that least benefit from evolution. We provide qualitative predictions regarding how trait evolution affects the response of communities to disturbance and habitat loss. This highlights the importance of eco-evolutionary dynamics at metacommunity level in determining food-chain length.

Local adaptation, phenotypic plasticity, and species coexistence

Understanding the mechanisms of species coexistence has always been a fundamental topic in ecology. Classical theory predicts that interspecific competition may select for traits that stabilize niche differences, although recent work shows that this is not strictly necessary. Here, we ask whether adaptive phenotypic plasticity could allow species coexistence (i.e., some stability at an equilibrium point) without ecological differentiation in habitat use. We used individual-based stochastic simulations defining a landscape composed of spatially uncorrelated or autocorrelated environmental patches, where two species with the same competitive strategies, not able to coexist without some form of phenotypic plasticity, expanded their ranges in the absence of a competition—colonization trade-off (a well-studied mechanism for species diversity). Each patch is characterized by a random environmental value that determines the optimal phenotype of its occupants. In such a scenario, only local adaptation and gene flow (migration) may interact to promote genetic variation and coexistence in the metapopulation. Results show that a competitively inferior species with adaptive phenotypic plasticity can coexist in a same patch with a competitively superior, non-plastic species, provided the migration rates and variances of the patches' environmental values are sufficiently large.

Colonization–competition dynamics of basal species shape food web complexity in island metacommunities

Exploring how food web complexity emerges and evolves in island ecosystems remains a major challenge in ecology. Food webs assembled from multiple islands are commonly recognized as highly complex trophic networks that are dynamic in both space and time. In the context of global climate change, it remains unclear whether food web complexity will decrease in a monotonic fashion when undergoing habitat destruction (e.g., the inundation of islands due to sea-level rise). Here, we develop a simple yet comprehensive patch-dynamic framework for complex food web metacommunities subject to the competition-colonization tradeoff between basal species. We found that oscillations in food web topological complexity (characterized by species diversity, mean food chain length and the degree of omnivory) emerge along the habitat destruction gradient. This outcome is robust to changing parameters or relaxing the assumption of a strict competitive hierarchy. Having oscillations in food web complexity indicates that small habitat changes could have disproportionate negative effects on species diversity, thus the success of conservation actions should be evaluated not only on changes in biodiversity, but also on system robustness to habitat alteration. Overall, this study provides a parsimonious mechanistic explanation for the emergence of food web complexity in island ecosystems, further enriching our understanding of metacommunity assembly.

Predicting evolution in experimental range expansions of an aquatic model system.

Predicting range expansion dynamics is an important goal of both fundamental and applied research in conservation and global change biology. However, this is challenging if ecological and evolutionary processes occur on the same time scale. Using the freshwater ciliate Paramecium caudatum, we combined experimental evolution and mathematical modeling to assess the predictability of evolutionary change during range expansions. In the experiment, we followed ecological dynamics and trait evolution in independently replicated microcosm populations in range core and front treatments, where episodes of natural dispersal alternated with periods of population growth. These eco-evolutionary conditions were recreated in a predictive mathematical model, parametrized with dispersal and growth data of the 20 founder strains in the experiment. We found that short-term evolution was driven by selection for increased dispersal in the front treatment and general selection for higher growth rates in all treatments. There was a good quantitative match between predicted and observed trait changes. Phenotypic divergence was further mirrored by genetic divergence between range core and front treatments. In each treatment, we found the repeated fixation of the same cytochrome c oxidase I (COI) marker genotype, carried by strains that also were the most likely winners in our model. Long-term evolution in the experimental range front lines resulted in the emergence of a dispersal syndrome, namely a competition-colonization trade-off. Altogether, both model and experiment highlight the potential importance of dispersal evolution as a driver of range expansions. Thus, evolution at range fronts may follow predictable trajectories, at least for simple scenarios, and predicting these dynamics may be possible from knowledge of few key parameters.

Ecological Determinants of Inter-Island Distributions through Occasional Dispersal of Two Closely Related Species, Varied Tit and Cinereous Tit, in the Volcanic Izu Archipelago, Japan

Inspired by ecologists R. H. MacArthur and E. O. Wilson, who pioneered the use of islands as ideal model systems for researching the formation and maintenance of biodiversity, we studied two sympatric passerine bird species on a volcanic archipelago, in Japan. We focused on the occupancy and co-occurrence of Varied Tit Sittiparus varius and Cinereous Tit Parus cinereus, both members of the Parus guild, on the Izu Islands. Their populations range from being stable, and co-occurring, to being temporally unstable (with local extinctions, invasions, and species replacement) and even totally absent. Using results from several surveys carried out on the islands over the last 60 years, and our own contemporary surveys, we tested the ecological drivers of inter-island occupancy through occasional dispersal of these two species. We found that both species had positive relationships with island size and vegetation diversity, while Cinereous Tit exhibited a negative associating with distance to the mainland (Honshu, Japan). Cinereous Tit was also negatively correlated with Varied Tit, but their co-occurrence was positively associated with island size, connectivity, and vegetation diversity. We suspect that resource constraints play a significant role in the distributions of the two species on the islands. Varied Tit is dominant over Cinereous Tit, but Cinereous Tit is able to utilize a wider range of habitats and resources. We also discuss the potential for a competition-colonization trade-off for the two species on the islands.

Ecological succession and the competition-colonization trade-off in microbial communities

BackgroundDuring range expansion in spatially distributed habitats, organisms differ from one another in terms of their patterns of localization versus propagation. To exploit locations or explore the landscape? This is the competition-colonization trade-off, a dichotomy at the core of ecological succession. In bacterial communities, this trade-off is a fundamental mechanism towards understanding spatio-temporal fluxes in microbiome composition.ResultsUsing microfluidics devices as structured bacterial habitats, we show that, in a synthetic two-species community of motile strains, Escherichia coli is a fugitive species, whereas Pseudomonas aeruginosa is a slower colonizer but superior competitor. We provide evidence highlighting the role of succession and the relevance of this trade-off in the community assembly of bacteria in spatially distributed patchy landscapes. Furthermore, aggregation-dependent priority effects enhance coexistence which is not possible in well-mixed environments.ConclusionsOur findings underscore the interplay between micron-scale landscape structure and dispersal in shaping biodiversity patterns in microbial ecosystems. Understanding this interplay is key to unleash the technological revolution of microbiome applications.

Breaking down the components of the competition-colonization trade-off: New insights into its role in diverse systems.

Performance trade-offs between competition and colonization can be an important mechanism facilitating regional coexistence of competitors. However, empirical evidence for this trade-off is mixed, raising questions about the extent to which it shapes diverse ecological communities. Here, we outline a framework that can be used to improve empirical tests of the competition-colonization trade-off. We argue that tests of the competition-colonization trade-off have been diverted into unproductive paths when dispersal mode and/or competition type have been inadequately defined. To generate comparative predictions of associations between dispersal and competitive performance, we develop a conceptual trait-based framework that clarifies how dispersal mode and type of competitor shape this trade-off at the stage of dispersal and establishment in a variety of systems. Our framework suggests that competition-colonization trade-offs may be less common for passively dispersing organisms when competitive dominants are those best able to withstand resource depletion (competitive response), and for active dispersers when traits for dispersal performance are positively associated with resource pre-emption (competitive effect). The framework presented here is designed to provide common ground for researchers working in different systems in order to prompt more effective assessment of this performance trade-off and its role in shaping community structure. By delineating key system properties that mediate the trade-off between competitive and colonization performance and their relationship to individual-level traits, researchers in disparate systems can structure their predictions about this trade-off more effectively and compare across systems more clearly.

Epidemiological dynamics of bacteriocin competition and antibiotic resistance.

Bacteriocins, toxic peptides involved in the competition between bacterial strains, are extremely diverse. Previous work on bacteriocin dynamics has highlighted the role of non-transitive ‘rock–paper–scissors’ competition in maintaining the coexistence of different bacteriocin profiles. The focus to date has primarily been on bacteriocin interactions at the within-host scale (i.e. within a single bacterial population). Yet in species such as Streptococcus pneumoniae, with relatively short periods of colonization and limited within-host diversity, ecological outcomes are also shaped by processes at the epidemiological (between-host) scale. Here, we first investigate bacteriocin dynamics and diversity in epidemiological models. We find that in these models, bacteriocin diversity is more readily maintained than in within-host models, and with more possible combinations of coexisting bacteriocin profiles. Indeed, maintenance of diversity in epidemiological models does not require rock–paper–scissors dynamics; it can also occur through a competition–colonization trade-off. Second, we investigate the link between bacteriocin diversity and diversity at antibiotic resistance loci. Previous work has proposed that bacterial duration of colonization modulates the fitness of antibiotic resistance. Due to their inhibitory effects, bacteriocins are a plausible candidate for playing a role in the duration of colonization episodes. We extend the epidemiological model of bacteriocin dynamics to incorporate an antibiotic resistance locus and demonstrate that bacteriocin diversity can indeed maintain the coexistence of antibiotic-sensitive and -resistant strains.

Quantifying mechanisms of coexistence in disease ecology.

Pathogen coexistence depends on ecological processes operating at both within and between-host scales, making it difficult to quantify which processes may promote or prevent coexistence. Here, we propose that adapting modern coexistence theory-traditionally applied in plant communities-to pathogen systems provides an exciting approach for examining mechanisms of coexistence operating across different spatial scales. We first overview modern coexistence theory and its mechanistic decomposition; we subsequently adapt the framework to quantify how spatial variation in pathogen density, host resources and immunity, and their interaction may promote pathogen coexistence. We apply this derivation to an example two pathogen, multiscale model comparing two scenarios with generalist and strain-specific immunity: one with demographic equivalency among pathogens and one with demographic trade-offs among pathogens. We then show how host-pathogen feedbacks generate spatial heterogeneity that promote pathogen coexistence and decompose those mechanisms to quantify how each spatial heterogeneity contributes to that coexistence. Specifically, coexistence of demographically equivalent pathogens occurs due to spatial variation in host resources, immune responses, and pathogen aggregation. With a competition-colonization trade-off, the superior colonizer requires spatial heterogeneity to coexist, whereas the superior competitor does not. Finally, we suggest ways forward for linking theory and empirical tests of coexistence in disease systems.

Coexistence under Hierarchical Resource Exploitation: The Role of the R*-Preemption Trade-Off.

AbstractResource competition theory predicts coexistence and exclusion patterns based on species' R*s, the minimum resource values required for a species to persist. A central assumption of the theory is that all species have equal access to resources. However, many systems are characterized by preemption exploitation, where some species deplete resources before their competitors can access them (e.g., asymmetric light competition, contest competition among animals). We hypothesized that coexistence under preemption requires an R*-preemption trade-off-that is, the species with the priority access should have a higher R* (lower "efficiency"). Thus, we developed an extension of resource competition theory to investigate partial and total preemption (in the latter, the preemptor is unaffected by species with lower preemption rank). We found that an R*-preemption trade-off is a necessary condition for coexistence in all models. Moreover, under total preemption, the trade-off alone is sufficient for coexistence. In contrast, under partial preemption, more conditions are needed, which restricts the parameter space of coexistence. Finally, we discuss the implications of our finding for seemingly distinct trade-offs, which we view as special cases of the R*-preemption trade-off. These trade-offs include the digger-grazer trade-off, the competition-colonization trade-off, and trade-offs related to light competition between trees and understories.

Vertical Stratification of Insect Species Developing in Water-Filled Tree Holes

Forest ecosystems have a distinct vertical dimension, but the structuring of communities in this three-dimensional space is not well understood. Water-filled tree holes are natural microcosms structured in metacommunities. Here, we used these microcosms as model systems to analyze how insect communities and the occurrence and abundance of individual species are influenced by biotic and abiotic microhabitat characteristics, the vertical position of the tree hole, and stand-scale habitat availability. We found that both the characteristics of water-filled tree holes and their insect communities differ significantly between canopy and ground level. Individual insect species showed contrasting responses to the vertical position of the tree holes when important environmental factors at the stand and the tree-hole scale were considered. While some species, such as the mosquito Aedes geniculatus and the beetle Prionocyphon serricornis, decreased in abundance with increasing tree-hole height, the biting midge Dasyhelea sp., the non-biting midge Metriocnemus cavicola and the hoverfly Myiatropa florea increased in abundance. Our results suggest that vertical stratification in forests is most likely driven not only by variation in tree-hole microhabitat properties, i.e., niche separation, but also by individual species traits, such as adult dispersal propensity, food preferences and mating behavior of adult stages, and interspecific competition of larval stages. Therefore, communities of insect species developing in tree holes are likely structured by competition–colonization trade-offs predicted by metacommunity theory.

Evolutionary coexistence in a metacommunity: Competition-colonization trade-off, ownership effects, environmental fluctuations

We study the adaptive dynamics of the colonization rate of species living in a patchy habitat when there is a trade-off with the competitive strength for individual patches. To that end, we formulate a continuous-time competition-colonization model that also includes ownership effects as well as random disturbance affecting the mortality rate. We find that intermediate disturbance (as measured by the fluctuation intensity of the mortality rate), a strong competition-colonization trade-off, and a weak ownership effect are necessary conditions for evolutionary branching and hence for the emergence of polymorphisms (i.e., coexistence) by small evolutionary steps. Specifically, concerning ownership we find that with low-intermediate disturbance, a weak ownership advantage favours evolutionary branching while ownership disadvantage does not. This asymmetry disappears at the higher-intermediate disturbance. Moreover, at a low-intermediate disturbance, the effect of the strength of the competition-colonization trade-off on evolutionary branching is non-monotonic disappears because the possibility of branching disappears again when the trade-off is too strong. We also find that there can be multiple evolutionary attractors for polymorphic populations, each with its own basin of attraction. With small but non-zero random evolutionary steps and depending on the initial polymorphic condition just after branching, a coevolutionary trajectory may come arbitrarily close to the shared boundary of two such basins and may even jump from one side to the other, which can lead to various kinds of long-term evolutionary dynamics, including evolutionary branching-extinction cycles.

No evidence for dominance–discovery trade-offs in Pheidole (Hymenoptera: Formicidae) assemblages

Understanding the mechanisms that allow species coexistence across spatial scales is of great interest to ecologists. Many such proposed mechanisms involve trade-offs between species in different life-history traits, with distinct trade-offs being expected to be prevalent at varying temporal and spatial scales. The dominance–discovery trade-off posits that species differ in their ability to find and use resources quickly, in contrast to their ability to monopolize those resources, a mechanism analogous to the competition-colonization trade-off. We investigated the occurrence of this structuring mechanism in the genus Pheidole Westwood, 1839 (Hymenoptera: Formicidae) assemblages in Atlantic Forest remnants. According to the dominance–discovery trade-off, consistent interspecific variation should be observed along the axis of discovery and dominance. We established 55 sampling units across two sites, with each unit consisting of a sardine bait monitored for 3 h. There was no distinction among Pheidole species in their ability to find or dominate food sources, suggesting that the dominance–discovery trade-off does not explain their coexistence. The low levels of aggression between Pheidole species could prevent the establishment of dominance hierarchies, whereas the species order of arrival at food sources could allow for resource partitioning through priority effects.

On the Cost-Effective Design of Agglomeration Bonus Schemes for the Conservation of Multiple Competing Species

An important mechanism of species co-existence in spatially structured landscapes is the competition-colonisation trade-off which states that co-existence of competing species is possible if, all other things equal, the better competitor is the worse coloniser. The effectiveness of this trade-off for the facilitation of co-existence, however, is likely to depend on the spatial arrangement of the habitat, because too strong agglomeration of the habitat may overly benefit the strong competitor (being the poor disperser), implying extinction of the inferiour competitor, while too much dispersion of the habitat may drive the superiour competitor (being the inferiour coloniser) to extinction. In working landscapes, biodiversity conservation is often induced through conservation payments that offset the forgone profits incurred by the conservation measure. To control the spatial arrangement of conservation measures and habitats in a conservation payment scheme, the agglomeration bonus has been proposed to provide financial incentives for allocating conservation measures in the vicinity of other sites with conservation measures. This paper presents a generic spatially explicit ecological-economic simulation model to explore the ability of the agglomeration bonus to cost-effectively conserve multiple competing species that differ by their competition strengths, their colonisation rates and their dispersal ranges. The interacting effects of the agglomeration bonus and different species traits and their trade-offs on the species richness in the model landscape are analysed. Recommendations for the biodiversity-maximising design of agglomeration bonus schemes are derived.