Limited niche differentiation within remarkable co‐occurrences of congeneric species: Monomorium ants in the Australian seasonal tropics

Niche theory predicts that few closely related species can co‐occur because such species tend to be ecologically similar and niche differentiation is required to avoid competitive exclusion. We analyse the co‐occurrence of a remarkable 10–15 species of the ant genus Monomorium occurring within single 10 × 10 m plots in a tropical savanna of northern Australia. Most of the species are undescribed, so we use genetic analysis to validate our species demarcations. We document nest dispersion patterns, and investigate differentiation in the three primary niche dimensions: space, time and food. We also examine species differences in competitive abilities, by describing rates of foraging activity, foraging ranges, worker aggression, and levels of behavioural dominance. Analyses of nest and forager distributions showed very limited evidence of spatial segregation within plots. The great majority of species foraged either exclusively or primarily during daylight hours. Body size and isotopic analyses indicated very limited dietary differentiation. Such limited niche partitioning occurred despite the species differing markedly in their competitive abilities as measured by rates of resource discovery, recruitment and monopolization. Our findings defy the traditional assumption that multiple closely related and ecologically similar species of highly interactive taxa cannot co‐occur. It seems very likely that species coexistence in our study system is determined to a very large degree by stochastic processes relating to dispersal and establishment, as predicted by neutral theory. However, neutral theory assumes competitive equivalence, whereas we found very marked differences in the competitive abilities of our co‐occurring species. We suggest that competitive exclusion is prevented by the modular nature of ant colonies, with competition limiting colony performance but not preventing occurrence. We conclude that other factors that allow species persistence, and not just competitive equivalence, can allow dispersal and establishment processes to drive species coexistence.     

Not enough niches: non‐equilibrial processes promoting species coexistence in diverse ant communities

Abstract Explanations for species coexistence in ant communities have traditionally focused on niche partitioning, particularly relating to differences in diet, foraging times and nesting requirements. Although niche separation is undoubtedly important, it seems insufficient to account for the high levels of local species richness that are commonly observed. This paper explores alternative explanations for ant species coexistence, focusing on factors that prevent competitive exclusion in diverse ant communities experiencing high levels of behavioural dominance, such as characteristically occurs in Australia. Very high species densities require two conditions to be met: first, a large number of species must successfully establish; and second, there must be a high rate of species persistence once established. In this context I advance five propositions based around three sets of arguments. First, ant sociality and modularity confers a high level of persistence once colonies are established, so that species coexistence is determined to a significant extent by processes operating at the establishment phase, rather than just by interactions between established colonies. Second, competitive outcomes are highly conditioned by environmental variation, which severely limits competitive exclusion. Finally, dominant species are highly patchy in space, and cannot comprehensively monopolize resources, such that there will usually be room for low densities of subordinate species. These propositions have relevance to neutral theories of community ecology, and to understanding intercontinental differences in local species richness.     

Concurrent niche and neutral processes in the competition-colonization model of species coexistence

The importance of neutral dynamics is contentiously debated in the ecological literature. This debate focuses on neutral theory's assumption of fitness equivalency among individuals, which conflicts with stabilizing fitness that promotes coexistence through niche differentiation. I take advantage of competition-colonization trade-offs between species of aquatic micro-organisms (protozoans and rotifers) to show that equalizing and stabilizing mechanisms can operate simultaneously. Competition trials between species with similar colonization abilities were less likely to result in competitive exclusion than for species further apart. While the stabilizing mechanism (colonization differences) facilitates coexistence at large spatial scales, species with similar colonization abilities also exhibited local coexistence probably due to fitness similarities allowing weak stabilizing mechanisms to operate. These results suggest that neutral- and niche-based mechanisms of coexistence can simultaneously operate at differing temporal and spatial scales, and such a spatially explicit view of coexistence may be one way to reconcile niche and neutral dynamics.     

Niche partitioning at multiple scales facilitates coexistence among mosquito larvae

A theoretical dichotomy in community ecology distinguishes between mechanisms that stabilize species coexistence and those that cause neutral drift. Stable coexistence is predicted to occur in communities where competing species have niche-partitioning mechanisms that reduce interspecific competition. Neutral communities are predicted to be structured by stochastic processes that are not influenced by species identity, but that may be influenced by priority effects and dispersal limitation. Recent developments have suggested that neutral interactions may be more common at local scales, while niche structuring may be more common at larger scales. We tested for mechanisms that could promote either stable coexistence or neutral drift in a bromeliad-dwelling mosquito community by evaluating A) if a hypothesized within-bromeliad niche partitioning mechanism occurs in the community, B) if this mechanism correlates with local species co-occurrence patterns, and C) if patterns of coexistence at the larger (metacommunity) scale were consistent with those at the local scale. We found that mosquitoes in this community do partition space within containers, and that species with the strongest potential for competition co-occurred least. Species with overlapping spatial niches minimized co-occurrence by specialising in bromeliads of differing sizes, effectively changing the scale at which they coexist. In contrast, we found no evidence to support neutral dynamics in mosquito communities at either scale. In this community, a niche-based mechanism that is predicted to stabilize species coexistence explains co-occurrence patterns within and among bromeliads.     

Resisting annihilation: relationships between functional trait dissimilarity,assemblage competitive power and allelopathy

Allelopathic species can alter biodiversity. Using simulated assemblages that are characterised by neutrality, lumpy coexistence and intransitivity, we explore relationships between within‐assemblage competitive dissimilarities and resistance to allelopathic species. An emergent behaviour from our models is that assemblages are more resistant to allelopathy when members strongly compete exploitatively (high competitive power). We found that neutral assemblages were the most vulnerable to allelopathic species, followed by lumpy and then by intransitive assemblages. We find support for our modeling in real‐world time‐series data from eight lakes of varied morphometry and trophic state. Our analysis of this data shows that a lake's history of allelopathic phytoplankton species biovolume density and dominance is related to the number of species clusters occurring in the plankton assemblages of those lakes, an emergent trend similar to that of our modeling. We suggest that an assemblage's competitive power determines its allelopathy resistance.     

Estimates of species extinctions from species–area relationships strongly depend on ecological context

Species–area (SAR) and endemics–area (EAR) relationships are amongst the most common methods used to forecast species loss resulting from habitat loss. One critical, albeit often ignored, limitation of these area‐based estimates is their disregard of the ecological context that shapes species distributions. In this study, we estimate species loss using a spatially explicit mechanistic simulation model to evaluate three important aspects of ecological context: coexistence mechanisms (e.g. species sorting, competition–colonization tradeoffs and neutral dynamics), spatial distribution of environmental conditions, and spatial pattern of habitat loss. We found that 1) area‐based estimates of extinctions are sensitive to coexistence mechanisms as well as to the pattern of environmental heterogeneity; 2) there is a strong interaction between coexistence mechanisms and the pattern of habitat loss; 3) SARs always yield higher estimates of species loss than do EARs; and 4) SARs and EARs consistently underestimate the realized species loss. Our results highlight the need to integrate ecological mechanisms in area‐estimates of species loss.     

Habitat disturbance modifies dominance,coexistence, and competitive interactions in tropical ant communities

1. Interspecific competition is a major structuring principle in ecological communities. Despite their prevalence, the outcome of competitive interactions is hard to predict, highly context-dependent, and multiple factors can modulate such interactions. 2. We tested predictions concerning how competitive interactions are modified by anthropogenic habitat disturbance in ground-foraging ant assemblages inhabiting fragmented Inter-Andean tropical dry forests in southwestern Colombia, and investigated ant assemblages recruiting to baits in 10 forest fragments exposed to varying level of human disturbance. 3. Specifically, we evaluated how different components of competitive interactions (patterns of species co-occurrence, resource partitioning, numerical dominance, and interspecific trade-offs between discovery and dominance competition) varied with level of habitat disturbance in a human-dominated ecosystem. 4. Multiple lines of evidence suggest that the role of competitive interactions in structuring ground-foraging ant communities at baits varied with respect to habitat disturbance. As disturbance increased, community structure was more likely to exhibit random co-occurrence patterns, higher levels of monopolization of food resources by dominant ants, and disproportionate dominance of a single species, the little fire ant (Wasmannia auropunctata). At a regional scale, we found evidence for a trade-off between dominance and discovery abilities of the 15 most common species at baits. 5. Together, these results suggest that human disturbance modifies the outcome of competitive interactions in ground-foraging ant assemblages and may promote dominant species that reduce diversity and coexistence in tropical ecosystems.     

The effects of metapopulation structure on indirect interactions in host-parasitoid assemblages.

The interaction between two species that do not compete for resources but share a common natural enemy is known as apparent competition. In the absence of other limiting factors, such three-species interactions are impermanent, with one species being excluded from the assemblage by the natural enemy. Here, the effects of metapopulation structure are explored in a system of two hosts that experience apparent competition through a shared parasitoid. A coupled-map lattice model is developed and used to explore species coexistence and patterns of patch occupancy at the metapopulation scale. Linking local and regional dynamics favours coexistence by uncoupling the dynamics of the three species in space. Coexistence is promoted by the inferior species being either a fugitive or a sedentary species. The occurrence of these two mutually exclusive mechanisms of coexistence is influenced by the relative dispersal of the inferior apparent competitor.     

Phylogenetic relatedness and the determinants of competitive outcomes

Recent hypotheses argue that phylogenetic relatedness should predict both the niche differences that stabilise coexistence and the average fitness differences that drive competitive dominance. These still largely untested predictions complicate Darwin's hypothesis that more closely related species less easily coexist, and challenge the use of community phylogenetic patterns to infer competition. We field parameterised models of competitor dynamics with pairs of 18 California annual plant species, and then related species' niche and fitness differences to their phylogenetic distance. Stabilising niche differences were unrelated to phylogenetic distance, while species' average fitness showed phylogenetic structure. This meant that more distant relatives had greater competitive asymmetry, which should favour the coexistence of close relatives. Nonetheless, coexistence proved unrelated to phylogeny, due in part to increasing variance in fitness differences with phylogenetic distance, a previously overlooked property of such relationships. Together, these findings question the expectation that distant relatives should more readily coexist.     

Spatial Complementarity and the Coexistence of Species

Coexistence of apparently similar species remains an enduring paradox in ecology. Spatial structure has been predicted to enable coexistence even when population-level models predict competitive exclusion if it causes each species to limit its own population more than that of its competitor. Nevertheless, existing hypotheses conflict with regard to whether clustering favours or precludes coexistence. The spatial segregation hypothesis predicts that in clustered populations the frequency of intra-specific interactions will be increased, causing each species to be self-limiting. Alternatively, individuals of the same species might compete over greater distances, known as heteromyopia, breaking down clusters and opening space for a second species to invade. In this study we create an individual-based model in homogeneous two-dimensional space for two putative sessile species differing only in their demographic rates and the range and strength of their competitive interactions. We fully characterise the parameter space within which coexistence occurs beyond population-level predictions, thereby revealing a region of coexistence generated by a previously-unrecognised process which we term the triadic mechanism. Here coexistence occurs due to the ability of a second generation of offspring of the rarer species to escape competition from their ancestors. We diagnose the conditions under which each of three spatial coexistence mechanisms operates and their characteristic spatial signatures. Deriving insights from a novel metric — ecological pressure — we demonstrate that coexistence is not solely determined by features of the numerically-dominant species. This results in a common framework for predicting, given any pair of species and knowledge of the relevant parameters, whether they will coexist, the mechanism by which they will do so, and the resultant spatial pattern of the community. Spatial coexistence arises from complementary combinations of traits in each species rather than solely through self-limitation.     

Fusing spatial resource heterogeneity with a competition–colonization trade-off in model communities

Two commonly cited mechanisms of multispecies coexistence in patchy environments are spatial heterogeneity in competitive abilities caused by variation in resources and a competition–colonization trade-off. In this paper, a model that fuses these mechanisms together is presented and analyzed. The model suggests that spatial variation in resource ratios can lead to multispecies coexistence, but this mechanism by itself is weak when the number of resources for which species compete is small. However, spatial resource heterogeneity is a powerful mechanism for multispecies coexistence when it acts synergistically with a competition–colonization trade-off. The model also shows how resource supply can control the competitive balance between species that are weak competitors but superior colonizers and strong competitors/inferior colonizers. This provides additional theoretical support for a possible explanation of empirically observed hump-shaped relationships between species diversity and ecological productivity.     

Mobility unevenness in rock–paper–scissors models

We investigate a tritrophic system whose cyclic dominance is modelled by the rock–paper–scissors game. We consider that organisms of one or two species are affected by movement limitations, which unbalances the cyclic spatial game. Performing stochastic simulations, we show that mobility unevenness controls the population dynamics. In the case of one slow species, the predominant species depends on the level of mobility restriction, with the slow species being preponderant if the mobility limitations are substantial. If two species face mobility limitations, our outcomes show that being higher dispersive does not constitute an advantage in terms of population growth. On the contrary, if organisms move with higher mobility, they expose themselves to enemies more frequently, being more vulnerable to being eliminated. Finally, our findings show that biodiversity benefits in regions where species are slowed. Biodiversity loss for high mobility organisms, common to cyclic systems, may be avoided with coexistence probability being higher for robust mobility limitations. Our results may help biologists understand the dynamics of unbalanced spatial systems where organisms’ dispersal is fundamental to biodiversity conservation.     

Microbial expansion–collision dynamics promote cooperation and coexistence on surfaces

Microbes colonizing a surface often experience colony growth dynamics characterized by an initial phase of spatial clonal expansion followed by collision between neighboring colonies to form potentially genetically heterogeneous boundaries. For species with life cycles consisting of repeated surface colonization and dispersal, these spatially explicit “expansion‐collision dynamics” generate periodic transitions between two distinct selective regimes, “expansion competition” and “boundary competition,” each one favoring a different growth strategy. We hypothesized that this dynamic could promote stable coexistence of expansion‐ and boundary‐competition specialists by generating time‐varying, negative frequency‐dependent selection that insulates both types from extinction. We tested this experimentally in budding yeast by competing an exoenzyme secreting “cooperator” strain (expansion–competition specialists) against nonsecreting “defectors” (boundary–competition specialists). As predicted, we observed cooperator–defector coexistence or cooperator dominance with expansion–collision dynamics, but only defector dominance otherwise. Also as predicted, the steady‐state frequency of cooperators was determined by colonization density (the average initial cell–cell distance) and cost of cooperation. Lattice‐based spatial simulations give good qualitative agreement with experiments, supporting our hypothesis that expansion–collision dynamics with costly public goods production is sufficient to generate stable cooperator–defector coexistence. This mechanism may be important for maintaining public–goods cooperation and conflict in microbial pioneer species living on surfaces.     

Local‐scale spatial dynamics of ants in a temperate agroecosystem

At the local scale, spatial aggregations in ant distribution are often thought to be driven by competitive interactions among dominant ant species, although niche preferences and habitat heterogeneity might also lead to patchiness. Nevertheless, competitive interactions might be particularly important in agroecosystems that are structurally more homogeneous than natural habitats. The spatial patterns of ants in two Australian vineyards were investigated by intensive pitfall trapping to examine if non‐random patterns occur and whether these might be the result of competitive species interactions as well as the influence of woody vegetation adjacent to the vineyards. Null model analyses suggested competitive species interactions within ant assemblages that might have been driven by numerically dominant species, even though both positive and negative associations between these were found. Consistent spatial aggregations indicated significant spatial overlap in distributions of some species. Such overlap suggests that potential coexistence might be attributed to temporal partitioning or differences in foraging strategies. The presence of woody vegetation had a marked influence on ant assemblage structure and competitive interactions, and might facilitate coexistence by increasing resource heterogeneity. The implications of these findings for sampling strategies and ecological processes within vineyards are discussed.     

How variation between individuals affects species coexistence

Although the effects of variation between individuals within species are traditionally ignored in studies of species coexistence, the magnitude of intraspecific variation in nature is forcing ecologists to reconsider. Compelling intuitive arguments suggest that individual variation may provide a previously unrecognised route to diversity maintenance by blurring species‐level competitive differences or substituting for species‐level niche differences. These arguments, which are motivating a large body of empirical work, have rarely been evaluated with quantitative theory. Here we incorporate intraspecific variation into a common model of competition and identify three pathways by which this variation affects coexistence: (1) changes in competitive dynamics because of nonlinear averaging, (2) changes in species’ mean interaction strengths because of variation in underlying traits (also via nonlinear averaging) and (3) effects on stochastic demography. As a consequence of the first two mechanisms, we find that intraspecific variation in competitive ability increases the dominance of superior competitors, and intraspecific niche variation reduces species‐level niche differentiation, both of which make coexistence more difficult. In addition, individual variation can exacerbate the effects of demographic stochasticity, and this further destabilises coexistence. Our work provides a theoretical foundation for emerging empirical interests in the effects of intraspecific variation on species diversity.     

Scaling issues of neutral theory reveal violations of ecological equivalence for dominant Amazonian tree species

Neutral models are often used as null models, testing the relative importance of niche versus neutral processes in shaping diversity. Most versions, however, focus only on regional scale predictions and neglect local level contributions. Recently, a new formulation of spatial neutral theory was published showing an incompatibility between regional and local scale fits where especially the number of rare species was dramatically under‐predicted. Using a forward in time semi‐spatially explicit neutral model and a unique large‐scale Amazonian tree inventory data set, we show that neutral theory not only underestimates the number of rare species but also fails in predicting the excessive dominance of species on both regional and local levels. We show that although there are clear relationships between species composition, spatial and environmental distances, there is also a clear differentiation between species able to attain dominance with and without restriction to specific habitats. We conclude therefore that the apparent dominance of these species is real, and that their excessive abundance can be attributed to fitness differences in different ways, a clear violation of the ecological equivalence assumption of neutral theory.     

Coupling of two competitive systems via density dependent migration

Coupling of two Lotka–Volterra type competition systems with density-dependent migration was surveyed. We assumed that species x and y are each exclusively superior in subhabitat 1 and subhabitat 2, respectively, and that population densities that exert intra-and interspecific competitive effects also impose pressures for migration of individuals from a subhabitat. If the two species are, respectively, abundant in the subhabitats in which either species is competitively superior, and the migration has a mixing effect, then, it would be intuitively expected that, as potential migration rates increase, the two species are mixed well and coexist in the whole habitat. An analysis of this competitive situation using our model under the assumption of linear diffusion predicted that, even though weak mixing maintains coexistence in the whole habitat, strong mixing collapses coexistence and leads to the exclusion of one species. The assumption that migrations occur due to self- and cross-population pressures provides different predictions: (i) weak dominance and strong mixing destabilize the coexistence state and lead to a monopolizing equilibrium of either species (bi-stability of monopolizing equiliblia); (ii) conspicuous weakness of the inferior species makes the mixing equilibrium stable, regardless of the potential migration rate; and (iii) tri-stability exists in between situations (i) and (ii). In the third case, the attainable state is the mixing equilibrium or either of the monopolizing equilibria, depending on the initial state. Migration mechanisms with self- and cross-population pressures tends to mediate spatial segregation and makes coexistence possible, even with strong mixing.     

Negative density dependence can offset the effect of species competitive asymmetry: a niche-based mechanism for neutral-like patterns

The debate on the role of species differences in shaping biodiversity patterns, with its two extremes of pure niche theory and neutral theory, is still ongoing. It has been demonstrated that a slight difference in competitive ability of species severely affects the predictions of the neutral model. At the same time, neutral patterns seem to be ubiquitous. Here, we model both negative density dependence (NDD) and competitive asymmetry (CA) simultaneously. Our simulation results show that an appropriate intensity of NDD can offset the negative effect of CA (modeled as fecundity difference) on species coexistence and produce a neutral-like species abundance distribution. Therefore, our model provides a plausible mechanistic explanation of neutral-like patterns, but contrary to the neutral model, a species' relative abundance is positively related to its competitive ability in our model.     

Coexistence between native and exotic species is facilitated by asymmetries in competitive ability and susceptibility to herbivores

Differences between native and exotic species in competitive ability and susceptibility to herbivores are hypothesized to facilitate coexistence. However, little fieldwork has been conducted to determine whether these differences are present in invaded communities. Here, we experimentally examined whether asymmetries exist between native and exotic plants in a community invaded for over 200 years and whether removing competitors or herbivores influences coexistence. We found that natives and exotics exhibit pronounced asymmetries, as exotics are competitively superior to natives, but are more significantly impacted by herbivores. We also found that herbivore removal mediated the outcome of competitive interactions and altered patterns of dominance across our field sites. Collectively, these findings suggest that asymmetric biotic interactions between native and exotic plants can help to facilitate coexistence in invaded communities.