Litter identity mediates predator impacts on the functioning of an aquatic detritus-based food web

During past decades, several mechanisms such as resource quality and habitat complexity have been proposed to explain variations in the strength of trophic cascades across ecosystems. In detritus-based headwater streams, litter accumulations constitute both a habitat and a resource for detritivorous macroinvertebrates. Because litter edibility (which promotes trophic cascades) is usually inversely correlated with its structural complexity (which weakens trophic cascades), there is a great scope for stronger trophic cascades in litter accumulations that are dominated by easily degradable litter species. However, it remains unclear how mixing contrasting litter species (conferring both habitat complexity and high quality resource) may influence top–down controls on communities and processes. In enclosures exposed in a second-order stream, we manipulated litter species composition by using two contrasting litter (alder and oak), and the presence–absence of a macroinvertebrate predator (Cordulegaster boltonii larvae), enabling it to effectively exert predation pressure, or not, on detritivores (consumptive versus non-consumptive predation effects). Leaf mass loss, detritivore biomass and community structure were mostly controlled independently by litter identity and mixing and by predator consumption. However, the strength of predator control was mediated by litter quality (stronger on alder), and to a lesser extent by litter mixing (weaker on mixed litter). Refractory litter such as oak leaves may contribute to the structural complexity of the habitat for stream macroinvertebrates, allowing the maintenance of detritivore communities even when strong predation pressure occurs. We suggest that considering the interaction between top–down and bottom–up factors is important when investigating their influence on natural communities and ecosystem processes in detritus-based ecosystems.     

Metacommunity patterns in larval odonates

The growth of metacommunity ecology as a subdiscipline has increased interest in how processes at different spatial scales structure communities. However, there is still a significant knowledge gap with respect to relating the action of niche- and dispersal-assembly mechanisms to observed species distributions across gradients. Surveys of the larval dragonfly community (Odonata: Anisoptera) in 57 lakes and ponds in southeast Michigan were used to evaluate hypotheses about the processes regulating community structure in this system. We considered the roles of both niche- and dispersal-assembly processes in determining patterns of species richness and composition across a habitat gradient involving changes in the extent of habitat permanence, canopy cover, area, and top predator type. We compared observed richness patterns and species distributions in this system to patterns predicted by four general community models: species sorting related to adaptive trade-offs, a developmental constraints hypothesis, dispersal assembly, and a neutral community assemblage. Our results supported neither the developmental constraints nor the neutral-assemblage models. Observed patterns of richness and species distributions were consistent with patterns expected when adaptive tradeoffs and dispersal-assembly mechanisms affect community structure. Adaptive trade-offs appeared to be important in limiting the distributions of species which segregate across the habitat gradient. However, dispersal was important in shaping the distributions of species that utilize habitats with a broad range of hydroperiods and alternative top predator types. Our results also suggest that the relative importance of these mechanisms may change across this habitat gradient and that a metacommunity perspective which incorporates both niche- and dispersal-assembly processes is necessary to understand how communities are organized. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Inter‐specific differences in invader and native fish functional responses illustrate neutral effects on prey but superior invader competitive ability

相似文献   

Optimal foraging when predation risk increases with patch resources: an analysis of pollinators and ambush predators

Pollinators and their predators share innate and learned preferences for high quality flowers. Consequently, pollinators are more likely to encounter predators when visiting the most rewarding flowers. I present a model of how different pollinator species can maximize lifetime resource gains depending on the density and distribution of predators, as well as their vulnerability to capture by predators. For pollinator species that are difficult for predators to capture, the optimal strategy is to visit the most rewarding flowers as long as predator density is low. At higher predator densities and for pollinators that are more vulnerable to predator capture, the lifetime resource gain from the most rewarding flowers declines and the optimal strategy depends on the predator distribution. In some cases, a wide range of floral rewards provides near‐maximum lifetime resource gains, which may favor generalization if searching for flowers is costly. In other cases, a low flower reward level provides the maximum lifetime resource gain and so pollinators should specialize on less rewarding flowers. Thus, the model suggests that predators can have qualitatively different top‐down effects on plant reproductive success depending on the pollinator species, the density of predators, and the distribution of predators across flower reward levels.     

Resource limitation,biodiversity, and competitive effects interact to determine the invasibility of rock pool microcosms

The success of species invasions depends on both the characteristics of the invaded habitat and the traits of the invasive species. At local scales biodiversity may act as a barrier to invasion; however, the mechanism by which biodiversity confers invasion resistance to a community has been the subject of considerable debate. The purpose of this study was to test the hypothesis that productivity and diversity affected the ability of a regionally available species to colonize communities from which it is absent. We hypothesized that the invasibility of rock pool invertebrate communities would increase with increasing nutrients and decrease with increasing diversity. We tested this possibility using naturally invaded outdoor aquatic microcosms. We demonstrated that the invasibility of an experimental multi-trophic aquatic community by a competitive native midge species (Ceratopogonidae: Dasyhelea sp.) was determined by an interaction between resource availability, diversity, and the densities of two competitive ostracods species. Nutrient enrichment increased invasion success; however, within nutrient-enriched microcosms, invasion success was highest in the low-diversity treatments. Our results suggest that resource availability may in fact be the principal mechanism determining invasibility at local scales in multi-trophic rock pool communities; however resource availability can be determined by both nutrient input as well as by the diversity of the biotic community.     

The roles of stochasticity and biotic interactions in the spatial patterning of plant species in alpine communities

Questions

Plant community composition can be influenced by multiple biotic, abiotic, and stochastic factors acting on the local species pool to determine their establishment success and abundance and subsequently the diversity of the community. We asked if the influences of biotic interactions on the composition of plant species in communities, as indicated by patterns of plant species spatial associations (independent, positive or negative), vary across a productivity gradient within a single ecosystem type. Do dominant species of communities show spatial patterning suggestive of competitive interactions with interspecific neighbors? Do species that span multiple community types exhibit the same heterospecific interactions with neighbours in each community?

Location

Three alpine communities in the southern Rocky Mountains.

Methods

We measured the occurrence of species in a 1‐cm spatial grid within 2 m × 2 m plots to determine the spatial patterns of species pairs in the three communities. A null model of independent species spatial arrangements was used to determine whether species pairs were positively, negatively or independently associated, and how these patterns differed among the communities across the gradient of resource supply and environmental stress.

Results

Positive associations, indicative of facilitation between species, were most common in the most resource‐poor and least productive community. However negative associations, suggestive of competitive interactions among species, were not more common in the two more resource‐rich, productive communities. The dominant species of these communities did exhibit higher negative than positive associations with neighbours relative to positive patterning. Independent interspecific patterning was equally common relative to positive and negative patterns in all communities. Species that previously were shown to either facilitate other species or compete with neighbours exhibited spatial patterning consistent with the earlier experimental work.

Conclusions

A large number of species exhibit a lack of net biotic interactions, and stochastic factors appear to be as important as competition and facilitation in shaping the structure of the three alpine plant communities we studied.

     

Soil Heterogeneity Effects on Tallgrass Prairie Community Heterogeneity: An Application of Ecological Theory to Restoration Ecology

Spatial heterogeneity of resources can influence plant community composition and diversity in natural communities. We manipulated soil depth (two levels) and nutrient availability (three levels) to create four heterogeneity treatments (no heterogeneity, depth heterogeneity, nutrient heterogeneity, and depth + nutrient heterogeneity) replicated in an agricultural field seeded to native prairie species. Our objective was to determine whether resource heterogeneity influences species diversity and the trajectory of community development during grassland restoration. The treatments significantly increased heterogeneity of available inorganic nitrogen (N), soil water content, and light penetration. Plant diversity was indirectly related to resource heterogeneity through positive relationships with variability in productivity and cover established by the belowground manipulations. Diversity was inversely correlated with the average cover of the dominant grass, Switchgrass (Panicum virgatum), which increased over time in all heterogeneity treatments and resulted in community convergence among the heterogeneity treatments over time. The success of this cultivar across the wide range of resource availability was attributed to net photosynthesis rates equivalent to or higher than those of the native prairie plants in the presence of lower foliar N content. Our results suggest that resource heterogeneity alone may not increase diversity in restorations where a dominant species can successfully establish across the range of resource availability. This is consistent with theory regarding the role of ecological filters on community assembly in that the establishment of one species best adapted for the physical and biological conditions can play an inordinately important role in determining community structure.     

Your worst enemy could be your best friend: predator contributions to invasion resistance and persistence of natives

Native predators are postulated to have an important role in biotic resistance of communities to invasion and community resilience. Effects of predators can be complex, and mechanisms by which predators affect invasion success and impact are understood for only a few well-studied communities. We tested experimentally whether a native predator limits an invasive species’ success and impact on a native competitor for a community of aquatic insect larvae in water-filled containers. The native mosquito Aedes triseriatus alone had no significant effect on abundance of the invasive mosquito Aedes albopictus. The native predatory midge Corethrella appendiculata, at low or high density, significantly reduced A. albopictus abundance. This effect was not caused by trait-mediated oviposition avoidance of containers with predators, but instead was a density-mediated effect caused by predator-induced mortality. The presence of this predator significantly reduced survivorship of the native species, but high predator density also significantly increased development rate of the native species when the invader was present, consistent with predator-mediated release from interspecific competition with the invader. Thus, a native predator can indirectly benefit its native prey when a superior competitor invades. This shows the importance of native predators as a component of biodiversity for both biotic resistance to invasion and resilience of a community perturbed by successful invasion.     

Influence of microclimate and species interactions on the composition of plant and invertebrate communities in alpine northern Norway

We assessed the effects of both biotic processes and abiotic factors on the community composition of vascular plant species and invertebrates at a site in northern Norway. Plant species were assigned to functional (woody versus herbaceous) and biogeographic (boreal versus alpine) groups. Invertebrate species were classified as either herbivore or predator. When species interactions and effects of the abiotic environment were partitioned, boreal species appeared to influence the distribution of alpine species and woody species the distribution of herbaceous species. Analysis of partial correlations indicated that facilitation was the dominant mode of interaction between the two pairs of plant groups. Among abiotic factors, the thermal environment probably influenced all components of the plant and invertebrate communities, except for predatory invertebrates, and wind appeared important in determining the composition of woody and alpine components of the plant community but not the herbaceous component. The composition of the boreal component of the plant community apparently influenced the composition of all invertebrate communities, except for predatory invertebrates. The composition of the woody component of the plant community influenced the composition of both herbivore and predator communities. The alpine plant-community composition influenced predatory invertebrate community composition. Woody plant community composition influenced the composition of both herbivore and predator communities. Our analytic approach, based on two kinds of structural equation models (d-separation and path analysis), provides a useful method for identifying the biotic as well as abiotic factors that influence community structure.     

Fear in the dark? Community‐level effects of non‐lethal predators change with light regime

The total effect of predators on prey is a combination of direct consumption, and non‐consumptive effects (NCEs), such as predator‐induced changes to prey morphology, behaviour and life history. Past research into NCEs has tended to focus on pair‐wise interactions between predators and prey, while in natural ecosystems, species exist in complex communities with several trophic levels made up of multiple autotrophic and heterotropic species. To address how predator NCEs alter the photosynthetic and heterotrophic components of communities, we exposed microbial microcosms to one of three predator treatments: live predators (full predator effect), freeze‐killed predators (NCEs only) or no predators (control), and incubated them under either 12 h:12 h light:dark conditions or continual darkness. Under 12 h:12 h light:dark conditions, NCEs‐only communities never differed from predator‐free communities, but differed from live predator communities. Under conditions of continual darkness, the structure of NCEs‐only communities differed from predator‐free controls, but not from live predator communities, suggesting NCEs can be strong enough to structure communities. Predation threat may cause certain prey to induce defences, such as reductions in movement, which make them less competitive in a community setting. This reduction in competitive ability could lead to these species being driven to extinction through interspecific competition, resulting in similar communities to those in which live predators are present. Heterotrophic species whose rates of resource acquisition depend on movement rates may be affected to a greater extent than autotrophs by predator‐induced reductions in movement, accounting for our observed differences in predator NCEs in ‘dark’ and ‘light’ communities. Our results suggest that the community‐level consequences of fear are greater in the dark. Synthesis Predators affect prey through consumptive and non‐consumptive effects (NCEs) such as alterations to prey behaviour, morphology, and life history. However, predators and prey do not exist in isolated pairs, but in complex communities where they interact with many other species. Using a long term study (>10 predator generations), we show that predator NCEs alone can alter community structure under conditions of darkness, but not in a 12h:12h light:dark cycle. Our results demonstrate for the first time that although the community‐level consequences of predator NCEs may be dramatic, they depend upon the abiotic conditions of the ecosystem.     

Habitat subdivision causes changes in food web structure

Theory suggests that the response of communities to habitat subdivision depends on both species' characteristics and the extent to which species interact. For species with dynamics that are independent of other species, subdivision is expected to promote regional extinction as populations become small and isolated. By contrast, intermediate levels of subdivision can facilitate persistence of strongly interacting species. Consistent with this prediction, experimental subdivision lengthened persistence of some species, altering the extent of food web collapse through extinction. Extended persistence was associated with immigration rescuing a basal prey species from local extinction. As predicted by food web theory, habitat subdivision reduced population density of a top predator. Removal of this top predator from undivided microcosms increased the abundance of two other predator species, and these changes paralleled those produced by habitat subdivision. These results show that species interactions structured this community, and illustrate the need for investigations of other communities.     

Top predator removals have consistent effects on large species despite high environmental variability

Top predator losses affect a wide array of ecological processes, and there is growing evidence that top predators are disproportionately vulnerable to environmental changes. Despite increasing recognition of the fundamental role that top predators play in structuring communities and ecosystems, it remains challenging to predict the consequences of predator extinctions in highly variable environments. Both biotic and abiotic drivers determine community structure, and manipulative experiments are necessary to disentangle the effects of predator loss from other co‐occurring environmental changes. To explore the consistency of top predator effects in ecological communities that experience high local environmental variability, we experimentally removed top predators from arid‐land stream pool mesocosms in southeastern Arizona, USA, and measured natural background environmental conditions. We inoculated mesocosms with aquatic invertebrates from local streams, removed the top predator Abedus herberti (Hemiptera: Belostomatidae) from half of the mesocosms as a treatment, and measured community divergence at the end of the summer dry season. We repeated the experiment in two consecutive years, which represented two very different biotic and abiotic environments. We found that some of the effects of top predator removal were consistent despite significant differences in environmental conditions, community composition, and colonist sources between years. As in other studies, top predator removal did not affect overall species richness or abundance in either year, and we observed inconsistent effects on community and trophic structure. However, top predator removal consistently affected large‐bodied species (those in the top 1% of the community body size distribution) in both years, increasing the abundance of mesopredators and decreasing the abundance of detritivores, even though the identity of these species varied between years. Our findings highlight the vulnerability of large taxa to top predator extirpations and suggest that the consistency of observed ecological patterns may be as important as their magnitude.     

Testing intraguild predation theory in a field system: does numerical dominance shift along a gradient of productivity?

Although ecological theory exists to predict dynamics in communities with intraguild predation (IGP), few empirical tests have examined this theory. IGP theory, in particular, predicts that when two competitors interact via IGP, with increasing resource productivity: (1) the IG predator will increase in abundance as the IG prey declines, and (2) increasing dominance of the IG predator will cause resource density to increase. Here, we provide a first test of these predictions in a field community consisting of a scale insect and its two specialist parasitoids, Aphytis melinus (the IG predator) and Encarsia perniciosi (the IG prey). The shared resource, California red scale, is a pest of citrus, and its productivity varies across a threefold range among citrus cultivars. We examined both absolute and relative densities of parasitoids along this natural gradient of scale productivity in three citrus cultivars (orange, grapefruit and lemon). Although both parasitoid species were found in all three cultivars, their abundances reflected those predicted by IGP theory: the IG prey species dominated at low productivity and the IG predator dominated at high productivity. This relationship was caused by an increase in Aphytis density with productivity. In addition, the density of scale increased with the dominance of the IG predator. These results from a field system demonstrate the important dynamic outcomes for food webs with IGP.     

Interactions between invading benthivorous fish and native whitefish in subarctic lakes

相似文献   

Initial colonization,community assembly and ecosystem function: fungal colonist traits and litter biochemistry mediate decay rate

Priority effects are an important ecological force shaping biotic communities and ecosystem processes, in which the establishment of early colonists alters the colonization success of later‐arriving organisms via competitive exclusion and habitat modification. However, we do not understand which biotic and abiotic conditions lead to strong priority effects and lasting historical contingencies. Using saprotrophic fungi in a model leaf decomposition system, we investigated whether compositional and functional consequences of initial colonization were dependent on initial colonizer traits, resource availability or a combination thereof. To test these ideas, we factorially manipulated leaf litter biochemistry and initial fungal colonist identity, quantifying subsequent community composition, using neutral genetic markers, and community functional characteristics, including enzyme potential and leaf decay rates. During the first 3 months, initial colonist respiration rate and physiological capacity to degrade plant detritus were significant determinants of fungal community composition and leaf decay, indicating that rapid growth and lignolytic potential of early colonists contributed to altered trajectories of community assembly. Further, initial colonization on oak leaves generated increasingly divergent trajectories of fungal community composition and enzyme potential, indicating stronger initial colonizer effects on energy‐poor substrates. Together, these observations provide evidence that initial colonization effects, and subsequent consequences on litter decay, are dependent upon substrate biochemistry and physiological traits within a regional species pool. Because microbial decay of plant detritus is important to global C storage, our results demonstrate that understanding the mechanisms by which initial conditions alter priority effects during community assembly may be key to understanding the drivers of ecosystem‐level processes.     

Elevated success of multispecies bacterial invasions impacts community composition during ecological succession

Successful microbial invasions are determined by a species’ ability to occupy a niche in the new habitat whilst resisting competitive exclusion by the resident community. Despite the recognised importance of biotic factors in determining the invasiveness of microbial communities, the success and impact of multiple concurrent invaders on the resident community has not been examined. Simultaneous invasions might have synergistic effects, for example if resident species need to exhibit divergent phenotypes to compete with the invasive populations. We used three phylogenetically diverse bacterial species to invade two compositionally distinct communities in a controlled, naturalised in vitro system. By initiating the invader introductions at different stages of succession, we could disentangle the relative importance of resident community structure, invader diversity and time pre‐invasion. Our results indicate that multiple invaders increase overall invasion success, but do not alter the successional trajectory of the whole community.     

An evolutionary modelling approach to understanding the factors behind plant invasiveness and community susceptibility to invasion

Ecologists have had limited success in understanding which introduced species may become invasive. An evolutionary model is used to investigate which traits are associated with invasiveness. Translocation experiments were simulated in which species were moved into similar but evolutionarily younger communities. The main findings were that species that had previously been the most abundant in their original communities have significantly higher rates of establishment than did species that had previously occurred at low abundance in their original community. However, if establishment did occur, previously abundant and previously low-abundant species were equally likely to become dominant and were equally likely to exclude other species from their new community. There was a suggestion that the species that were most likely to establish and exclude others were 'genetically' different. When species that had evolved in different simulations (but with identical environmental conditions) were transplanted into communities that had also evolved in different simulations of the same conditions, the outcomes were difficult to predict. Observed rates of establishment and subsequent competitive dominance were observed to be species- and community combination-specific. This evolutionary study represents a novel in silico attempt to tackle invasiveness in an experimental framework and may provide a new methodology for tackling these issues.     

Spatial dynamics of keystone predation

1. I investigated the effects of dispersal on communities of keystone predators and prey. I obtained two key results. 2. First, a strong trade-off between competitive ability and predator susceptibility allows consumer coexistence over a large resource productivity range, but it also lowers the predator-susceptible superior competitor's abundance and increases its risk of extinction. Thus, unexpectedly, dispersal plays a more important role in coexistence when predator-mediated coexistence is strong rather than weak. The interplay between the trade-off, small population sizes resulting from transient oscillations, and dispersal leads to qualitatively different species distributions depending on the relative mobilities of the consumers and predator. These differences yield comparative predictions that can be tested with data on trade-off strength, dispersal rates, and species distributions across productivity gradients. 3. Second, there is an asymmetry between species in their dispersal effects: the predator-resistant inferior competitor's dispersal has a large effect, but the predator-susceptible superior competitor's dispersal has no effect, on coexistence and species' distributions. The inferior competitor's dispersal also mediates the predator's dispersal effects: the predator's dispersal has no effect when the inferior competitor is immobile, and a large effect when it is mobile. The net outcome of the direct and indirect effects of the inferior competitor's dispersal is a qualitative change in the species' distributions from interspecific segregation to interspecific aggregation. 4. The important point is that differences between species in how they balance resource acquisition and predator avoidance can lead to unexpected differences in their dispersal effects. While consumer coexistence in the absence of dispersal is driven largely by the top predator, consumer coexistence in the presence of dispersal is driven largely by the predator-resistant inferior competitor.     

Inside‐container effects drive mosquito community structure in Brazilian Atlantic forest

Metacommunity theory is a convenient framework in which to investigate how local communities linked by dispersal influence patterns of species distribution and abundance across large spatial scales. For organisms with complex life cycles, such as mosquitoes, different pressures are expected to act on communities due to behavioral and ecological partitioning of life stages. Adult females select habitats for oviposition, and resulting offspring are confined to that habitat until reaching adult stages capable of flight; outside‐container effects (OCE) (i.e., spatial factors) are thus expected to act more strongly on species distributions as a function of adult dispersal capability, which should be limited by geographic distances between sites. However, larval community dynamics within a habitat are influenced by inside‐container effects (ICE), mainly interactions with conspecifics and heterospecifics (e.g., through effects of competition and predation). We used a field experiment in a mainland‐island scenario to assess whether environmental, spatial, and temporal factors influence mosquito prey and predator distributions and abundances across spatial scales: within‐site, between‐site, and mainland‐island. We also evaluated whether predator abundances inside containers play a stronger role in shaping mosquito prey community structure than do OCE (e.g., spatial and environmental factors). Temporal influence was more important for predators than for prey mosquito community structure, and the changes in prey mosquito species composition over time appear to be driven by changes in predator abundances. There was a negligible effect of spatial and environmental factors on mosquito community structure, and temporal effects on mosquito abundances and distributions appear to be driven by changes in abundance of the dominant predator, perhaps because ICE are stronger than OCE due to larval habitat restriction, or because adult dispersal is not limited at the chosen spatial scales.     

Seed supply, recruitment, and assembly: quantifying relative seed and establishment limitation in a plant community context

There is a growing consensus that the relative constraints of seed limitation and establishment limitation in recruitment strongly influence abundance patterns in plant communities. Although these constraints have direct relevance to coexistence, most investigations utilize a seed addition approach that offers limited insight into these dynamics. Here we report the results of an assembly experiment with annual plant species from California grasslands to examine how propagule pool characteristics (dominant species abundance, functional diversity) influence establishment and seed limitation (density independence and density dependence across a gradient of seed supply) for each species, as well as how these constraints affect community diversity. Species were predominantly colimited by seed and establishment constraints, exhibiting saturating recruitment functions with increased seed supply. Consistent with competition-colonization trade-off predictions, recruitment constraints often depended on the degree of seed limitation of the competitive dominant, Brassica nigra; diversity was greatest in communities where Brassica was seed limited. Functional similarity within the propagule pool did not affect recruitment across a range of seed supply; likewise, functional diversity of the propagule pool was not related to community diversity. We conclude that seed limitation of the dominant species rather than niche similarity influences interspecific competition for safe sites and scales up to affect community-level diversity.