Testing the role of biotic stress in the stress gradient hypothesis. Processes and patterns in arid rangelands

Since many arid ecosystems are overstocked with domestic herbivores, biotic stress could have a stronger influence in modulating the balance of species interactions than expected from the stress gradient hypothesis (SGH). Here we tested a priori predictions about the effect of grazing on species interactions and fine scale spatial structure of grasses in water‐limited ecosystems. We used detailed vegetation mapping and spatial analysis, and performed a field experiment where the direct and indirect components of positive interactions were disentangled to provide evidence of links between process and pattern. We found associational resistance (biotic refuge) to be the dominant process in grazing situations, while competition, instead of direct facilitation, seemed to govern grass spatial patterns when herbivore pressure was relaxed. These results suggest that facilitation between grasses in arid communities may be related to herbivory rather than nurse plant effects. Associational resistance tends to have the strongest effect on spatial aggregation of species at intermediate grazing pressure. Results suggest that contrary to SGH, this physical clustering of species decreased when grazing pressure reached their maximum levels. Positive associations remained significant only when palatability differences between neighbours is large, suggesting that managing stocking rate is a key factor determining the persistence of herbivory refuges. These refuges are potential foci to initiate population recovery of high quality forage species in arid degraded areas.     

Release from competition and protection determine the outcome of plant interactions along a grazing gradient

Plant interactions are suggested to shift from competition to facilitation and collapse with increasing grazing pressure. The existence of this full range of plant interactions and the role of underlying mechanisms (i.e. release from competition and protecting effect) in response to herbivory remains poorly documented and mainly described in terrestrial systems. We use a large grazing disturbance gradient (five levels of grazing) to test its effect on the outcome of plant interactions and underlying mechanisms in freshwater ecosystems. In a mesocosm experiment, we manipulated the presence of neighbouring plants to test their negative (competition) or protective (facilitation) effects on target plants along the grazing pressure gradient. We predicted that plant interactions 1) shift from competition to indirect facilitation with increased grazing pressure, 2) indirect facilitation collapses at high levels of grazing, 3) release from competition mainly drives the outcome in lowly grazed conditions and, 4) decreased protection occurs in highly grazed conditions responsible for the collapse of facilitation. This study shows the occurrence of the full range of outcomes in plant interactions under a wide spectrum of grazing pressure and indicates how the complex combination of underlying mechanisms shapes variations in plant interactions. We show that both, the release from competition and the increased protection by neighbouring plants drove the shift from competition to indirect facilitation. Declined protection by neighbouring plants resulted in a collapse of indirect facilitation for survival under intense herbivory. Our study provides the first experimental evidence of indirect facilitation structuring freshwater ecosystems thereby validating important ecological concepts mainly developed for terrestrial ecosystems.     

Plant–plant facilitation increases with reduced phylogenetic relatedness along an elevation gradient

Environmental conditions can modify the intensity and sign of ecological interactions. The stress gradient hypothesis (SGH) predicts that facilitation becomes more important than competition under stressful conditions. To properly test this hypothesis, it is necessary to account for all (not a subset of) interactions occurring in the communities and consider that species do not interact at random but following a specific pattern. We aim to assess elevational changes in facilitation, in terms of species richness, frequency and intensity of the interaction as a function of the evolutionary relatedness between nurses and their associated species. We sampled nurse and their facilitated plant species in two 1000–2000 m. elevation gradients in Mediterranean Chile where low temperature imposes a mortality filter on seedlings. We first estimated the relative importance of facilitation as a mechanism adding new species to communities distributed along these gradients. We then tested whether the frequency and intensity of facilitation increases with elevation, taking into account the evolutionary relatedness of the nurse species and the facilitated species. We found that nurses increase the species richness of the community by up to 35%. Facilitative interactions are more frequent than competitive interactions (56% versus 44%) and facilitation intensity increased with elevation for interactions involving distantly related lineages. Our results highlight the importance of including an evolutionary dimension in the study of facilitation to have a clearer picture of the mechanisms enabling species to coexist and survive under stressful conditions. This knowledge is especially relevant to conserve vulnerable and threatened communities facing new climate scenarios, such as those located in Mediterranean-type ecosystems.     

Direct and indirect interactions co‐determine species composition in nurse plant systems

Facilitation by nurse plants plays an important role in determining community composition in severe environments. Although the unidirectional effect of nurses on beneficiary species has received considerable research interest, nurse‐mediated interactions among beneficiary species (so‐called indirect interactions) are less known. Consequently, community composition in nurse plant systems is generally considered as a simple consequence of the facilitative effect of the nurse even though beneficiary species may significantly contribute to community assembly and modulate the direct nurse effects on the community. In an observational study we assessed nurse effects and nurse‐mediated beneficiary interactions in two contrasting nurse plant systems in dry environments using a newly developed framework. We quantified plant–plant interaction intensity using the relative interaction index (RII) at the community and species level for three Retama sphaerocarpa shrub size‐classes in a semiarid shrubland and four Arenaria tetraquetra cushion plant communities differing in aspect and elevation in dry alpine gravel habitats. The observed RII was split into nurse and beneficiary effects, and related to individual mass, species frequency and abundance using generalized linear mixed models. Results showed predominantly positive nurse effects and negative beneficiary interactions. The effect size of nurse plants, however, was significantly higher than the effect size of beneficiary species in both systems. Individual plant mass and abundance of species was dependent on the combined effects of nurse and beneficiary species whereas species occurrence was related to nurse effects only. Despite evident differences, the semiarid and alpine nurse plant systems showed strong functional parallelisms. We found interdependence between the effects of nurse and beneficiary species on beneficiary plant assemblages emphasizing their combined role on community assembly in both systems. Our results highlight the need to consider indirect interactions to understand fully plant community dynamics.     

Shifting Mosaics in Grazed Woodlands Driven by the Alternation of Plant Facilitation and Competition

Abstract: Free-ranging large grazers, such as cattle and horses, are increasingly reintroduced to former agricultural areas in Western Europe in order to restore natural and diverse habitats. In this review we outline mechanisms by which large grazers induce and maintain structural diversity in the vegetation (mosaics of grasslands, shrub thickets and trees). This variation in vegetation structure is considered to be important for the conservation of biodiversity of various plant and animal groups. The process of spatial association with unpalatable plants (as-sociational resistance) enables palatable plants to establish in grasslands maintained by large grazers. In this way, short unattractive (thorny, low quality or toxic) species facilitate taller unattractive shrubs, which facilitate palatable trees, which in turn outshade the species that facilitated their recruitment. Established trees can, therefore, not regenerate under their own canopy, leading to cyclic patch dynamics. Since this cyclic dynamic occurs on a local scale, this contributes to shifting mosaics. The mechanisms involved in creating and maintaining the resulting shifting mosaics are described for temperate flood-plain and heathland ecosystems, including the effects on nutrient transport within grazed landscapes. How grazing leads to shifting mosaics is described in terms of plant functional types, allowing potential generalisation to other ecosystems. The resulting interaction web of grasses, unpalatable forbs and shrubs, palatable light-demanding trees and shade-tolerant trees is discussed, and was found to contain various interesting direct and indirect effects. The key process contributing to spatial diversity in vegetation structure is the alternation of positive (facilitation) interactions between plant species at one life cycle stage, and competitive displacement at another stage. Grazing thus causes directional successional sequences to change to shifting mosaics. The implications of this theory for nature conservation are discussed, including the relevant management problems, possible choices and practical solutions. We conclude that the theoretical framework outlined in this review provides helpful insights when coping with nature conservation issues in temperate woodland habitats.     

Grazing increases functional richness but not functional divergence in Tibetan alpine meadow plant communities

Plant community diversity and ecosystem function are conditioned by competition among co-occurring species for multiple resources. Previous studies suggest that removal of standing biomass by grazing decreases competition for light, but coincident grazing effects on competition for soil nutrients remain largely unknown in Tibetan rangelands where grazing tends to deplete soil phosphorus availability. We measured five functional traits indicative of plant productivity and stoichiometry leaf carbon concentration (LCC), leaf nitrogen concentration (LNC), leaf phosphorus concentration (LPC), specific leaf area (SLA), leaf dry matter content (LDMC) for component species of plant communities in grazed and ungrazed plots in five Tibetan alpine meadows. We examined the diversity of traits singly Rao index of functional diversity (FDrao) and in aggregate functional richness (FRic), functional divergence (FDiv), and functional evenness (FEve) in response to grazing. We tested whether foliar trait diversity increases with nutrient competition but decreases with light competition when competitive exclusion is reduced by grazing. The FDrao of LPC significantly increased under grazing, but FDrao for LCC, LNC and SLA tended to decrease. The FDrao of LDMC increased at the drier site but decreased at the wettest site. There was a strong negative association between increase in FDrao of LPC and decrease in soil nutrients, especially soil phosphorus availability. The FRic for all five traits together increased with species diversity following grazing, but neither FDiv nor FEve differed significantly between grazed and ungrazed plots at most sites. Grazing in Tibetan alpine meadows tends to increase competition for soil phosphorus while decreasing competition for light, resulting in an increase in the functional richness in grazed plant communities without any significant changes in the overall functional diversity of foliar traits. Our study highlights the potential importance of grazing mediated competition for multiple resources in alpine meadow ecosystems.     

Functional traits determine plant co-occurrence more than environment or evolutionary relatedness in global drylands

Plant–plant interactions are driven by environmental conditions, evolutionary relationships (ER) and the functional traits of the plants involved. However, studies addressing the relative importance of these drivers are rare, but crucial to improve our predictions of the effects of plant–plant interactions on plant communities and of how they respond to differing environmental conditions. To analyze the relative importance of – and interrelationships among – these factors as drivers of plant–plant interactions, we analyzed perennial plant co-occurrence at 106 dryland plant communities established across rainfall gradients in nine countries. We used structural equation modelling to disentangle the relationships between environmental conditions (aridity and soil fertility), functional traits extracted from the literature, and ER, and to assess their relative importance as drivers of the 929 pairwise plant–plant co-occurrence levels measured. Functional traits, specifically facilitated plants’ height and nurse growth form, were of primary importance, and modulated the effect of the environment and ER on plant–plant interactions. Environmental conditions and ER were important mainly for those interactions involving woody and graminoid nurses, respectively. The relative importance of different plant–plant interaction drivers (ER, functional traits, and the environment) varied depending on the region considered, illustrating the difficulty of predicting the outcome of plant–plant interactions at broader spatial scales. In our global-scale study on drylands, plant–plant interactions were more strongly related to functional traits of the species involved than to the environmental variables considered. Thus, moving to a trait-based facilitation/competition approach help to predict that: (1) positive plant–plant interactions are more likely to occur for taller facilitated species in drylands, and (2) plant–plant interactions within woody-dominated ecosystems might be more sensitive to changing environmental conditions than those within grasslands. By providing insights on which species are likely to better perform beneath a given neighbour, our results will also help to succeed in restoration practices involving the use of nurse plants.     

Plant Trait Assembly Affects Superiority of Grazer's Foraging Strategies in Species-Rich Grasslands

Background

Current plant – herbivore interaction models and experiments with mammalian herbivores grazing plant monocultures show the superiority of a maximizing forage quality strategy (MFQ) over a maximizing intake strategy (MI). However, there is a lack of evidence whether grazers comply with the model predictions under field conditions.

Methodology/Findings

We assessed diet selection of sheep (Ovis aries) using plant functional traits in productive mesic vs. low-productivity dry species-rich grasslands dominated by resource-exploitative vs. resource-conservative species respectively. Each grassland type was studied in two replicates for two years. We investigated the first grazing cycle in a set of 288 plots with a diameter of 30 cm, i.e. the size of sheep feeding station. In mesic grasslands, high plot defoliation was associated with community weighted means of leaf traits referring to high forage quality, i.e. low leaf dry matter content (LDMC) and high specific leaf area (SLA), with a high proportion of legumes and the most with high community weighted mean of forage indicator value. In contrast in dry grasslands, high community weighted mean of canopy height, an estimate of forage quantity, was the best predictor of plot defoliation. Similar differences in selection on forage quality vs. quantity were detected within plots. Sheep selected plants with higher forage indicator values than the plot specific community weighted mean of forage indicator value in mesic grasslands whereas taller plants were selected in dry grasslands. However, at this scale sheep avoided legumes and plants with higher SLA, preferred plants with higher LDMC while grazing plants with higher forage indicator values in mesic grasslands.

Conclusions

Our findings indicate that MFQ appears superior over MI only in habitats with a predominance of resource-exploitative species. Furthermore, plant functional traits (LDMC, SLA, nitrogen fixer) seem to be helpful correlates of forage quality only at the community level.     

Direct and indirect facilitation affect community productivity through changes in functional diversity in an alpine system

Background and AimsFacilitation is an important ecological process for plant community structure and functional composition. Although direct facilitation has accrued most of the evidence so far, indirect facilitation is ubiquitous in nature and it has an enormous potential to explain community structuring. In this study, we assess the effect of direct and indirect facilitation on community productivity via taxonomic and functional diversity.MethodsIn an alpine community on the Tibetan Plateau, we manipulated the presence of the shrub Dasiphora fruticosa and graminoids in a fenced meadow and a grazed meadow to quantify the effects of direct and indirect facilitation. We measured four plant traits: height, lateral spread, specific leaf area (SLA) and leaf dry matter content (LDMC) of forbs; calculated two metrics of functional diversity [range of trait and community-weighted mean (CWM) of trait]; and assessed the responses of functional diversity to shrub facilitation. We used structural equation modelling to explore how shrubs directly and indirectly drove community productivity via taxonomic diversity and functional diversity.Key ResultsWe found stronger effects from herbivore-mediated indirect facilitation than direct facilitation on productivity and taxonomic diversity, regardless of the presence of graminoids. For functional diversity, the range and CWM of height and SLA, rather than lateral spread and LDMC, generally increased due to direct and indirect facilitation. Moreover, we found that the range of traits played a primary role over taxonomic diversity and CWM of traits in terms of shrub effects on community productivity.ConclusionsOur study reveals that the mechanism of shrub direct and indirect facilitation of community productivity in this alpine community is expanding the realized niche (i.e. expanding range of traits). Our findings indicate that facilitators might increase trait dispersion in the local community, which could alleviate the effect of environmental filters on trait values in harsh environments, thereby contributing to ecosystem functioning.     

Nurse plants,tree saplings and grazing pressure: changes in facilitation along a biotic environmental gradient

Current conceptual models predict that an increase in stress shifts interactions between plants from competitive to facilitative; hence, facilitation is expected to gain in ecological importance with increasing stress. Little is known about how facilitative interactions between plants change with increasing biotic stress, such as that incurred by consumer pressure or herbivory (i.e. disturbance sensu Grime). In grazed ecosystems, the presence of unpalatable plants is reported to protect tree saplings against cattle grazing and enhance tree establishment. In accordance with current conceptual facilitation-stress models, we hypothesised a positive relationship between facilitation and grazing pressure. We tested this hypothesis in a field experiment in which tree saplings of four different species (deciduous Fagus sylvatica, Acer pseudoplatanus and coniferous Abies alba, Picea abies) were planted either inside or outside of the canopy of the spiny nurse shrub Rosa rubiginosa in enclosures differing in grazing pressure (low and high) and in exclosures. During one grazing season we followed the survival of the different tree saplings and the level of browsing on these; we also estimated browsing damage to the nurse shrubs. Shrub damage was highest at the higher grazing pressure. Correspondingly, browsing increased and survival decreased in saplings located inside the canopy of the shrubs at the high grazing pressure compared to the low grazing pressure. Saplings of both deciduous species showed a higher survival than the evergreens, while sapling browsing did not differ between species. The relative facilitation of sapling browsing and sapling survival – i.e. the difference between saplings inside and outside the shrub canopy – decreased at high grazing pressure as the facilitative species became less protective. Interestingly, these findings do not agree with current conceptual facilitation-stress models predicting increasing facilitation with abiotic stress. We used our results to design a conceptual model of facilitation along a biotic environmental gradient. Empirical studies are needed to test the applicability of this model. In conclusion, we suggest that current conceptual facilitation models should at least consider the possibility of decreasing facilitation at high levels of stress.     

Removal of livestock alters native plant and invasive mammal communities in a dry grassland–shrubland ecosystem

The impacts of domesticated herbivores on ecosystems that did not evolve with mammalian grazing can profoundly influence community composition and trophic interactions. Also, such impacts can occur over long time frames by altering successional vegetation trajectories. Removal of domesticated herbivores to protect native biota can therefore lead to unexpected consequences at multiple trophic levels for native and non-native species. In the eastern South Island of New Zealand large areas of seral grassland–shrubland have had livestock (sheep and cattle) removed following changes in land tenure. The long-term (>10 years) outcomes for these communities are complex and difficult to predict: land may return to a native-dominated woody plant community or be invaded by exotic plants and mammals. We quantified direct and indirect effects of livestock removal on this ecosystem by comparing plant and invasive mammal communities at sites where grazing by livestock ceased c.10–35 years ago (conservation sites) with paired sites where pastoralism has continued to the present (pastoral sites). There was higher total native plant richness and reduced richness of exotic plants on conservation sites compared with pastoral sites. Further, there were differences in the use of conservation and pastoral sites by invasive mammals: rabbits and hedgehogs favoured sites grazed by livestock whereas house mice, brushtail possums and hares favoured conservation sites. Changes in the relative abundance of invasive mammal species after removal of domesticated livestock may compromise positive outcomes for conservation in successional plant communities with no evolutionary history of mammalian grazing.     

A community‐level test of the leaf‐height‐seed ecology strategy scheme in relation to grazing conditions

Question: Is the assumption of trait independence implied in Westoby's (1998) leaf‐height‐seed (LHS) ecology strategy scheme upheld in a Mediterranean grazing system dominated by annuals? Is the LHS approach applicable at the community level? Location: Northern Israel. Methods: LHS traits (specific leaf area [SLA], plant height and seed mass), and additional leaf traits (leaf dry matter content [LDMC], leaf area, and leaf content of nitrogen [LNC], carbon [LCC], and phosphorus [LPC]), were analyzed at the species and community levels. Treatments included manipulations of grazing intensity (moderate and heavy) and protection from grazing. We focused on species comprising 80% of biomass over all treatments, assuming that these species drive trait relationships and ecosystem processes. Results: At the species level, SLA and seed mass were negatively correlated, and plant height was positively correlated to LCC. SLA, seed mass, and LPC increased with protection from grazing. At the community level, redundancy analysis revealed one principal gradient of variation: SLA, correlated to grazing, versus seed mass and plant height, associated with protection from grazing. We divided community functional parameters into two groups according to grazing response: (1) plant height, seed mass, LDMC, and LCC, associated with protection from grazing, and (2) SLA, associated with grazing. Conclusions: The assumption of independence between LHS traits was not upheld at the species level in this Mediterranean grazing system. At the community level, the LHS approach captured most of the variation associated with protection from grazing, reflecting changes in dominance within the plant community.     

Comparing Direct Abiotic Amelioration and Facilitation as Tools for Restoration of Semiarid Grasslands

Desertification can be an irreversible process due to positive feedback among degraded plant and soil dynamics. The recovery of semiarid degraded ecosystems may need human intervention. In restoration practices, the abiotic conditions often need to be improved to overcome the positive plant–soil feedback loops. Using nurse‐plants to improve abiotic conditions for introduced individuals (facilitation) has been suggested as an alternative to direct abiotic amelioration. Here, we compared direct abiotic amelioration and facilitation as tools for restoration of semiarid grasslands in Spain. Seedlings and seeds of Lygeum spartum and Salsola vermiculata were planted and sown in a stably degraded semiarid area in Northeast Spain. Two levels of direct abiotic amelioration (ploughing and damming) and indirect abiotic amelioration through facilitation by Suaeda vera nurse shrubs were compared with a control with no amelioration treatment. The control treatment showed low plant establishment, confirming the practical irreversibility of the degraded state. Plant establishment was significantly higher in the three treatments with interventions than in the control treatment. The best treatment depended on the plant trait considered, but damming was in most cases better than plant facilitation. However, facilitation maintained the nutrient‐rich topsoil layer. Given the relative success of facilitation, revegetation using the facilitative effect of nurse‐plants would, in principle, be recommended for restoring semiarid grasslands. Direct abiotic amelioration would be needed under extreme degradation or harsh climatic conditions.     

Positive effects of shrubs on plant species diversity do not change along a gradient in grazing pressure in an arid shrubland

Facilitative or positive interactions among species are driven mainly by the environmental amelioration or protection from grazing provided by nurse plants. Some studies have suggested that protection from grazing is inconsequential in water-limited environments because of low herbivore densities and their grazing effects. Others, however, argue that herbivores have a major effect on semi-arid plant communities, and that protection from grazing is a significant factor driving positive plant–plant interactions in such environments. We identified a gradient in grazing pressure in a semi-arid shrubland in south-eastern Australia along which we compared soil condition, incident radiation and plant composition beneath two nurse shrub species with open (shrub-free) interspaces. Our aim was to assess the degree of microclimatic amelioration provided by both shrubs, and changes in the interactions (intensity, importance and frequency) between both nurse shrubs and their understorey species, and their effects on species richness at the community level. Both the relative interaction intensity (RII) and interaction importance (I_imp) indices of plant–plant interactions were generally positive and independent of grazing pressure. Soil beneath both nurse plants had significantly greater indices of nutrient cycling and infiltration, and contained more C and N than soil in the open. Almost twice as many species occurred under the canopies of both shrubs (44 species) than in the open (23 species), and the composition of species differed significantly among microsites. Fifty-four percent of all perennial plant species occurred exclusively under shrubs. Our results suggest that environmental amelioration is a stronger driver of the facilitatory effect of shrubs on their understorey species than protection from grazing. Our conclusions are based on the fact that the substantial effect of plant–plant interactions on plant species richness was largely independent of grazing pressure. Irrespective of the underlying mechanism for this effect, our study illustrates the ecological role of shrubs as refugia for understorey plants in semi-arid environments and cautions against management practices aimed at reducing shrub populations.     

Plant traits link hypothesis about resource-use and response to herbivory

Grazing by large herbivores, in interplay with environmental productivity, is a key driver of the composition of the vegetation with important consequences on the ecosystem and, consequently, for land management. We tested the predictions of the resource availability – resource–acquisition theory by assessing the extent to which community averages of plant traits, known to be related to plant growth, competitive ability and response to grazing were correlated with resource gradients within local (200 km²) geographical ranges. Second, we assessed the applicability of the same set of plant traits to make inferences on ecological effects of grazing by sheep in alpine ecosystems in Norway, using a data set consisting of 16 sites in central Norway. We estimated grazing intensity by free-ranging sheep based on GPS telemetry, soil properties, plant species composition and species traits i.e. specific leaf area (SLA), leaf dry matter content (LDMC), leaf size and plant height. Soil fertility and the interaction between soil fertility and grazing, but not grazing intensity alone, were significantly related to plant species and traits composition. Generally, average SLA showed lower correspondence with soil fertility and grazing than the other traits. Leaf size and plant height were lowest at sites with high grazing intensity and in sites with low fertility, and increased with soil fertility in little and moderately grazed sites, but declined at high fertility sites when grazing was intense. LDMC showed the opposite trend. Grazing intensity was more related to the variability in plant composition and average plant traits when environmental productivity was high. Our results therefore are indicative of a convergence of responses to grazing and nutrient limitation.     

Identifying qualitative effects of different grazing types on below‐ground communities and function in a long‐term field experiment

Herbivory is an important modulator of plant biodiversity and productivity in grasslands, but our understanding of herbivore‐induced changes on below‐ground processes and communities is limited. Using a long‐term (17 years) experimental site, we evaluated impacts of rabbit and invertebrate grazers on some soil functions involved in carbon cycling, microbial diversity, structure and functional composition. Both rabbit and invertebrate grazing impacted soil functions and microbial community structure. All functional community measures (functions, biogeochemical cycling genes, network association between different taxa) were more strongly affected by invertebrate grazers than rabbits. Furthermore, our results suggest that exclusion of invertebrate grazers decreases both microbial biomass and abundance of genes associated with key biogeochemical cycles, and could thus have long‐term consequences for ecosystem functions. The mechanism behind these impacts are likely to be driven by both direct effects of grazing altering the pattern of nutrient inputs and by indirect effects through changes in plant species composition. However, we could not entirely discount that the pesticide used to exclude invertebrates may have affected some microbial community measures. Nevertheless, our work illustrates that human activity that affects grazing intensity may affect ecosystem functioning and sustainability, as regulated by multi‐trophic interactions between above‐ and below‐ground communities.     

The interplay of nurse and target plant traits influences magnitude and direction of facilitative interactions under different combinations of stress and disturbance intensities in Andean dry grassland

Aims Facilitation is a key process in vegetation dynamics, driving the response to natural and anthropogenic pressures. In harsh-grazed systems, palatable plants mainly survive when nested under unpalatable tussocks and shrubs. The magnitude and direction of positive interactions are driven by resource availability, extent of herbivory and type of nurse species. We hypothesized that different combinations of disturbance and environmental stress affect community composition in the dry Puna (southern Peruvian Andes) by modifying nurse types and plant interactions in magnitude and specific associations. We investigated whether different combinations of stress and disturbance influence species richness, type and frequency of occurrence of nurse and beneficiary species and magnitude and patterns of plant interactions; whether nurse species influence these interactions and target species change their interactions under different combinations of stress and disturbance and whether plant functional traits differ in the studied communities and influence the pattern of spatial interactions.Methods We selected three plant communities subject to different precipitation and management regimes: in each we laid a number of transects proportional to its extension. Data collected include species presence/absence, type of spatial interactions with nurse species and functional traits. We calculated species richness and rarefaction patterns, described the patterns of plant–plant spatial interactions and investigated the associations between nurse and other species in the three communities using indicator species analysis (ISA). We performed ISA and correlation analysis to investigate whether plant functional traits influenced facilitative interactions.Important findings We found that different combinations of stress and disturbance shaped a complex set of responses, including changes in the nurse species set. Nurse composition influenced magnitude and direction of plant interactions under different stress intensities. Heavy disturbance increased the relative importance of facilitation, even if the overall number of facilitated species decreased. Under equivalent disturbance regimes, increased abiotic stress led to a greater importance of facilitation. Different combinations of stress and disturbance affected the community assemblage also by changing the behaviour of some non-nurse species. Both heavy disturbance and strong stress led to a decrease of trait states; with certain combinations of stress and disturbance, preferential distribution of these states was observed. We also found that plant traits were of key importance in determining facilitative interactions. Some traits were mainly associated with one type of spatial interaction: plant architecture, life cycle and root type influenced the type of interaction between nurses and beneficiaries under different combinations of stress and disturbance. Our results also demonstrate that in plant interaction research the object of observations (species per se, species percentage, etc.) might influence outputs, and to effectively assess the impact of different stress and disturbance intensities on plant interactions it is necessary to work at the community level to consider the whole species pool.     

Direct and indirect effects of episodic frost on plant growth and reproduction in subalpine wildflowers

Frost is an important episodic event that damages plant tissues through the formation of ice crystals at or below freezing temperatures. In montane regions, where climate change is expected to cause earlier snow melt but may not change the last frost‐free day of the year, plants that bud earlier might be directly impacted by frost through damage to flower buds and reproductive structures. However, the indirect effects of frost mediated through changes in plant–pollinator interactions have rarely been explored. We examined the direct and pollinator‐mediated indirect effects of frost on three wildflower species in southwestern Colorado, USA, Delphinium barbeyi (Ranunculaceae), Erigeron speciosus (Asteraceae), and Polemonium foliosissimum (Polemoniaceae), by simulating moderate (?1 to ?5°C) frost events in early spring in plants in situ. Subsequently, we measured plant growth, and upon flowering measured flower morphology and phenology. Throughout the flowering season, we monitored pollinator visitation and collected seeds to measure plant reproduction. We found that frost had species‐specific direct and indirect effects. Frost had direct effects on two of the three species. Frost significantly reduced flower size, total flowers produced, and seed production of Erigeron. Furthermore, frost reduced aboveground plant survival and seed production for Polemonium. However, we found no direct effects of frost on Delphinium. When we considered the indirect impacts of frost mediated through changes in pollinator visitation, one species, Erigeron, incurred indirect, negative effects of frost on plant reproduction through changes in floral traits and pollinator visitation, along with direct effects. Overall, we found that flowering plants exhibited species‐specific direct and pollinator‐mediated indirect responses to frost, thus suggesting that frost may play an important role in affecting plant communities under climate change.     

Comparison of methods for assessing the impact of different disturbances and nutrient conditions upon functional characteristics of grassland communities

Background and Aims

Predicting the response of plant communities to variation in resources and disturbance is still a challenge, because findings depend on how ecological gradients are characterized and how grassland functional composition is assessed. Focusing on leaf dry matter content (LDMC), the efficacy of different methods for evaluating the best response of plant communities to either environmental or disturbance change is examined.

Methods

Data were collected on 69 grasslands located at four sites in the Pyrenees and Massif Central. N-Ellenberg indices and plant nutrient content (Ni) were compared to assess fertility, and either LDMC (meas) measured or calculated from a trait database for which traits were measured under the same environmental conditions (db). Management regime (MR) was characterized in terms of categories (grazing, cutting) and plant height.

Key Results

LDMCdb was positively correlated to LDMCmeas, but depended significantly on site temperature. N-Ellenberg and Ni were significantly correlated, and there was a significant effect of MR and temperature. LDMC responded to fertility, MR and temperature. Replacing MR by plant height in an REML analysis reduced the uncertainty of the LDMC prediction. LDMC was correlated to plant height at community level, whereas the correlation was weak at species level. Differences in LDMC between plant communities under any of the management regimes were significantly correlated to the standing herbage mass.

Conclusion

The N-Ellenberg index is a better indicator of fertility than Ni which is short-term and environment-dependent. LDMC taken from a database allows plant trait variation due to species abundance (excluding variation due to trait plasticity in response to management) to be captured. So the former is better suited for assessing agricultural services that mainly depend on plant phenology and tissue composition. LDMC responded to defoliation regime in addition to fertility because plant height is roughly correlated with LDMC at plant community level.     

Changes in semi-arid plant species associations along a livestock grazing gradient

In semi-arid ecosystems, vegetation is heterogeneously distributed, with plant species often associating in patches. These associations between species are not constant, but depend on the particular response of each species to environmental factors. Here, we investigated how plant species associations change in response to livestock grazing in a semi-arid ecosystem, Cabo de Gata-Níjar Natural Park in South East Spain. We established linear point-intercept transects at four sites with different grazing intensity, and recorded all species at each point. We investigated plant associations by comparing the number of times that each pair of species occurred at the same spatial point (co-occurrences), with the expected number of times based on species abundances. We also assessed associations for each shrub and grass species by considering all their pairs of associations and for the whole plant community by considering all pairs of associations on each site. At all sites, the plant community had a negative pattern of association, with fewer co-occurrences than expected. Negative association in the plant community increased at maximum grazing intensity. Most species associated as expected, particularly grass species, and positive associations were most important at intermediate grazing intensities. No species changed its type of association along the grazing gradient. We conclude that in the present plant community, grazing-resistant species compete among themselves and segregate in space. Some shrub species act as refuges for grazing-sensitive species that benefit from being spatially associated with shrub species, particularly at intermediate grazing intensities where positive associations were highest. At high grazing intensity, these shrubs can no longer persist and positive associations decrease due to the disappearance of refuges. Spatial associations between plant species and their response to grazing help identify the factors that organize plant communities, and may contribute to improving management of semi-arid ecosystems.