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.     

Tradeoffs, competition, and coexistence in eastern deciduous forest ant communities

Ecologists have long sought to explain the coexistence of multiple potentially competing species in local assemblages. This is especially challenging in species-rich assemblages in which interspecific competition is intense, as it often is in ant assemblages. As a result, a suite of mechanisms has been proposed to explain coexistence among potentially competing ant species: the dominance–discovery tradeoff, the dominance–thermal tolerance tradeoff, spatial segregation, temperature-based niche partitioning, and temporal niche partitioning. Through a series of observations and experiments, we examined a deciduous forest ant assemblage in eastern North America for the signature of each of these coexistence mechanisms. We failed to detect evidence for any of the commonly suggested mechanisms of coexistence, with one notable exception: ant species appear to temporally partition foraging times such that behaviourally dominant species foraged more intensely at night, while foraging by subdominant species peaked during the day. Our work, though focused on a single assemblage, indicates that many of the commonly cited mechanisms of coexistence may not be general to all ant assemblages. However, temporal segregation may play a role in promoting coexistence among ant species in at least some ecosystems, as it does in many other organisms.     

Trait adaptation promotes species coexistence in diverse predator and prey communities

Species can adjust their traits in response to selection which may strongly influence species coexistence. Nevertheless, current theory mainly assumes distinct and time‐invariant trait values. We examined the combined effects of the range and the speed of trait adaptation on species coexistence using an innovative multispecies predator–prey model. It allows for temporal trait changes of all predator and prey species and thus simultaneous coadaptation within and among trophic levels. We show that very small or slow trait adaptation did not facilitate coexistence because the stabilizing niche differences were not sufficient to offset the fitness differences. In contrast, sufficiently large and fast trait adaptation jointly promoted stable or neutrally stable species coexistence. Continuous trait adjustments in response to selection enabled a temporally variable convergence and divergence of species traits; that is, species became temporally more similar (neutral theory) or dissimilar (niche theory) depending on the selection pressure, resulting over time in a balance between niche differences stabilizing coexistence and fitness differences promoting competitive exclusion. Furthermore, coadaptation allowed prey and predator species to cluster into different functional groups. This equalized the fitness of similar species while maintaining sufficient niche differences among functionally different species delaying or preventing competitive exclusion. In contrast to previous studies, the emergent feedback between biomass and trait dynamics enabled supersaturated coexistence for a broad range of potential trait adaptation and parameters. We conclude that accounting for trait adaptation may explain stable and supersaturated species coexistence for a broad range of environmental conditions in natural systems when the absence of such adaptive changes would preclude it. Small trait changes, coincident with those that may occur within many natural populations, greatly enlarged the number of coexisting species.     

Does “supersaturated coexistence” resolve the “paradox of the plankton”?

In contradiction with field observations, theory predicts that the number of coexisting plankton species at equilibrium cannot exceed the number of limiting resources, which is called the "paradox of the plankton". Recently, Huisman & Weissing (1999 , 2000 ) showed, in a model study, that the number of coexisting species may exceed the number of limiting resources when internal system feedback induces oscillations or chaos. In this paper, we use the term "supersaturated coexistence" for this phenomenon. On the basis of these findings, they claimed that the paradox of the plankton is solved. We investigated the prerequisites for supersaturated coexistence in the same model. Our results indicate that supersaturated coexistence is a rare phenomenon in parameter space, requires a very precise parameterization of the community members and is sensitive to the introduction of new species and the removal of the present species. This raises the question of whether supersaturated coexistence is likely to occur in nature. We conclude that the claim by Huisman & Weissing (1999 , 2000 ) is premature.     

Exploring taxonomic filtering in urban environments

Question: Several mechanisms have been proposed that control the spatio‐temporal pattern of species coexistence. Among others, the species pool hypothesis states that the large‐scale species pool is an important factor in controlling small‐scale species richness through filtering of species that can persist within a species assemblage on the basis of their tolerance of the abiotic environment. Because of the process of environmental filtering, co‐occurring species that experience similar environmental conditions are likely to be more taxonomically similar than ecologically distant species. This is because, due to the conservatism of many species traits during evolutionary diversification, the ability of species to colonize the same ecological space is thought to depend at least partially on their taxonomic similarity. The question for this study is: Under the assumption of trait conservatism, does environmental filtering lead to nonrandom species assemblages with respect to their taxonomic structure? Methods: The significance of taxonomic filtering in regulating species coexistence is tested using data from 15 local species assemblages from the urban flora of Rome (Italy). To find out whether the taxonomic structure of the selected’ local’ species assemblages was significantly different from random, we used a Monte Carlo simulation in which for each local species assemblage, the actual taxonomic diversity was compared to the taxonomic diversity of 1000 virtual species lists of the same size extracted at random from a larger ‘regional’ species pool. Results: We found that in most cases the local species assemblages have a higher degree of taxonomic similarity than would be expected by chance showing a phenomenon of ‘species condensation’ in a small number of higher‐level taxa. Conclusions: Our observations support the species pool hypothesis and imply that environmental filtering is an important mechanism in shaping the taxonomic structure of species assemblages. Therefore, the incorporation of taxonomic diversity into landscape and community ecology may be beneficial for a better understanding of the processes that regulate species coexistence.     

Species coexistence of communities with intraguild predation: The role of refuges used by the resource and the intraguild prey

In this paper, we develop a three-species intraguild predation model which incorporates refuges used by the resource and the intraguild prey, and focus on the effects of refuges on the three species coexistence. The invasion condition and parameter region for coexistence are obtained using invasion analysis. The new invasion condition requires that all boundary states with one missing species can be invaded by the missing species. Numerical simulations show that refuges have a major influence on species coexistence of intraguild predation system, and the results strongly depend on the types of refuges introduced into the model. Our study also shows that prey's refuges are detrimental to species coexistence except the resource using refuges. In contrast to previous research, we find that spatial structure may play an important role in effects of refuges on species coexistence of intraguild predation systems. Our results may shed new light on understanding the mechanisms and the persistence of multi-species predators-prey system.     

Coexistence of nearly neutral species

Aims The neutral theory of biodiversity provides a powerful framework for modeling macroecological patterns and interpreting species assemblages. However, there remain several unsolved problems, including the effect of relaxing the assumption of strict neutrality to allow for empirically observed variation in vital rates and the 'problem of time'—empirically measured coexistence times are much shorter than the prediction of the strictly neutral drift model. Here, we develop a nearly neutral model that allows for differential birth and death rates of species. This model provides an approach to study species coexistence away from strict neutrality.Methods Based on Moran's neutral model, which assumes all species in a community have the same competitive ability and have identical birth and death rates, we developed a model that includes birth–death trade-off but excludes speciation. This model describes a wide range of asymmetry from strictly neutral to nearly neutral to far from neutral and is useful for analyzing the effect of drift on species coexistence. Specifically, we analyzed the effects of the birth–death trade-off on the time and probability of species coexistence and quantified the loss of biodiversity (as measured by Simpson's diversity) due to drift by varying species birth and death rates.Important findings We found (i) a birth–death trade-off operating as an equalizing force driven by demographic stochasticity promotes the coexistence of nearly neutral species. Species near demographic trade-offs (i.e. fitness equivalence) can coexist even longer than that predicted by the strictly neutral model; (ii) the effect of birth rates on species coexistence is very similar to that of death rates, but their compensatory effects are not completely symmetric; (iii) ecological drift over time produces a march to fixation. Trade-off-based neutral communities lose diversity more slowly than the strictly neutral community, while non-neutral communities lose diversity much more rapidly; and (iv) nearly neutral systems have substantially shorter time of coexistence than that of neutral systems. This reduced time provides a promising solution to the problem of time.     

Trait plasticity alters the range of possible coexistence conditions in a competition–colonisation trade‐off

Most of the classical theory on species coexistence has been based on species‐level competitive trade‐offs. However, it is becoming apparent that plant species display high levels of trait plasticity. The implications of this plasticity are almost completely unknown for most coexistence theory. Here, we model a competition–colonisation trade‐off and incorporate trait plasticity to evaluate its effects on coexistence. Our simulations show that the classic competition–colonisation trade‐off is highly sensitive to environmental circumstances, and coexistence only occurs in narrow ranges of conditions. The inclusion of plasticity, which allows shifts in competitive hierarchies across the landscape, leads to coexistence across a much broader range of competitive and environmental conditions including disturbance levels, the magnitude of competitive differences between species, and landscape spatial patterning. Plasticity also increases the number of species that persist in simulations of multispecies assemblages. Plasticity may generally increase the robustness of coexistence mechanisms and be an important component of scaling coexistence theory to higher diversity communities.     

Temporal resource partitioning mediates vertebrate coexistence at carcasses: The role of competitive and facilitative interactions

Unravelling how biodiversity is maintained despite species competition for shared resources has been a central question in community ecology, and is gaining relevance amidst the current biodiversity crisis. Yet, we have still a poor understanding of the mechanisms that regulate species coexistence and shape the structure of assemblages in highly competitive environments such as carrion pulsed resources. Here, we study how large vertebrates coexist in scavenger assemblages by adapting their diel activity at large ungulate carcasses in NW Spain. We used camera traps to record vertebrate scavengers consuming 34 carcasses of livestock and hunted wild ungulates, which allowed us to assess also differences regarding carcass origin. To evaluate temporal resource partition among species, we estimated the overlap of diel activity patterns and the mean times of each scavenger at carcasses. We recorded 16 species of scavengers, 7 mammals and 9 birds, and found similar richness at both types of carcasses. Birds and mammals showed contrasting diel activity patterns, with birds using carcasses during daytime (mean= 11:38 h) and mammals mostly at night (23:09 h). The unimodal activity patterns of scavengers showed asynchronous peaks among species. Subordinate species modified their activity patterns at carcasses used by apex species to reduce temporal overlap. Also, diel activity patterns of vultures closely followed those of corvids, suggesting facilitation processes in which corvids would enhance carcass detection by vultures. Two mammal species (12.5%) increased nocturnality at carcasses of hunted ungulates, which could be a response to human disturbance. Our results suggest that both temporal segregation and coupling mediate the coexistence of large vertebrates at carcasses. These mechanisms might lead to richer scavenger assemblages and thereby more efficient ones in driving critical ecosystem functions related to carrion consumption, such as energy and nutrient recycling and biodiversity maintenance.     

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.     

Coexistence via resource partitioning fails to generate an increase in community function

Classic ecological theory suggests that resource partitioning facilitates the coexistence of species by reducing inter-specific competition. A byproduct of this process is an increase in overall community function, because a greater spectrum of resources can be used. In contrast, coexistence facilitated by neutral mechanisms is not expected to increase function. We studied coexistence in laboratory microcosms of the bactivorous ciliates Paramecium aurelia and Colpidium striatum to understand the relationship between function and coexistence mechanism. We quantified population and community-level function (biomass and oxygen consumption), competitive interactions, and resource partitioning. The two ciliates partitioned their bacterial resource along a size axis, with the larger ciliate consuming larger bacteria than the smaller ciliate. Despite this, there was no gain in function at the community level for either biomass or oxygen consumption, and competitive effects were symmetrical within and between species. Because other potential coexistence mechanisms can be ruled out, it is likely that inter-specific interference competition diminished the expected gain in function generated by resource partitioning, leading to a system that appeared competitively neutral even when structured by niche partitioning. We also analyzed several previous studies where two species of protists coexisted and found that the two-species communities showed a broad range of biomass levels relative to the single-species states.     

Does predator abundance influence species diversity of equilibrium macroinvertebrate assemblages in spring fens?

1. Predation may significantly control number and density of coexisting species. The effects of predation on species diversity have traditionally been tested in experiments and theoretical models of simple trophic systems. In complex natural ecosystems, however, disentangling multiple sources of variation is difficult. In groundwater-fed environments, a significant effect of predation can be expected due to the relatively stable environmental conditions; however, it has never been properly examined.
2. We analysed species diversity and total abundance of macroinvertebrate assemblages in 48 Western Carpathian spring fens, separately for whole sites and mesohabitat/season, and partitioned the effects of predation intensity from those of environmental variables in robust models using a bootstrapping technique. We verified our results by accounting for taxa resistant to predation.
3. The assumption that predation-mediated coexistence of species is the main mechanism responsible for the relatively species-rich assemblages in the Western Carpathian spring fens was not supported. However, predation may significantly influence abundance of non-predatory species and, under some conditions, it may contribute to explaining patterns in species diversity.
4. The effect of predation did not differ between the mesohabitats with different stability. However, we found higher environmental control in spring and a stronger effect of predators in autumn, which suggests that different mechanisms influence fen assemblages in different seasons.
5. Our study provides a new robust approach how to test the effect of predation on natural macroinvertebrate assemblages. The importance of predation was lower than expected in equilibrium assemblages but it may vary in time.

     

Phytoplankton Assemblage Characteristics in Recurrently Fluctuating Environments

Annual variations in biogeochemical and physical processes can lead to nutrient variability and seasonal patterns in phytoplankton productivity and assemblage structure. In many coastal systems river inflow and water exchange with the ocean varies seasonally, and alternating periods can arise where the nutrient most limiting to phytoplankton growth switches. Transitions between these alternating periods can be sudden or gradual and this depends on human activities, such as reservoir construction and interbasin water transfers. How such activities might influence phytoplankton assemblages is largely unknown. Here, we employed a multispecies, multi-nutrient model to explore how nutrient loading switching mode might affect characteristics of phytoplankton assemblages. The model is based on the Monod-relationship, predicting an instantaneous growth rate from ambient inorganic nutrient concentrations whereas the limiting nutrient at any given time was determined by Liebig’s Law of the Minimum. Our simulated phytoplankton assemblages self-organized from species rich pools over a 15-year period, and only the surviving species were considered as assemblage members. Using the model, we explored the interactive effects of complementarity level in trait trade-offs within phytoplankton assemblages and the amount of noise in the resource supply concentrations. We found that the effect of shift from a sudden resource supply transition to a gradual one, as observed in systems impacted by watershed development, was dependent on the level of complementarity. In the extremes, phytoplankton species richness and relative overyielding increased when complementarity was lowest, and phytoplankton biomass increased greatly when complementarity was highest. For low-complementarity simulations, the persistence of poorer-performing phytoplankton species of intermediate R*s led to higher richness and relative overyielding. For high-complementarity simulations, the formation of phytoplankton species clusters and niche compression enabled higher biomass accumulation. Our findings suggest that an understanding of factors influencing the emergence of life history traits important to complementarity is necessary to predict the impact of watershed development on phytoplankton productivity and assemblage structure.     

Structure of multi-species ant assemblages (Hymenoptera,Formicidae) in the Mountain Crimea

The multi-species ant assemblages were studied in the Crimea Mountains, in 9 types of plant associations in the plateau pastures as well as on the southern and northern slopes of the Main Range. A list of 18 ant species with data on their distribution in biotopes and also layers and the structure of their foraging territories is given. The hierarchical organization of multi-species ant assemblages and different strategies of nest location of the influents depending on the dominant species and its territoriality were investigated. Territoriality of each dominant species was determined by the frequency of foraging areas without nests of other species. Depending on dominant territoriality and the number of dominant species, the following classification of multi-species assemblages is proposed. (1) Monodominant assemblages with weakly expressed territoriality of the dominant species. Low density of the workers of Camponotus aethiops in the foraging territory determines weak interactions with influent species. The settling pattern of the influents shows no evident dependence on the dominant. Weakly expressed territoriality of the dominant allows non-territorial species to exist on the dominant’s territory. Such species were not found in other types of ant assemblages. (2) Monodominant assemblages with strongly expressed territoriality of the dominant species. The settling pattern of the influents shows the subordinate species to avoid those of higher ranks in the hierarchy, thus decreasing the probability of conflicts. The influents settle in the neutral zones as well as in the foraging territory of the dominant, so as to reduce competition with the subdominant species Formica cunicularia. Such a strategy reduces the probability of interspecific conflicts in biotopes with fewer vegetation layers than in the forests. (3) Bidominant assemblages with a variable dendrobiont dominant are typical among forest assemblages. Coexistence of two dominant species is possible when foraging of one of them is completely or partly restricted to one vegetation layer. The subdominant species avoid settling in the foraging territory of the dominant that occupies the same layer. When the dendrobiont dominant Crematogaster schmidti drops out of the assemblage, it (and its role in the assemblage) is replaced by other dendrobiont species Lasius emarginatus which otherwise acts as the 1st order subdominant in C. schmidti assemblages. (4) Polydominant assemblages are characterized by the presence of as many as 3 dominant species, namely Crematogaster schmidti, Camponotus aethiops, and Plagiolepis tauricus. The coexistence of the dominants is possible due to use of different layers (one species is a dendrobiont, the other two are herpeto-hortobionts), different activity periods (round-the-clock in Crematogaster schmidti, night in Camponotus aethiops, and daytime in P. tauricus), and different size classes of the workers (3rd, 5th, and 1st, respectively). The different size classes of workers reduce competition for food. Thus, when the number of dominant species in the assemblage increases, so does the number of mechanisms facilitating their coexistence.     

Life-history trade-offs and ecological dynamics in the evolution of longevity

Longevity is a life-history trait that is shaped by natural selection. An unexplored consequence is how selection on this trait affects diversity and diversification in species assemblages. Motivated by the diverse rockfish (Sebastes) assemblage in the North Pacific, the effects of trade-offs in longevity against competitive ability are explored. A competition model is developed and used to explore the potential for species diversification and coexistence. Invasion analyses highlight that life-history trait trade-offs in longevity can mitigate the effects of competitive ability and favour the coexistence of a finite number of species. Our results have implications for niche differentiation, limiting similarity and assembly dynamics in multispecies interactions.     

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.     

The Mechanisms of Coexistence and Competitive Exclusion in Complex Plankton Ecosystem Models

The biodiversity of plankton ecosystems may no longer be a paradox, but the mechanisms that determine coexistence of explicit competitors in ecosystems remain a mystery. This is particularly so in ecosystem models, where competitive exclusion remains the dominant process. Climate and fisheries models require plankton ecosystem sub-models that maintain competing plankton functional types extant, but coexistence can be reproduced in only a few ‘just so’ theoretical models. This limits our ability to predict the impacts of climate change and fisheries on ocean biota. We consider ecosystems of Kolmogorov form that conserve mass (CK systems). These systems describe a general class of ecosystem models that includes many theoretical and applied models. We develop heuristics that illuminate the key mechanisms that allow the coexistence of explicit competitors in these systems. These heuristics facilitate the identification of a large class of models with the structural property that all species coexist for all time. Our approach unifies many theoretical and applied models in a common biogeochemical framework, providing a powerful tool with the potential to generate new insights into the properties of complex ecosystems.     

Community assembly in temperate forest birds: habitat filtering,interspecific interactions and priority effects

Recent attempts to understand the processes governing community assembly have increasingly focused on patterns of phylogenetic relatedness and functional similarity among co-existing species. Considerations of the species pool, the number and identity of functional traits and the metrics used to identify patterns have come under scrutiny as possible influences on the detection of non-random patterns. Most mechanistic explanations of community assembly based on functional and phylogenetic approaches rely on deterministic explanations, while ignoring the role of stochastic processes and historical contingency, despite the prominent historical role of both types of explanations of species coexistence. We evaluated the phylogenetic and functional structure of 20 temperate forest bird assemblages in northeastern North America. We compared three approaches for characterizing the functional structure of assemblages. Regardless of approach, assemblages were generally no different than expected by chance. In contrast, phylogenetic structures of bird assemblages were overdispersed, clumped or consistent with random assembly depending on the site. Nonetheless, we found little evidence for differences in phylogenetic structure arising as a consequence of the identity of the species pool. We identified a strong relationship between the proportion of residents and phylogenetic relatedness that was unrelated to the species richness of assemblages. Our results suggest that different assembly mechanisms may structure resident and migratory subsets of temperate breeding bird communities. Resident assemblages are likely structured by interspecific interactions and habitat filtering prior to arrival of migrants. In contrast, the composition of migrant assemblages may be a consequence of priority effects in which the presence and abundance of residents and earliest arriving species affect the ability of subsequent migrants to colonize sites. This phenomenon enhances the likelihood of multiple alternative community structures in similar environments.     

Disappearing refuges in time and space: how environmental change threatens species coexistence

Understanding the impacts of environmental changes on species survival is a major challenge in ecological research, especially when shifting from single- to multispecies foci. Here, we apply a spatially explicit two-species simulation model to analyze the effects of geographic range shifting and habitat isolation on different coexistence mechanisms. The model explicitly considers dispersal, local competition, and growth on a single resource. Results highlight that both range shifting and habitat isolation severely impact coexistence. However, the strength of these impacts depends on the underlying coexistence mechanisms. Neutrally coexisting species are particularly sensitive to habitat isolation, while stabilized coexistence through overcompensatory density regulation is much more sensitive to range shifting. We conclude that, at the community level, the response to environmental change sensitively depends on the underlying coexistence mechanisms. This suggests that predictions and management recommendations should consider differences between neutral versus stabilized community structures whenever possible.