Predator dispersal determines the effect of connectivity on prey diversity

Linking local communities to a metacommunity can positively affect diversity by enabling immigration of dispersal-limited species and maintenance of sink populations. However, connectivity can also negatively affect diversity by allowing the spread of strong competitors or predators. In a microcosm experiment with five ciliate species as prey and a copepod as an efficient generalist predator, we analysed the effect of connectivity on prey species richness in metacommunities that were either unconnected, connected for the prey, or connected for both prey and predator. Presence and absence of predator dispersal was cross-classified with low and high connectivity. The effect of connectivity on local and regional richness strongly depended on whether corridors were open for the predator. Local richness was initially positively affected by connectivity through rescue of species from stochastic extinctions. With predator dispersal, however, this positive effect soon turned negative as the predator spread over the metacommunity. Regional richness was unaffected by connectivity when local communities were connected only for the prey, while predator dispersal resulted in a pronounced decrease of regional richness. The level of connectivity influenced the speed of richness decline, with regional species extinctions being delayed for one week in weakly connected metacommunities. While connectivity enabled rescue of prey species from stochastic extinctions, deterministic extinctions due to predation were not overcome through reimmigration from predator-free refuges. Prey reimmigrating into these sink habitats appeared to be directly converted into increased predator abundance. Connectivity thus had a positive effect on the predator, even when the predator was not dispersing itself. Our study illustrates that dispersal of a species with strong negative effects on other community members shapes the dispersal-diversity relationship. When connections enable the spread of a generalist predator, positive effects of connectivity on prey species richness are outweighed by regional extinctions through predation.     

A plant diversity×water chemistry experiment in subalpine grassland

Plant diversity has been shown to drive important ecosystem functions such as productivity. At the same time, plant diversity and species composition are altered in alpine ecosystems by human impacts such as skiing. Therefore, we investigated impacts of decreased species richness and ski piste treatments on ecosystem functions in subalpine grassland.Species richness manipulations were combined with nutrient input from snow cover treated with snow additives that are commonly used on ski pistes. Three different species richness levels containing 1, 3 or 9 species randomly selected from a larger pool plus unmanipulated meadow plots were treated with four water types to simulate melt water. One water type contained the snow additive ammonium nitrate. Invasion into the communities was prevented by weeding during 2 years and allowed in three subsequent years.Higher species richness increased plant cover and biomass and decreased their variation. The number of functional groups in a plant assemblage had a positive effect on plant growth. Ammonium nitrate strongly increased biomass and plant cover after a single application but decreased species richness in originally diverse meadow plots. There was no significant interaction between species richness and water-type treatments.After the cessation of weeding, the species richness of different plot types converged within 3 years due to invasion. Nevertheless, relationships between initial species richness and plant cover remained positive.The results suggest that the diversity and species composition of alpine vegetation are important factors influencing cover and biomass, in particular during re-colonization of bare ground after disturbances such as ski-piste construction. In slow-growing alpine vegetation, initially positive diversity effects may remain even after successional convergence of species richness due to invasion. The negative effect of ammonium nitrate on species richness suggests the snow additives should only be used with care.     

Species loss and secondary extinctions in simple and complex model communities

1. The loss of a species from an ecological community can trigger a cascade of secondary extinctions. Here we investigate how the complexity (connectance) of model communities affects their response to species loss. Using dynamic analysis based on a global criterion of persistence (permanence) and topological analysis we investigate the extent of secondary extinctions following the loss of different kinds of species. 2. We show that complex communities are, on average, more resistant to species loss than simple communities: the number of secondary extinctions decreases with increasing connectance. However, complex communities are more vulnerable to loss of top predators than simple communities. 3. The loss of highly connected species (species with many links to other species) and species at low trophic levels triggers, on average, the largest number of secondary extinctions. The effect of the connectivity of a species is strongest in webs with low connectance. 4. Most secondary extinctions are due to direct bottom-up effects: consumers go extinct when their resources are lost. Secondary extinctions due to trophic cascades and disruption of predator-mediated coexistence also occur. Secondary extinctions due to disruption of predator-mediated coexistence are more common in complex communities than in simple communities, while bottom-up and top-down extinction cascades are more common in simple communities. 5. Topological analysis of the response of communities to species loss always predicts a lower number of secondary extinctions than dynamic analysis, especially in food webs with high connectance.     

Environmental drivers of spider community composition at multiple scales along an urban gradient

Broad-scale modification of natural ecosystems associated with urbanisation often leads to localised extinctions and reduced species richness. Despite this, habitats within the urban matrix are still capable of supporting biodiversity to varying degrees. As species have different responses to anthropogenic habitat modification, the species composition of urban areas can depend greatly on the habitat characteristics of the local and surrounding areas. The aim of this study was to compare the community composition of spiders in private gardens, urban parks, patches of remnant vegetation and continuous bushland sites, so as to identify habitat variables associated with variation in spider populations along and within the urban gradient and matrix. Overall spider abundances and richness were highest in remnant vegetation patches and were associated with increased vegetation cover at microhabitat and landscape-scales. While gardens were not as diverse as remnant patches, they did support a surprisingly high diversity of spiders. We also found that species composition differed significantly between gardens and other urban green spaces. Higher richness within gardens was also associated with greater vegetation cover, indicating the importance of private management decisions on local biodiversity. Differences in community composition between land-use types were driven by a small number of urban-tolerant species, and spider guilds showed different responses to habitat traits such as vegetation cover and human population densities. This study demonstrates that urban land-uses support unique spider communities and that maintaining vegetation cover within the urban matrix is essential in order to support diverse spider communities in cities.     

茂县土地岭植被恢复过程中物种多样性动态特征

植被恢复是退化生态系统重建的重要途径,植被恢复过程物种多样性的变化反映了植被的恢复程度.通过群落调查和多样性分析,研究了岷江上游土地岭植被恢复过程中群落物种多样性特征.结果表明： 恢复过程中6类不同类型群落分别表现其对于不同环境特征、干扰及更新方式等的响应;森林是较灌丛更适合当地环境状况的植被类型;人工恢复无干扰和轻度干扰群落的多样性相对较高,是较好的恢复模式.重度干扰使得1年生植物与地下芽植物比例增加,其它口食性较好的多年生草本减少.较强的干扰是群落无法更新、长期处于灌丛阶段且多样性较低的重要原因.本地区人工恢复群落在更新进程和多样性维持上优于自然更新群落,种植华山松加速了本地区植被演替进程.建议以适合恢复区域的多种恢复配置方式进行造林,并避免较强干扰,可以加速群落演替进程并保持恢复群落较高的物种丰富度与多样性.     

Testing for local species saturation with nonindependent regional species pools

Identifying the factors controlling local community structure is a central problem in ecology. Ecologists frequently use regression to test for a nonlinear saturating relationship between local community richness and regional species pool richness, suggesting that species interactions limit the number of locally coexisting species. However, communities in different regions are not independent if regions share species. We present a Monte Carlo test for whether an observed local-regional richness relationship is significantly different from that expected when regions are nonindependent and species interactions do not limit community membership. We illustrate this test with data from experimental microcosm communities. A conventional F -test suggests a significant saturating relationship between realized community richness and species pool richness. However, the Monte Carlo test fails to reject the null hypothesis that species interactions do not affect community richness. Strong species interactions do not necessarily set an absolute upper limit to the number of locally coexisting species.     

Invertebrate community response to fire and rodent activity in the Mojave and Great Basin Deserts

Recent increases in the frequency and size of desert wildfires bring into question the impacts of fire on desert invertebrate communities. Furthermore, consumer communities can strongly impact invertebrates through predation and top‐down effects on plant community assembly. We experimentally applied burn and rodent exclusion treatments in a full factorial design at sites in both the Mojave and Great Basin deserts to examine the impact that fire and rodent consumers have on invertebrate communities. Pitfall traps were used to survey invertebrates from April through September 2016 to determine changes in abundance, richness, and diversity of invertebrate communities in response to fire and rodent treatments. Generally speaking, rodent exclusion had very little effect on invertebrate abundance or ant abundance, richness or diversity. The one exception was ant abundance, which was higher in rodent access plots than in rodent exclusion plots in June 2016, but only at the Great Basin site. Fire had little effect on the abundances of invertebrate groups at either desert site, with the exception of a negative effect on flying‐forager abundance at our Great Basin site. However, fire reduced ant species richness and Shannon's diversity at both desert sites. Fire did appear to indirectly affect ant community composition by altering plant community composition. Structural equation models suggest that fire increased invasive plant cover, which negatively impacted ant species richness and Shannon's diversity, a pattern that was consistent at both desert sites. These results suggest that invertebrate communities demonstrate some resilience to fire and invasions but increasing fire and spread of invasive due to invasive grass fire cycles may put increasing pressure on the stability of invertebrate communities.     

Secondary succession and summer herbivory in a subarctic grassland: community structure and diversity

A field experiment was established in a subarctic grassland in the Finnish Lapland to study the role of summer herbivory in plant community succession Perennial vegetation and moss cover were removed in an area of 324 m² The site was divided into four blocks, of which two were fenced to prevent herbivory by large mammals (reindeer, hare)
Early successional changes in the vegetation were assessed Mean species richness per 3 × 3 m plot was consistently higher in the fenced area, indicating that herbivory can suppress small-scale diversity Herbivory affected the height of several plant species However, there was no correlation between frequency and height of individual species There was a weak indication that taller species were more successful m early succession when grazed Light competition is apparently not a key process determining successional change Thus, in early stage of succession, summer herbivory has little effect on diversity by limiting light competition, and most species are equally successful in grazed and ungrazed plots There was some indirect evidence about competitive interactions in the developing community However, unlike temperate grasslands, large mammal herbivory and competition for light seem not to be important determinants of community change in this subarctic grassland (at least what concernes early successional stages) This may be explained by the harshness of local climate, and abundance of light due to the polar day     

Browsing‐mediated shrub canopy changes drive composition and species richness in forest‐tundra ecosystems

Changes in climate and in browsing pressure are expected to alter the abundance of tundra shrubs thereby influencing the composition and species richness of plant communities. We investigated the associations between browsing, tundra shrub canopies and their understory vegetation by utilizing a long‐term (10–13 seasons) experiment controlling reindeer and ptarmigan herbivory in the subarctic forest tundra ecotone in northwestern Fennoscandia. In this area, there has also been a consistent increase in the yearly thermal sum and precipitation during the study period. The cover of shrubs increased 2.8–7.8 fold in exclosures and these contrasted with browsed control areas creating a sharp gradient of canopy cover of tundra shrubs across a variety of vegetation types. Browsing exclusions caused significant shifts in more productive vegetation types, whereas little or no shift occurred in low‐productive tundra communities. The increased deciduous shrub cover was associated with significant losses of understory plant species and shifts in functional composition, the latter being clearest in the most productive plant community types. The total cover of understory vegetation decreased along with increasing shrub cover, while the cover of litter showed the opposite response. The cover of cryptogams decreased along with increasing shrub cover, while the cover of forbs was favoured by a shrub cover. Increasing shrub cover decreased species richness of understory vegetation, which was mainly due to the decrease in the cryptogam species. The effects were consistent across different types of forest tundra vegetation indicating that shrub increase may have broad impacts on arctic vegetation diversity. Deciduous shrub cover is strongly regulated by reindeer browsing pressure and altered browsing pressure may result in a profound shrub expansion over the next one or two decades. Results suggest that the impact of an increase in shrubs on tundra plant richness is strong and browsing pressure effectively counteracts the effects of climate warming‐driven shrub expansion and hence maintains species richness.     

Initial soil amendments still affect plant community composition after nine years in succession on a heavy metal contaminated mountainside

Many efforts to restore disturbed landscapes seek to meet ecological goals over timescales from decades to centuries. It is thus crucial to know how different actions available to restoration practitioners may affect ecosystems in the long term, yet few such data exist. Here, we test the effects of seed and compost applications on plant community composition 9 years after their application, by taking advantage of a well‐controlled restoration experiment on a mountainside severely degraded by over 80 years of zinc smelting emissions. We asked whether plots have converged on similar plant communities regardless of initial seed and compost treatments, or if these initial treatments have given rise to lasting differences in whole plant communities or in the richness and abundance of native, exotic, and planted species. We found that compost types significantly affected plant communities 9 years later, but seed mix species composition did not. Observed differences in species richness and vegetative cover were negatively correlated, and both were related to the differences in plant communities associated with different compost types. These observed differences are due primarily to the number and abundance of species not in original seed mixes, of which notably many are native. Our results underscore the importance of soils in shaping the aboveground composition of ecosystems. Differences in soil characteristics can affect plant diversity and cover, which are both common restoration targets. Even in highly polluted and devegetated sites, compost and seed application can reinstate high vegetative cover and allow continued colonization of native species.     

Effects of Species Richness on Resident and Target Species Components in a Prairie Restoration

The ecological role of biodiversity in achieving successful restoration has been little explored in restoration ecology. We tested the prediction that we are more likely to create persistent, species‐rich plant communities by increasing the number of species sown, and, to some degree, by varying functional group representation, in experimental prairie plantings. There were 12 treatments consisting of 1‐, 2‐, 3‐, 4‐, 8‐, 12‐, and 16‐species mixtures of native perennials representing four functional groups (C₄ grasses, C₃ grasses, nitrogen‐fixing species, and late‐flowering composites) that predominate within Central Plains tallgrass prairies. In 2000, species were seeded into square plots (6 × 6 m), with five replicates per treatment, on former agricultural land. Annually, we measured total species richness and evenness, target species richness and cover, and richness and cover of resident species (i.e., those emerging from the seed bank). Both target species richness and rate of establishment of target communities were highest in the most species‐rich mixtures, but there was no additional benefit for treatments that contained more than eight species. Richness of resident species did not vary with target species richness; however, cover by resident species was lower in the higher target species treatments. Our results, indicating that establishment of species‐rich prairie mimics can be enhanced by starting with larger numbers of species at the outset, have implications for grassland restoration in which community biodiversity creation and maintenance are key goals.     

Altitude effects on spatial components of vascular plant diversity in a subarctic mountain tundra

Environmental gradients are caused by gradual changes in abiotic factors, which affect species abundances and distributions, and are important for the spatial distribution of biodiversity. One prominent environmental gradient is the altitude gradient. Understanding ecological processes associated with altitude gradients may help us to understand the possible effects climate change could have on species communities. We quantified vegetation cover, species richness, species evenness, beta diversity, and spatial patterns of community structure of vascular plants along altitude gradients in a subarctic mountain tundra in northern Sweden. Vascular plant cover and plant species richness showed unimodal relationships with altitude. However, species evenness did not change with altitude, suggesting that no individual species became dominant when species richness declined. Beta diversity also showed a unimodal relationship with altitude, but only for an intermediate spatial scale of 1 km. A lack of relationships with altitude for either patch or landscape scales suggests that any altitude effects on plant spatial heterogeneity occurred on scales larger than individual patches but were not effective across the whole landscape. We observed both nested and modular patterns of community structures, but only the modular patterns corresponded with altitude. Our observations point to biotic regulations of plant communities at high altitudes, but we found both scale dependencies and inconsistent magnitude of the effects of altitude on different diversity components. We urge for further studies evaluating how different factors influence plant communities in high altitude and high latitude environments, as well as studies identifying scale and context dependencies in any such influences.     

Global versus local extinction in a network model of plant–pollinator communities

The loss of a species from an ecological community can trigger a cascade of additional extinctions; the complex interactions that comprise ecological communities make the dynamics and impacts of such a cascade challenging to predict. Previous studies have typically considered global extinctions, where a species cannot re-enter a community once it is lost. However, in some cases a species only becomes locally extinct, and may be able to reinvade from surrounding communities. Here, we use a dynamic, Boolean network model of plant–pollinator community assembly to analyze the differences between global and local extinction events in mutualistic communities. As expected, we find that compared to global extinctions, communities respond to local extinctions with lower biodiversity loss, and less variation in topological network properties. We demonstrate that in the face of global extinctions, larger communities suffer greater biodiversity loss than smaller communities when similar proportions of species are lost. Conversely, smaller communities suffer greater loss in the face of local extinctions. We show that targeting species with the most interacting partners causes more biodiversity loss than random extinctions in the case of global, but not local, extinctions. These results extend our understanding of how mutualistic communities respond to species loss, with implications for community management and conservation efforts.     

Cascade effects of crop species richness on the diversity of pest insects and their natural enemies

Understanding how plant species richness influences the diversity of herbivorous and predatory/parasitic arthropods is central to community ecology.We explore the effects of crop species richness on the diversity of pest insects and their natural enemies.Using data from a four-year experiment with five levels of crop species richness,we found that crop species richness significantly affected the pest species richness,but there were no significant effects on richness of the pests’natural enemies.In contrast,the species richness of pest insects significantly affected their natural enemies.These findings suggest a cascade effect where trophic interactions are strong between adjacent trophic levels,while the interactions between connected but nonadjacent trophic levels are weakened by the intermediate trophic level.High crop species richness resulted in a more stable arthropod community compared with communities in monoculture crops.Our results highlight the complicated cross-trophic interactions and the crucial role of crop diversity in the food webs of agro-ecosystems.     

Impacts and negative feedbacks in community recovery over eight years following removal of habitat-forming macroalgae

The decline or loss of habitat-forming species has affected rocky shore marine communities worldwide. Many short-term studies have documented the initiation of cascading effects due to canopy losses of macroalgae, but relatively few studies have followed recovery dynamics over many years. Here, we show that the experimental removal of a dominant intertidal fucoid in southern New Zealand had numerous community effects up to 8 years later. Even though the dominant fucoid returned to a nearly closed canopy, there remained many differences between disturbed and control communities. The disturbed treatments had lower plant density and biomass of the dominant fucoid, fewer species and more turfing coralline algae than controls. Plots with press disturbances were more affected than those with pulse disturbances. The negative feedback of turfing algae on the recovery of fucoid recruitment resulted in effects on cover and diversity in the wider community being evident after 8 years. We discuss the feedbacks between fucoids, benthic turfing algae and community recovery and argue that if biodiversity impacts on marine rocky reefs are to be understood, the role of non-trophic interactions in structure, function and dynamics must be better delineated.     

Changes in the dominant assembly mechanism drive species loss caused by declining resources

The species–energy hypothesis predicts that more productive areas support higher species richness. Conversely, when resources are reduced, species richness is reduced. Empirical tests of whether extinctions are predominantly caused by environmental constraints or competitive exclusion are lacking. We experimentally reduced dead wood to c. 15% of the initial amount after a major windstorm and examined changes in assembly mechanisms by combining trait‐based and evolutionary species dissimilarities of eight taxonomic groups, differing in their dependence on dead wood (saproxylic/non‐saproxylic). Species richness and assembly mechanisms of non‐saproxylic taxa remained largely unaffected by removal of dead wood. By contrast, extinctions of saproxylic species were caused by reversing the predominant assembly mechanisms (e.g. increasing importance of competitive exclusion for communities assembled through environmental filtering or vice versa). We found no evidence for an intensification of the predominant assembly mechanism (e.g. competitive exclusion becoming stronger in a competitively structured community).     

The effect of overstorey proteas on plant species richness in South African mountain fynbos

Two South African mountain fynbos sites, similar in drainage, elevation, slope angle, slope aspect and soil type but with differing fire histories, were studied to measure how the effect of high densities of overstorey proteas in one fire cycle affects the α-diversity levels of the plant community in the following fire-cycle, how their repeated absence due to several short fire-cycles affects their species richness and finally, at what spatial scale such patterns are most appropriately measured. High prefire canopy cover percentages and densities of overstorey proteas increase the postfire α-diversity of understorey species. In addition, the increase in species richness observed occurred for all higher plant life history types present. At sites where one or more short fire cycles resulted in the repeated absence of overstorey proteas, the number of plant species present in the understorey was lower than at a site where overstorey proteas persisted. These results are dependent on the spatial scale at which the α-diversity of understorey species is measured. At small quadrat sizes (< 5 m²), overstorey proteas decrease the number of understorey species present, while at larger quadrat sizes (100 m²) higher species richness is observed. The contradiction in conclusions when α-diversity is measured at different spatial scales can be attributed to the patchiness of fynbos communities. Overstorey proteas play an important role in maintaining the patchiness component of fynbos communities by diminishing the effect of understorey resprouting species, making available regeneration niches for the maintenance of plant species richness. Where small quadrats are used, the effect of patchiness on the dynamics of the mountain fynbos community is lost. Thus, it is the fire history prior to the last fire and how it affects overstorey proteas that is important in the determination of α-diversity levels in mountain fynbos plant communities.     

Keystone species and vulnerable species in ecological communities: strong or weak interactors?

The loss of a species from an ecological community can trigger a cascade of secondary extinctions. The probability of secondary extinction to take place and the number of secondary extinctions are likely to depend on the characteristics of the species that is lost--the strength of its interactions with other species--as well as on the distribution of interaction strengths in the whole community. Analysing the effects of species loss in model communities we found that removal of the following species categories triggered, on average, the largest number of secondary extinctions: (a) rare species interacting strongly with many consumers, (b) abundant basal species interacting weakly with their consumers and (c) abundant intermediate species interacting strongly with many resources. We also found that the keystone status of a species with given characteristics was context dependent, that is, dependent on the structure of the community where it was embedded. Species vulnerable to secondary extinctions were mainly species interacting weakly with their resources and species interacting strongly with their consumers.     

Different Assembly Processes Drive Shifts in Species and Functional Composition in Experimental Grasslands Varying in Sown Diversity and Community History

Background

The prevalence of different biotic processes (limiting similarity, weaker competitor exclusion) and historical contingency due to priority effects are in the focus of ongoing discussions about community assembly and non-random functional trait distributions.

Methodology/Principal Findings

We experimentally manipulated assembly history in a grassland biodiversity experiment (Jena Experiment) by applying two factorially crossed split-plot treatments to all communities: (i) duration of weeding (never weeded since sowing or cessation of weeding after 3 or 6 years); (ii) seed addition (control vs. seed addition 4 years after sowing). Spontaneous colonization of new species in the control treatment without seed addition increased realized species richness and functional richness (FRic), indicating continuously denser packing of niches. Seed addition resulted in forced colonization and increased realized species richness, FRic, functional evenness (FEve) and functional divergence (FDiv), i.e. higher abundances of species with extreme trait values. Furthermore, the colonization of new species led to a decline in FEve through time, suggesting that weaker competitors were reduced in abundance or excluded. Communities with higher initial species richness or with longer time since cessation of weeding were more restricted in the entry of new species and showed smaller increases in FRic after seed addition than other communities. The two assembly-history treatments caused a divergence of species compositions within communities originally established with the same species. Communities originally established with different species converged in species richness and functional trait composition over time, but remained more distinct in species composition.

Conclusions/Significance

Contrasting biotic processes (limiting similarity, weaker competitor exclusion) increase functional convergence between communities initially established with different species. Historical contingency with regard to realized species compositions could not be eradicated by cessation of weeding or forced colonization and was still detectable 5 years after application of these treatments, providing evidence for the role of priority effects in community assembly.     

Genotypic richness and dissimilarity opposingly affect ecosystem functioning

Biodiversity is an essential determinant of ecosystem functioning. Numerous studies described positive effects of diversity on the functioning of communities arising from complementary resource use and facilitation. However, high biodiversity may also increase competitive interactions, fostering antagonism and negatively affecting community performance. Using experimental bacterial communities we differentiated diversity effects based on genotypic richness and dissimilarity. We show that these diversity characteristics have opposite effects on ecosystem functioning. Genotypic dissimilarity governed complementary resource use, improving ecosystem functioning in complex resource environments. Contrastingly, genotypic richness drove allelopathic interactions, mostly reducing ecosystem functioning. The net biodiversity effect on community performance resulted from the interplay between the genetic structure of the community and resource complexity. These results demonstrate that increasing richness, without concomitantly increasing dissimilarity, can decrease ecosystem functioning in simple environments due to antagonistic interactions, an effect insufficiently considered so far in mechanistic models of the biodiversity-ecosystem functioning relationship.