Asymmetric facilitation can reduce size inequality in plant populations resulting in delayed density‐dependent mortality

Size inequality in plant populations is a ubiquitous feature that has received much attention due to ecological and evolutionary implications. The mechanisms driving size inequality were mainly attributed to different modes of competition (symmetric versus asymmetric), while the potential effects of different modes of facilitation (symmetric versus asymmetric) to this pattern have not yet been fully explored. We employed an individual‐based model to explore the relative roles of both competition and facilitation simultaneously along an environmental stress gradient. Special emphasis was given to the assessment of symmetric facilitation (plants receive benefit from each other equally or proportionally to benefactors’ sizes) and asymmetric facilitation (beneficiary plants receive benefits from benefactor plants that are higher than proportional to the benefactors’ size) in altering plant size inequality. We found that independent of the particular mode of competition, symmetric facilitation generally increased size inequality, whereas asymmetric facilitation decreased it. This pattern was consistent along the stress gradient. Because of their different effects on size inequality, symmetric facilitation accelerated self‐thinning, whereas asymmetric facilitation delayed the onset of density‐dependent mortality, promoting survival under intermediate stress conditions. We compared our model predictions with both 1) a previous modelling study focusing on the effect of (symmetric) facilitation on the size inequality, and 2) re‐analysed data from a published experiment generating asymmetric facilitation of plants against enhanced ultraviolet‐B (UV‐B). Whereas our model predictions and the results of the empirical experiment were consistent, we found that previous theoretical results that solely relied on symmetric facilitation need to be re‐adjusted. Our study showed that combinations of different modes of competition and facilitation can alter size inequality in different ways and with important consequences for the onset of density‐dependent mortality during population development. Explicitly considering different modes and mechanisms of interactions (both facilitation and competition) will improve mechanistic understanding in plant ecology.     

Effects of Competition and Facilitation on Species Assemblage in Two Types of Tropical Cloud Forest

Competition and facilitation between tree individuals are two kinds of non-random processes influencing the structure and functioning of forest communities, but how these two plant-plant interactions change along gradient of resources or environments remains very much a matter of debate. We developed a null model to test the size-distance regression, and assessed the effects of competition and facilitation (including interspecific interactions, intraspecific interactions and overall species interactions) on each adult tree species assemblage [diameter at breast height (dbh) ≥5 cm] across two types of tropical cloud forest with different environmental and resource regimes. The null model test revealed that 17% to 27% tree species had positive dbh-distance correlations while 11% to 19% tree species showed negative dbh-distance correlations within these two forest types, indicating that both competition and facilitation processes existed during the community assembly. The importance of competition for heterospecific species, and the intensity of competition for both heterospecific and overall species increased from high to low resources for all the shared species spanning the two forests. The importance of facilitation for conspecific and overall species, as well as that the intensity of facilitation for both heterospecific and conspecific species increased with increasing low air temperature stress for all the shared species spanning the two forests. Our results show that both competition and facilitation processes simultaneously affect parts of species assemblage in the tropical cloud forests. Moreover, the fact that nearly 50% species assemblage is not detected with our approaches suggest that tree species in these tropical forest systems are assembled with multiple ecological processes, and that there is a need to explore the processes other than the two biotic interactions in further researches.     

Facilitation and competition interact with seed dormancy to affect population dynamics in annual plants

Seed dormancy increases population size via bet-hedging and by limiting negative interactions (e.g., competition) among individuals. On the other hand, individuals also interact positively (e.g., facilitation), and in some systems, facilitation among juveniles precedes competition among adults in the same generation. Nevertheless, studies of the benefits of seed dormancy typically ignore facilitation. Using a population growth model, we ask how the facilitation–competition balance interacts with seed dormancy rate to affect population dynamics in constant and variable environments. Facilitation increases the growth rate and equilibrium size (in both constant and variable environments) and reduces the extinction rate of populations (in a variable environment), and a higher rate of seed dormancy allows populations with facilitation to reach larger sizes. However, the combined benefits of facilitation and a high dormancy rate only occur in large populations. In small populations, weak facilitation does not affect the growth rate, but does induce a weak demographic Allee effect (where population growth decreases with decreasing population size). Our results suggest that facilitation within populations can interact with bet-hedging traits (such as dormancy) or other traits that mediate density to affect population dynamics. Further, by ensuring survival but limiting reproduction, ontogenetic switches from facilitation to competition may enable populations to persist but limit their maximum size in variable environments. Such intrinsic regulation of populations could then contribute to the maintenance of similar species within communities.     

Neighbours matter: natural selection on plant size depends on the identity and diversity of the surrounding community

Plant diversity can affect ecological processes such as competition and herbivory, and these ecological processes can act as drivers of evolutionary change. However, surprisingly little is known about how ecological variation in plant diversity can alter selective regimes on members of the community. Here, we examine how plant diversity at two different scales (genotypic and species diversity) impacts natural selection on a focal plant species, the common evening primrose (Oenothera biennis). Because competition is frequently relaxed in both genotypically and species rich plant communities, we hypothesized that increasing diversity would weaken selection on competitive ability. Changes in plant diversity can also affect associated arthropod communities. Therefore, we hypothesized that diversity would alter selection on plant traits mediating these interactions, such as herbivory related traits. We grew 24 focal O. biennis genotypes within four different neighbourhoods: genotypic monocultures or polycultures of O. biennis, and species monocultures or polycultures of old-field species that commonly co-occur with O. biennis. We then measured genotypic selection on nine plant traits known to be ecologically important for competition and herbivory. Focal O. biennis plants were smaller, flowered for shorter periods of time, had lower fitness, and experienced greater attack from specialist predispersal seed predators when grown with conspecifics versus heterospecifics. While neither conspecific nor heterospecific diversity altered trait means, both types of diversity altered the strength of selection on focal O. biennis plants. Specifically, selection on plant biomass was stronger in conspecific monocultures versus polycultures, but weaker in heterospecific monocultures versus polycultures. We found no evidence of selection on plant traits that mediate insect interactions, despite differences in arthropod communities on plants surrounded by conspecifics versus heterospecifics. Our data demonstrate that plant genotypic and species diversity can act as agents of natural selection, potentially driving evolutionary changes in plant communities.     

Pollination outcomes reveal negative density‐dependence coupled with interspecific facilitation among plants

Pollination is thought to be under positive density‐dependence, destabilising plant coexistence by conferring fitness disadvantages to rare species. Such disadvantage is exacerbated by interspecific competition but can be mitigated by facilitation and intraspecific competition. However, pollinator scarcity should enhance intraspecific plant competition and impose disadvantage on common over rare species (negative density‐dependence, NDD). We assessed pollination proxies (visitation rate, pollen receipt, pollen tubes) in a generalised plant community and related them to conspecific and heterospecific density, expecting NDD and interspecific facilitation due to the natural pollinator scarcity. Contrary to usual expectations, all proxies indicated strong intraspecific competition for common plants. Moreover interspecific facilitation prevailed and was stronger for rare than for common plants. Both NDD and interspecific facilitation were modulated by specialisation, floral display and pollinator group. The combination of intraspecific competition and interspecific facilitation fosters plant coexistence, suggesting that pollination can be a niche axis maintaining plant diversity.     

When can two plant species facilitate each other's pollination?

Facilitation occurs when an increase in the density of one species causes an increase in the population growth rate or the density of a second species. In plants, ample evidence demonstrates that one species can facilitate another by ameliorating abiotic conditions, but the hypothesis that pollination facilitation – in which the presence of one flowering species increases pollinator visits to a second species – can also occur remains controversial. To identify the necessary conditions for pollination facilitation to occur, we constructed population models of two plant species that share the same pollinator and compete for establishment sites, and we assumed that heterospecific pollen can interfere with successful seed set. We found that facilitation for pollination occurs only when the pollinator visitation rate is an initially accelerating function of the combined numbers of flowering plants of both species in a patch. The presence of a second species can allow populations of a focal species either to persist for a longer amount of time before going extinct ("weak facilitation") or to persist indefinitely at a stable equilibrium density ("strong facilitation"). When only a single plant of either species can occupy a site, the plant species with the higher initial density can experience strong facilitation but will eventually out-compete the other species. However, when site occupancy was not exclusive, strong facilitation sometimes led to coexistence of the two species. Increasing the extent of pollen carryover increased the range of initial population densities leading to strong facilitation. In light of our theoretical results, we discuss the apparent rarity of pollination facilitation in nature.     

Forest tree neighborhoods are structured more by negative conspecific density dependence than by interactions among closely related species

Interactions among neighbors influence the structure of communities of sessile organisms. Closely related species tend to share habitat and resource requirements and to interact with the same mutualists and natural enemies so that the strength of interspecific interactions tends to decrease with evolutionary divergence time. Nevertheless, the degree to which such phylogenetically related ecological interactions structure plant communities remains unclear. Using data from five large mapped forest plots combined with a DNA barcode mega‐phylogeny, we employed an individual‐based approach to assess the collective effects of focal tree size on neighborhood phylogenetic relatedness. Abundance‐weighted average divergence time for all neighbors (ADT_all) and for heterospecific neighbors only (ADT_hetero) were calculated for each individual of canopy tree species. Within local neighborhoods, we found phylogenetic composition changed with focal tree size. Specifically, significant increases in ADT_all with focal tree size were evident at all sites. In contrast, there was no significant change in ADT_hetero with tree size in four of the five sites for both sapling‐sized and all neighbors, even at the smallest neighbourhood scale (0–5 m), suggesting a limited role for phylogeny‐dependent interactions. However, there were inverse relationships between focal tree size and the proportion of heterospecific neighbors belonging to closely related species at some sites, providing evidence for negative phylogenetic density dependence. Overall, our results indicate that negative interaction with conspecifics had a much greater impact on neighborhood assemblages than interactions among closely related species and could contribute to community structure and diversity maintenance in different forest communities.     

Facilitation among plants can accelerate density‐dependent mortality and steepen self‐thinning lines in stressful environments

The speed and slope of plant self‐thinning are all affected by plant–plant interactions across environmental gradients. Possible mechanisms driving the self‐thinning dynamics include the relative strength of root versus shoot competition, and the interplay between competition and facilitation. Although these mechanisms often act in concert, their relative importance has not yet been fully explored. We used both a one‐layer and a two‐layer zone‐of‐influence (ZOI) model to examine how competition and facilitation drive self‐thinning across stress gradients. As a development of the traditional ZOI model, the two‐layer version explicitly models shoot and root growth and neighbor interactions, and thus the overall size‐symmetry of competition is regulated by the relative strength of root versus shoot competition. One‐layer model simulations revealed that increasingly asymmetric competition accelerated thinning, and steepened (slope ranged from about –1 to –4/3) and lowered self‐thinning lines. Stress slowed down density‐dependent mortality considerably when competition was not completely symmetric. Stress significantly decreased the self‐thinning intercept, while facilitation simply counteracted stress effects. Both stress and facilitation showed little effect on the slope. In the two‐layer model, both stress and facilitation affected mortality in the same way as in the one‐layer version when competition was not completely symmetric. Different from the one‐layer model, the two‐layer version showed that the effects of stress and facilitation on the self‐thinning slope were mediated by the asymmetry of competition. As stress increased, the overall asymmetry of competition shifted from asymmetric to symmetric due to increased relative strength of root competition. High stress thus dramatically flattened self‐thinning lines, whereas the inclusion of facilitation counteracted stress and led to steeper self‐thinning lines. Our two‐layer model is based on the current knowledge of plant–plant interactions, and better represents ecological realities. It can help elaborate experiments for testing the role of competition and facilitation in driving plant population dynamics.     

LOCAL ADAPTATION WHEN COMPETITION DEPENDS ON PHENOTYPIC SIMILARITY

Recent work incorporating demographic–genetic interactions indicates the importance of population size, gene flow, and selection in influencing local adaptation. This work typically assumes that density‐dependent survival affects individuals equally, but individuals in natural population rarely compete equally. Among‐individual differences in resource use generate stronger competition between more similar phenotypes (frequency‐dependent competition) but it remains unclear how this additional form of selection changes the interactions between population size, gene flow, and local stabilizing selection. Here, we integrate migration–selection dynamics with frequency‐dependent competition. We developed a coupled demographic‐quantitative genetic model consisting of two patches connected by dispersal and subject to local stabilizing selection and competition. Our model shows that frequency‐dependent competition slightly increases local adaptation, greatly increases genetic variance within patches, and reduces the amount that migration depresses population size, despite the increased genetic variance load. The effects of frequency‐dependence depend on the strength of divergent selection, trait heritability, and when mortality occurs in the life cycle in relation to migration and reproduction. Essentially, frequency‐dependent competition reduces the density‐dependent interactions between migrants and residents, the extent to which depends on how different and common immigrants are compared to residents. Our results add new dynamics that illustrate how competition can alter the effects of gene flow and divergent selection on local adaptation and population carrying capacities.     

Local interactions drive size dependent space competition between coral and crustose coralline algae

For many species, the outcome of competition for space in homogeneous habitats depends upon relative rates of growth and overgrowth. Size dependence in competition occurs when this balance shifts due to the growth of one or both species. For example, the ability of coral to compete with certain species of crustose coralline algae (CCA) may depend on whether coral colonies are large enough to avoid being overgrown. Spatially implicit models suggest size refuges from competition can improve the persistence of species with a vulnerable life stage. We use spatially explicit simulation models to explore size dependence in competition between coral and competitively dominant CCA in well lit habitat. We determine what conditions allow coral to use size refuges and whether refuges improve the recovery of coral after disturbance. Local interactions in explicit space prevent the maturation of coral into size refuges unless coral grows more rapidly than CCA or coral colonies are allowed to fuse, and mortality mechanisms can limit long‐term persistence even if the refuge is achieved. We contrast results with analogous differential equation models, with and without an explicit maturation delay, to demonstrate how the predicted outcome of competition is frequently reversed when local interactions and individual‐based dynamics are included in models of size‐dependent competition.     

Evaluating the effects of pollinator‐mediated interactions using pollen transfer networks: evidence of widespread facilitation in south Andean plant communities

Information about the relative importance of competitive or facilitative pollinator‐mediated interactions in a multi‐species context is limited. We studied interspecific pollen transfer (IPT) networks to evaluate quantity and quality effects of pollinator sharing among plant species on three high‐Andean communities at 1600, 1800 and 2000 m a.s.l. To estimate the sign of the effects (positive, neutral or negative), the relation between conspecific and heterospecific pollen deposited on stigmas was analysed with GLMMs. Network analyses showed that communities were characterised by the presence of pollen hub‐donors and receptors. We inferred that facilitative and neutral pollinator‐mediated interactions among plants prevailed over competition. Thus, the benefits from pollinator sharing seem to outweigh the costs (i.e. heterospecific deposition and conspecific pollen loss). The largest proportion of facilitated species was found at the highest elevation community, suggesting that under unfavourable conditions for the pollination service and at lower plant densities facilitation can be more common.     

Physiologically structured models – from versatile technique to ecological theory

A ubiquitous feature of natural communities is the variation in size that can be observed between organisms, a variation that to a substantial degree is intraspecific. Size variation within species by necessity implies that ecological interactions vary both in intensity and type over the life cycle of an individual. Physiologically structured population models (PSPMs) constitute a modelling approach especially designed to analyse these size‐dependent interactions as they explicitly link individual level processes such as consumption and growth to population dynamics. We discuss two cases where PSPMs have been used to analyse the dynamics of size‐structured populations. In the first case, a model of a size‐structured consumer population feeding on a non‐structured prey was successful in predicting both qualitative (mechanisms) and quantitative (individual growth, survival, cycle amplitude) aspects of the population dynamics of a planktivorous fish population. We conclude that single generation cycles as a result of intercohort competition is a general outcome of size‐structured consumer–resource interactions. In the second case, involving both cohort competition and cannibalism, we show that PSPMs may predict double asymptotic growth trajectories with individuals ending up as giants. These growth trajectories, which have also been observed in field data, could not be predicted from individual level information, but are emergent properties of the population feedback on individual processes. In contrast to the size‐structured consumer–resource model, the dynamics in this case cannot be reduced to simpler lumped stage‐based models, but can only be analysed within the domain of PSPMs. Parameter values used in PSPMs adhere to the individual level and are derived independently from the system at focus, whereas model predictions involve both population level processes and individual level processes under conditions of population feedback. This leads to an increased ability to test model predictions but also to a larger set of variables that is predicted at both the individual and population level. The results turn out to be relatively robust to specific model assumptions and thus render a higher degree of generality than purely individual‐based models. At the same time, PSPMs offer a much higher degree of realism, precision and testing ability than lumped stage‐based or non‐structured models. The results of our analyses so far suggest that also in more complex species configurations only a limited set of mechanisms determines the dynamics of PSPMs. We therefore conclude that there is a high potential for developing an individual‐based, size‐dependent community theory using PSPMs.     

Signs of stabilisation and stable coexistence

Many empirical studies motivated by an interest in stable coexistence have quantified negative density dependence, negative frequency dependence, or negative plant–soil feedback, but the links between these empirical results and ecological theory are not straightforward. Here, we relate these analyses to theoretical conditions for stabilisation and stable coexistence in classical competition models. By stabilisation, we mean an excess of intraspecific competition relative to interspecific competition that inherently slows or even prevents competitive exclusion. We show that most, though not all, tests demonstrating negative density dependence, negative frequency dependence, and negative plant–soil feedback constitute sufficient conditions for stabilisation of two‐species interactions if applied to data for per capita population growth rates of pairs of species, but none are necessary or sufficient conditions for stable coexistence of two species. Potential inferences are even more limited when communities involve more than two species, and when performance is measured at a single life stage or vital rate. We then discuss two approaches that enable stronger tests for stable coexistence‐invasibility experiments and model parameterisation. The model parameterisation approach can be applied to typical density‐dependence, frequency‐dependence, and plant–soil feedback data sets, and generally enables better links with mechanisms and greater insights, as demonstrated by recent studies.     

Relationship between size hierarchy and density of trees in a tropical dry deciduous forest of western India

Questions: Density dependence is thought to restrict exponential growth as well as give rise to size structure in populations. Size hierarchy in trees from tropical dry deciduous forests is studied to ask (1) whether nature of competition is symmetric or asymmetric and (2) what is the self thinning trajectory under a natural gradient of tree density. Location: Western India. Methods: Density was measured as the number of trees in 10‐m radius circular plots (n= 96) and size was measured at DBH. Size variation was evaluated by the Gini coefficient (n= 1239 trees). Results: Size inequality between neighbours decreased with density but in a non‐linear manner. In the backdrop of existing theory this indirectly suggests that competitive interactions may be symmetric over a ‘depletive’ resource such as below‐ground water (rather than a ‘pre‐emptive’ resource such as light), which is very plausible in a semi‐arid environment. The self thinning coefficient derived from the relationship between stem diameter and density (γ～?1/4), is higher than expected from existing models of allometric plant growth (γ=?1/3) which are based on above‐ground interactions alone. Seen in conjunction, these results suggest that above‐ground structures, such as stem size, do not adequately represent the outcome of competitive interactions when below‐ground resources, such as water, may be more important under semi‐arid conditions. Conclusions: The non‐linear relationship between size inequality and density indicates that there exists a density threshold beyond which investment in above‐ground biomass becomes sluggish in semi‐arid, deciduous forests. Since current allometric models do not incorporate below‐ground biomass for trees, these data suggest that a more comprehensive allometric growth model may have higher predictive power and wider applicability.     

Eavesdropping in solitary large carnivores: Black bears advance and vocalize toward cougar playbacks

Large carnivore behavioral responses to the cues of their competitors are rarely observed, but may mediate competition between these top predators. Playback experiments, currently limited to interactions involving group‐living large carnivores, demonstrate that attending to cues indicative of the immediate presence of heterospecific competitors plays a substantial role in influencing competition among these species. Group‐living species vocalize regularly to signal to one another, and competitors can readily “eavesdrop” on these acoustic cues. Solitary large carnivores also vocalize to conspecifics, but much less frequently, reducing the ease with which heterospecific competitors can eavesdrop. Eavesdropping could nonetheless play a substantive role in mediating competition among solitary large carnivores if the benefits of responding to the acoustic cues of heterospecific competitors (reducing risk or locating resources) are sufficiently large. Behavioral interactions between solitary large carnivore species are almost never observed, and there have been no experimental tests of their reactions to cues indicative of the immediate presence of other solitary large carnivores. We used an automated playback system to test the responses of a solitary large carnivore (black bear, Ursus americanus) to vocalizations of their similarly solitary competitor (cougar, Puma concolor), presenting both cougar and control vocalizations to free‐living bears foraging along shorelines in British Columbia, Canada. Both mothers with cubs and solitary bears were significantly more likely to advance and vocalize toward cougar than control playbacks, mothers producing one or both of two distinct vocalizations and solitary bears producing just one. Cougars could either represent a potential risk to bears (particularly cubs), or a source of resources, as bears are known to regularly scavenge cougar kills. Our results are consistent with bears eavesdropping on cougars for both these reasons. As with group‐living species, eavesdropping may be common among solitary large carnivores, and may be an important driver of competition between these species.     

From facilitation to competition: temperature‐driven shift in dominant plant interactions affects population dynamics in seminatural grasslands

Biotic interactions are often ignored in assessments of climate change impacts. However, climate‐related changes in species interactions, often mediated through increased dominance of certain species or functional groups, may have important implications for how species respond to climate warming and altered precipitation patterns. We examined how a dominant plant functional group affected the population dynamics of four co‐occurring forb species by experimentally removing graminoids in seminatural grasslands. Specifically, we explored how the interaction between dominants and subordinates varied with climate by replicating the removal experiment across a climate grid consisting of 12 field sites spanning broad‐scale temperature and precipitation gradients in southern Norway. Biotic interactions affected population growth rates of all study species, and the net outcome of interactions between dominants and subordinates switched from facilitation to competition with increasing temperature along the temperature gradient. The impacts of competitive interactions on subordinates in the warmer sites could primarily be attributed to reduced plant survival. Whereas the response to dominant removal varied with temperature, there was no overall effect of precipitation on the balance between competition and facilitation. Our findings suggest that global warming may increase the relative importance of competitive interactions in seminatural grasslands across a wide range of precipitation levels, thereby favouring highly competitive dominant species over subordinate species. As a result, seminatural grasslands may become increasingly dependent on disturbance (i.e. traditional management such as grazing and mowing) to maintain viable populations of subordinate species and thereby biodiversity under future climates. Our study highlights the importance of population‐level studies replicated under different climatic conditions for understanding the underlying mechanisms of climate change impacts on plants.     

Population dynamics of three songbird species in a nestbox population in Central Europe show effects of density,climate and competitive interactions

Unravelling the contributions of density‐dependent and density‐independent factors in determining species population dynamics is a challenge, especially if the two factors interact. One approach is to apply stochastic population models to long‐term data, yet few studies have included interactions between density‐dependent and density‐independent factors, or explored more than one type of stochastic population model. However, both are important because model choice critically affects inference on population dynamics and stability. Here, we used a multiple models approach and applied log‐linear and non‐linear stochastic population models to time series (spanning 29 years) on the population growth rates of Blue Tits Cyanistes caeruleus, Great Tits Parus major and Pied Flycatchers Ficedula hypoleuca breeding in two nestbox populations in southern Germany. We focused on the roles of climate conditions and intra‐ and interspecific competition in determining population growth rates. Density dependence was evident in all populations. For Blue Tits in one population and for Great Tits in both populations, addition of a density‐independent factor improved model fit. At one location, Blue Tit population growth rate increased following warmer winters, whereas Great Tit population growth rates decreased following warmer springs. Importantly, Great Tit population growth rate also decreased following years of high Blue Tit abundance, but not vice versa. This finding is consistent with asymmetric interspecific competition and implies that competition could carry over to influence population dynamics. At the other location, Great Tit population growth rate decreased following years of high Pied Flycatcher abundance but only when Great Tit population numbers were low, illustrating that the roles of density‐dependent and density‐independent factors are not necessarily mutually exclusive. The dynamics of this Great Tit population, in contrast to the other populations, were unstable and chaotic, raising the question of whether interactions between density‐dependent and density‐independent factors play a role in determining the (in) stability of the dynamics of species populations.     

Competition,facilitation and the Allee effect

Individuals of different species may interact in many different ways, such as competition, mutualism, or predation, to name but a few. Recent theory and experiments reveal that whether an interaction is beneficial or detrimental to the dynamics of a population often depends on species densities and other environmental factors. Here, we explore how, for suitable densities, facilitation may arise between two competing species with an Allee effect. We consider two different mechanisms for the Allee effect: 1) plant species with obligate insect pollination, and 2) generalist predation. In the first case, a second plant species, competing for nutrients, may have a facilitative effect by attracting more pollinators. In the second case, another potentially competing species may serve to satiate the same generalist predator and thereby have a facilitative effect. We explore three aspects of facilitation in each of the two systems. The focal species may benefit from the presence of a ‘competitor’ if it experiences 1) the removal of the Allee threshold, 2) a lowering of the Allee threshold, or 3) an increase in carrying capacity. We find that the latter two effects occur in both study systems whereas the first only occurs for the generalist predation system but not for the plant‐pollination system. We give precise conditions on when such a facilitative effect can be expected. We also demonstrate several unexpected outcomes of these two‐species interactions with multiple steady states, such as obligate co‐occurence; we draw parallels to the dynamics of species known as ‘ecosystem engineers’, and we discuss implications for conservation and management.     

Effects of altered resource consumption rates by one consumer species on a competitor

Non‐mechanistic models of competition suggest that harming one of two competing species will increase the population density of the other. These models also suggest that any change in a fitness component of one competitor will make the densities of the two competitors change in opposite directions. However, models of competition that incorporate resource dynamics show that neither conclusion holds generally. Reducing the consumption abilities of one competitor may decrease the population size of the other by decreasing resource overexploitation by the first and thereby increasing its density. It is also possible for decreased consumption abilities of one species to increase the population densities of both species, when the increased density of the focal species is offset by its decreased ability to consume the main resources of its competitor. Finally, decreases in consumption may have the effects predicted by phenomenological models; a decrease in the focal species and an increase in its competitor. Unstable systems may exhibit more complicated patterns of changes in densities with changes in consumption rates. These counterintuitive effects depend on the presence of overexploitation of biotic resources, about which little is known. More generally, there have been few theoretical or empirical studies examining the indirect effects of changes in consumption rates of a focal species in a food web; these are termed ‘trait‐initiated indirect effects’. A better understanding of the potential consequences of altered consumption rates will be important for understanding biotic shifts in communities undergoing environmental change, and in using simple community modules to understand larger food webs.     

Density Effects on the Size Structure of Annual Plant Populations: An Indication of Neighbourhood Competition

Size variability among plants has been observed to increasewith higher stand density, leading to the speculation that resourcedistribution among competing plants is primarily asymmetricrather than symmetric. The relationships between size variability,stand density, and type of resource distribution among competingplants were investigated using a spatially explicit, individual-plantmodel of annual plant population dynamics. When plants variedin neighbourhood competition, size variability increased withhigher stand densities whether shared resources were symmetricallyor asymmetrically distributed among competing plants. Size variabilitydid not increase with higher stand densities when neighbourhoodcompetition was constant for all plants. These simulations indicatethat increased size variability among competing plants doesnot distinguish between symmetric and asymmetric resource distribution,but rather is direct evidence for neighbourhood competition. Size hierarchy, neighbourhood competition, density effects, asymmetric competition, symmetric competition