The distribution of positive and negative species interactions across environmental gradients on a dual-lattice model

There has been considerable recent interest in understanding the role of positive inter-specific interactions within ecology, and significant progress has been made both empirically and theoretically. Similarly, considerable progress has been made in improving our understanding of the mechanisms that limit species' ranges. In this contribution, we seek to understand the setting of species' borders when some species within the assemblage exhibit positive inter-specific interactions. We use a spatially explicit dual-lattice simulation model to explore the distribution of different interactions across environmental gradients. We first simulate community dynamics when there is either a gradient in reproductive rate or in mortality. We then consider what happens when gradients in reproduction and mortality run in parallel or perpendicular to one another. If the stress gradient impacts on reproductive potential, positive interactions are found where there is high abiotic stress. In this instance, the mutualists are able to tolerate an environment that the cheaters cannot. However, when the stress gradient influences mortality, we find that the mutualists occur as a stripe surrounded by cheaters both towards the better and the harsher ends of the gradient. Previous theory and most empirical evidence tend to indicate that net positive interactions are likely to occur in environments characterized by high abiotic stress. However, evidence from some stress gradients suggests that the distribution of positive and negative interactions can be more complex, with the most stressful environments being occupied by individuals engaging in negative rather than positive interactions. Our results provide a potential theoretical explanation for these recent field observation, and highlight the need for further theoretical and empirical work to better our understanding of how positive and negative interactions act to determine the limits to species' ranges.     

Florivores decrease pollinator visitation in a self-incompatible plant

Different biotic interactions may influence one another to produce complex patterns of direct and indirect effects, which together influence plant reproductive success. However, so far most studies on plant-animal interactions have focused on single interactions in isolation. In this study, we studied the effect of florivory by the weevil Cionus nigritarsis on pollinator visitation rate in the self-incompatible perennial herb Verbascum nigrum by combining observations of florivory and pollination in natural populations with records of pollinator visitation to plants with different levels of experimentally inflicted damage.Increasing levels of damage through either natural or simulated florivory resulted in fewer pollinator visits per plant and per flower. As expected, the magnitude of the indirect effect of florivory on pollinator visitation was proportional to the intensity of florivory. Our results indicate that biotic non-pollinating agents, such as florivores, may induce substantial changes in pollinator availability. Therefore, studies addressing different plant-animal interactions in parallel are necessary to better comprehend the factors influencing the reproductive performance and demography of flowering plants.     

Species removal and experimental warming in a subarctic tundra plant community

Neighbor interactions are likely to play an important role in subarctic plant communities. We conducted experiments in Interior Alaska in which we crossed species removal with greenhouse warming manipulations. We examined changes in community biomass, and in plant survival and growth of individual species in response to experimental warming and to: (1) removal of whole species versus an equivalent amount of biomass across many species, and (2) removal of subdominant (locally common) versus minor (locally uncommon) plants. Community biomass indicated compensation in growth after removal of minor species and after biomass removal without elimination of entire species, but under-compensation after removal of subdominants. Growth and survival of individual species showed facilitation between some species. Warming increased growth of dominant vascular plants, but at the same time reduced survival, and these impacts were greater for larger, more mesic species than for the smaller species associated with drier habitats. Growth of mosses was reduced by the warming. Removal effects did not differ between warming and ambient conditions. The results indicate that common species are able to reduce resources for others (competitive effect) and increase their growth after neighbor removal, whereas locally uncommon species are not able to respond rapidly to increased resources made available by neighbor removal. Therefore, the impact of the presence of common species on locally uncommon species was facilitative overall, but not vice versa. The balance between disturbances such as changes in temperature and species losses from the community will likely be crucial in determining shifts in subsequent community composition.     

Effects of floral traits and plant genetic composition on pollinator behavior

Pollinator-mediated selection plays a major role in floral evolution and speciation. Floral traits that influence animal pollinator behavior are the target of pollinator-mediated selection, but can only evolve if floral phenotypes have underlying genetic variation. Thus, understanding the genetic basis of a floral trait is a crucial step in studying pollinator-mediated selection. In this study I tested the effect of quantitative trait loci (QTL) underlying floral traits on pollinator behavior in recombinant inbred lines (RILs) in the common sunflower, Helianthus annuus L. and its crop relative. The indirect effects of QTL on pollinator behavior, mediated by floral phenotypes, were analyzed for six insect visitor types using structural equation modeling (SEM) and path analysis. For three of the six visitor types (large and small bees and non-bee insects) valid models were revealed when all three levels (QTL, floral traits, and pollinator behavior) were incorporated. Nested model without genetics were validated for five of the six visitor types. The results suggest that insect behavior as a reaction to floral phenotypes is affected by the genetic architecture of floral traits. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Heikki Hokkanen     

The role of reproductive plant traits and biotic interactions in the dynamics of semi-arid plant communities

The dynamics of semi-arid plant communities are determined by the interplay between competition and facilitation among plants. The sign and strength of these biotic interactions depend on plant traits. However, the relationships between plant traits and biotic interactions, and the consequences for plant communities are still poorly understood. Our objective here was to investigate, with a modelling approach, the role of plant reproductive traits on biotic interactions, and the consequences for processes such as plant succession and invasion. The dynamics of two plant types were modelled with a spatially-explicit integrodifferential model: (1) a plant with seed dispersal (colonizer of bare soil) and (2) a plant with local vegetative propagation (local competitor). Both plant types were involved in facilitation due to a local positive feedback between vegetation biomass and soil water availability, which promoted establishment and growth. Plants in the system also competed for limited water. The efficiency in water acquisition (dependent on reproductive and growth plant traits) determined which plant type dominated the community at the steady state. Facilitative interactions between plant types also played an important role in the community dynamics, promoting establishment in the driest conditions and recovery from low biomass. Plants with vegetative propagation took advantage of the ability of seed dispersers to establish on bare soil from a low initial biomass. Seed dispersers were good invaders, maintained high biomass at intermediate and high rainfall and showed a high ability in taking profit from the positive feedback originated by plants with vegetative propagation under the driest conditions. However, seed dispersers lost competitiveness with an increasing investment in fecundity. All together, our results showed that reproductive plant traits can affect the balance between facilitative and competitive interactions. Understanding this effect of plant traits on biotic interactions provides insights in processes such as plant succession and shrub encroachment.     

Convergence patterns and multiple species interactions in a designed plant mixture of five species

It is known that convergence and divergence can occur in complex plant communities, but the relative importance of biotic and abiotic factors driving these processes is less clear. We addressed this issue in an experiment using a range of mixed stands of five species that are common in Swiss fens (Carex elata, C. flava, Lycopus europaeus, Lysimachia vulgaris and Mentha aquatica) and two levels of water and nutrients. One hundred and seventy-six experimental mixtures were maintained in large pots (75 l) for two consecutive growing seasons in an experimental garden. The stands varied systematically in the initial relative abundance of each of the five species and in overall initial stand abundance. The changes in biomass over 2 years were modelled as linear functions of treatments and the initial biomass of each species. The dynamics of the system were mainly driven by differences in the identity of species and by a negative feedback mechanism but also by different abiotic conditions. In all mixtures, C. elata became more dominant over time, which caused an overall convergence of community composition. In addition, the rate of change of each species’ biomass was negatively related to its own initial abundance. Thus, a negative feedback further contributed to the convergence of communities. Species responded differently to water level and nutrient supply, causing community dynamics to differ among treatments. However, the different abiotic conditions only slightly modified the overall convergence pattern. Competitive interactions between more than two species were weaker than the negative feedback but still significantly influenced the species’ final relative abundance. The negative feedback suggests that there is niche partitioning between the species, which permits their coexistence. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Recovery of a northern New England salt marsh plant community from winter icing

High latitude salt marsh plant communities are frequently exposed to conspicuous winter ice disturbances, which trigger secondary succession. In this paper, we document the recovery of a northern New England salt marsh from a severe winter icing event in 1998. Ice disturbances that killed plants but that left the underlying peat intact recovered rapidly. However, ice damage that killed plants and removed the underlying peat, led to areas of physiologically harsh edaphic conditions, specifically waterlogged and anoxic soils that limited plant recolonization. A transplant experiment revealed that only the most stress-tolerant plants were capable of invading the most stressful portions of ice disturbances. A second experiment that artificially dried disturbance patches accelerated patch recovery. These data suggest that recovery from intense ice disturbance is dependent on stress-tolerant plants invading edaphically harsh disturbances, eventually facilitating the recolonization of the community. This process likely takes longer than a decade for full recovery to occur in the areas where both plants and the peat base are removed.     

Heterospecific attraction and food resources in migrants' breeding patch selection in northern boreal forest

We studied experimentally how heterospecific attraction may affect habitat selection of migrant passerine birds in Finnish Lapland. We manipulated the densities of resident tit species (Parus spp.). In four study plots residents were removed before the arrival of the migrants in the first study year, and in four other plots their densities were increased by releasing caught individuals. In the second year the treatments of the areas were reversed, allowing paired comparisons within each plot. We also investigated the relative abundance of arthropods in the study plots by the sweep-net method. This allowed us to estimate the effect of food resources on the abundance of birds. The heterospecific attraction hypothesis predicts that densities of migrant species (especially habitat generalists) would be higher during increased resident density. Results supported this prediction. Densities and number of the most abundant migrant species were significantly higher when resident density was increased than when they were removed. On the species level the redwing (Turdus iliacus) showed the strongest positive response to the increased abundance of tits. Migrant bird abundances seemed not to vary in parallel with relative arthropod abundance, with the exception of the pied flycatcher (Ficedula hypoleuca) which showed a strongly positive correlation with many arthropod groups. The results of the experiment indicate that migrants can use resident tit species as a cue to a profitable breeding patch. The relationship between the abundance of the birds and arthropods suggests that annual changes in food resources during the breeding season probably do not have a very important effect on bird populations in these areas. The results stress the importance of positive interspecific interactions in structuring northern breeding bird communities. Received: 1 September 1997 / Accepted: 22 January 1998     

Nested species interactions promote feasibility over stability during the assembly of a pollinator community

The foundational concepts behind the persistence of ecological communities have been based on two ecological properties: dynamical stability and feasibility. The former is typically regarded as the capacity of a community to return to an original equilibrium state after a perturbation in species abundances and is usually linked to the strength of interspecific interactions. The latter is the capacity to sustain positive abundances on all its constituent species and is linked to both interspecific interactions and species demographic characteristics. Over the last 40 years, theoretical research in ecology has emphasized the search for conditions leading to the dynamical stability of ecological communities, while the conditions leading to feasibility have been overlooked. However, thus far, we have no evidence of whether species interactions are more conditioned by the community''s need to be stable or feasible. Here, we introduce novel quantitative methods and use empirical data to investigate the consequences of species interactions on the dynamical stability and feasibility of mutualistic communities. First, we demonstrate that the more nested the species interactions in a community are, the lower the mutualistic strength that the community can tolerate without losing dynamical stability. Second, we show that high feasibility in a community can be reached either with high mutualistic strength or with highly nested species interactions. Third, we find that during the assembly process of a seasonal pollinator community located at The Zackenberg Research Station (northeastern Greenland), a high feasibility is reached through the nested species interactions established between newcomer and resident species. Our findings imply that nested mutualistic communities promote feasibility over stability, which may suggest that the former can be key for community persistence.     

Effects of spatial pattern and relatedness in an experimental plant community

Many plant species show limited dispersal resulting in spatial and genetic substructures within populations. Consequently, neighbours are often related between each other, resulting in sibling competition. Using seed families of the annuals Capsella bursa-pastoris and Stachys annua we investigated effects of spatial pattern (i.e. random versus aggregated) on total and individual performance at the level of species and seed families under field conditions. At the level of species, we expected that inferior competitors increase, while superior competitors decrease their performance within neighbourhoods of conspecifics. Thus, we expected a species by spatial pattern interaction. Sibling competition, however, might reduce the performance of competitors, when genetically related, rather than non-related individuals are competing. Therefore, aggregations at the level of seed families could decrease the performance of competitors. Alternatively, if the opposite outcome would be observed, kin selection might be hypothesized to have occurred in the past. Because heavy seeds are expected to disperse less than light seeds, we further hypothesized that kin selection might be more likely to occur in superior competitors with heavy, locally dispersed seeds (e.g. Stachys) compared to inferior competitors with light, more distantly dispersed seeds (e.g. Capsella). We found a significant species by spatial pattern interaction. Indeed, the inferior competitor, Capsella, showed increased reproductive biomass production in aggregated compared to random patterns. Whereas, the performance of the superior competitor, Stachys, was to some extent decreased by intraspecific aggregation. Although statistically not significant, effects of intrafamily aggregations tended to be rather negative in Capsella but positive in Stachys. Our results confirmed that spatial patterns affect growth and reproduction of plant species promoting coexistence in plant communities. Although, we could not provide strong evidence for sibling competition or kin selection, our results suggested that competition among relatives was more severe for Capsella (lighter seeds) compared to Stachys (heavier seeds).     

Soil as a mediator in plant‐plant interactions in a semi‐arid community

Abstract. Competition and facilitation may occur simultaneously in plant communities, and the prevalence of either process depends on abiotic conditions. Here we attempt a community‐wide approach in the analysis of plant interactions, exploring whether in a semi‐arid environment positive or negative interactions predominate and whether there are differences among co‐occurring shrub species. Most shrubs in our plot exerted significant effects on their understorey communities, ranging from negative to positive. We found a clear case of interference and another case where the effect was neutral, but facilitation predominated and the biomass of annuals under most shrubs in our community was larger than in gaps. Effects on soil water and fertility were revealed as the primary source of facilitation; the build‐up of soil organic matter changed soil physical properties and improved soil water relations. Facilitation by shrubs involved decoupling of soil temperature and moisture. Sheltering from direct radiation had an effect on productivity, but significant differences in understorey biomass did not parallel understorey light environment. A positive balance of the interaction among plants, essentially mediated by changes in soil properties, is the predominant outcome of plant interactions in this semi‐arid community.     

Positive and negative effects of a dominant competitor on the settlement, growth, and survival of competing species in an epibenthic community

Because dominant competitors can monopolize resources, any positive effects they have on other species can have large community impacts. The solitary tunicate Ascidia ceratodes is a dominant competitor in the fouling community in Bodega Harbor, CA. This tunicate preempts primary substratum from competitors, but its thick tunic also allows other species to grow on its surface. The net effect of Ascidia on the community as a whole therefore depends on the balance between competitive and facilitative effects. In this study we evaluate the facilitative effects of Ascidia on different life stages of common competing species. We quantified larval settlement onto Ascidia compared to unoccupied space; we quantified the growth rate on Ascidia of small colonies of two common species; and we measured whether established colonies could escape overgrowth by Ascidia by growing onto its tunic. We found evidence for high rates of settlement on Ascidia, with some species showing higher and others lower settlement relative to that observed on free space. The growth rate of settlers was generally lower on Ascidia compared to primary substratum. We also found that colonial species established on primary space commonly escape overgrowth from Ascidia by growing onto Ascidia, as this occurs in about half of all encounters. This study indicates that the total effect of Ascidia on the community will depend on species-specific and life stage-specific effects, with likely long-term consequences for the diversity and composition of the community.     

Positive interactions, discontinuous transitions and species coexistence in plant communities

The population and community level consequences of positive interactions between plants remain poorly explored. In this study we incorporate positive resource-mediated interactions in classic resource competition theory and investigate the main consequences for plant population dynamics and species coexistence. We focus on plant communities for which water infiltration rates exhibit positive dependency on plant biomass and where plant responses can be improved by shading, particularly under water limiting conditions. We show that the effects of these two resource-mediated positive interactions are similar and additive. We predict that positive interactions shift the transition points between different species compositions along environmental gradients and that realized niche widths will expand or shrink. Furthermore, continuous transitions between different community compositions can become discontinuous and bistability or tristability can occur. Moreover, increased infiltration rates may give rise to a new potential coexistence mechanism that we call controlled facilitation.     

Little evidence for negative effects of an invasive alien plant on pollinator services

Many invasive alien plants occur in large populations with abundant flowers which are highly attractive to pollinators, and thus might affect pollination of co-occurring native species. This study focuses on the invasive Heracleum mantegazzianum and distance-dependent effects on pollination of Mimulus guttatus in abandoned grassland over 2 years. First, we examined pollinator abundance in yellow traps at 0, 10, 30 and 60–200 m from H. mantegazzianum. We then placed M. guttatus plants at the same distances to monitor effects of the invasive species on pollinator visitation and seed set of neighbouring plants. Finally, we conducted a garden experiment to test if deposition of H. mantegazzianum pollen reduces seed set in M. guttatus. No distance effect was found for the number of bumblebees in traps, although the invasive species attracted a diverse assemblage of insects, and visitation of M. guttatus was enhanced close to H. mantegazzianum. This positive effect was not reflected by seed set of M. guttatus, and heterospecific pollen decreased seed set in these plants. Overall there is little evidence for negative effects of the invasive species on pollination of neighbouring plants, and flower visitation even increases close to the invaded patches. The functional role of the invader and suitable control strategies need further clarification, since removal of H. mantegazzianum may actually damage local pollinator populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Impacts of simulated livestock grazing on Utah prairie dogs (Cynomys parvidens) in a low productivity ecosystem

Allometric foraging theory suggests that herbivores of greatly differing size should co-exist through niche segregation, but a few studies of large–small herbivore foraging relationships have reported competitive interactions. This study addresses the potential roles of habitat productivity and large herbivore grazing intensities on large–small herbivore foraging interactions. We examined effects of different intensity simulated grazing treatments on forage abundance and quality for Utah prairie dogs (Cynomys parvidens) in a low productivity ecosystem, and consequent effects on prairie dog individual growth rates, foraging preferences, and activity budgets. We hypothesized that simulated grazing would have predominantly facilitative impacts on Utah prairie dogs, as was found for black-tailed prairie dogs in higher productivity ecosystems. To test this hypothesis, we measured the effects of simulated grazing on forage nitrogen, digestibility, and biomass. Simulated grazing increased average forage nitrogen and digestibility while decreasing forage biomass. These effects were associated with reduced individual growth rates, increased juvenile foraging time, and reduced juvenile vigilance. Results suggest that the negative effects of reduced vegetation biomass greatly outweighed positive treatment effects in this study. However, prairie dogs in the moderate intensity defoliation treatment showed some preference for “grazed” plots over “ungrazed” plots, and this preference increased with time. Our study lends support to the idea that habitat productivity and herbivore densities may mediate shifts between facilitative and competitive interactions between different-sized herbivores.     

Resource limitation,habitat segregation,and species interactions of british tree-hole mosquitoes in nature

Summary The insect fauna of water-filled tree holes in southern Britain consists primarily of the mosquitoes Aedes geniculatus, Anopheles plumbeus, Culex torrentium, and a benthic detritivorous fauna that includes primarily the scirtid beetle Prionocyphon serricornis and the chironomid midge Metriocnemus martinii. Culex torrentium has been documented only relatively recently in tree holes but all three species of mosquitoes partition the resource in space and time. When mosquito larvae were forced to coexist in natural tree holes at limiting densities and at higher than natural levels of interspecific encounter, there was no evidence that Aedes geniculatus or Anopheles plumbeus affected pupation success, pupal weight, or development time of the other or that either Aedes geniculatus or C. torrentium affected the survivorship, pupation success, pupal weight, and biomass yield of the other. When A. geniculatus at limiting densities were forced in natural tree holes to live without or to coexist with natural or twice natural densities of P. serricornis and M. martinii, the presence, absence, or superabundance of the benthic insects did not affect pupation success or pupal weight of A. geniculatus; development time of A. geniculatus was faster when a superabundance of the benthic fauna was present. Effects of the benthic fauna on A. geniculatus are slight and the only significant interaction is facilitative, not competitive. The pattern of habitat segregation among treehole mosquitoes in southern Britain is characteristic of their respective genera and we propose that this pattern is more likely (but not certain) to have arisen through a process of independent evolution than through competitively driven niche shifts among already coexisting species.     

Positive vs. negative interactions in Picris hieracioides L., a mid-successional species of Mediterranean secondary succession

A field experiment was designed to evaluate the importance offacilitative and competitive interactions in Picrishieracioides, a facultative biennial that colonises the early andthemid-stages of secondary succession in the Mediterranean region. Seedlings ofPicris hieracioides from populations of the early- (1year)and the mid-stages of field abandonment (15–40 years) were transplantedintwo adjacent old fields, abandoned for 4 (F4) and 20 years (F20) and thatdiffered markedly in floristic composition and vegetation structure. For twoyears, we experimentally manipulated competition (no-neighbours vs. naturalvegetation) and resource availability (addition of water and fertiliser vs.controls) in an attempt to evaluate their influence on survival, reproductivetiming, growth and reproductive output throughout the life cycle. Earlymortality was higher in non vegetated plots in both fields. Mortality ofseedlings was mainly due to herbivory by larvae of genusAgriotes. Flowering throughout the whole experiment wasalso facilitated by vegetation in the F4 field as a result of the positiveeffect of annual vegetation and remained unaffected in the F20 field because ofthe high competitive effect of established perennial vegetation. The additionofresources altered the effect of facilitation and competition on late seedlingsurvival. Survival was enhanced in the vegetated plots of the F4 field, becauseresource addition increased the shade provided by the canopy of vegetation,protected seedlings from temperature extremes and reduced water loss. Seedlingmortality also decreased in the F20 field but in a similar manner to vegetatedand non-vegetated subplots, and consequently the outcome of positive andnegative interactions remained neutral. The net effect of facilitation andcompetition resulted in interference later in the life cycle and appearedthrough final lower growth and reproduction in both fields. However, thegreatercompetition in 1994 than in 1995 in both fields, probably because the size ofthe rosettes, makes them less susceptible to competition, illustrates thedifficulty in predicting the outcome of competition solely of one season forfacultative biennial plants. The relative competition intensity calculatedusingonly survivors (RCI₁) was unaffected by habitat fertility in bothfields. In striking contrast, the relative competition intensity calculatedusing seedling mortality (RCI₂) was significantly higher in subplotswithout resource addition in both fields because of high seedling mortality invegetated subplots. Finally, there were no differences in the net effect offacilitation and interference processes among populations from early and midsuccessional stages showing that phenotypic plasticity buffers theenvironmentalselective pressures linked to successional processes.