Once seen as anomalous, facilitative interactions among plants and their importance for community structure and functioning are now widely recognized. The growing body of modelling, descriptive and experimental studies on facilitation covers a wide variety of terrestrial and aquatic systems throughout the globe. However, the lack of a general body of theory linking facilitation among different types of organisms and biomes and their responses to environmental changes prevents further advances in our knowledge regarding the evolutionary and ecological implications of facilitation in plant communities. Moreover, insights gathered from alternative lines of inquiry may substantially improve our understanding of facilitation, but these have been largely neglected thus far.Despite over 15 years of research and debate on this topic, there is no consensus on the degree to which plant-plant interactions change predictably along environmental gradients (i.e. the stressgradient hypothesis), and this hinders our ability to predict how plant-plant interactions may affect the response of plant communities to ongoing global environmental change. The existing controversies regarding the response of plant-plant interactions across environmental gradients can be reconciled when clearly considering and determining the species-specificity of the response, the functional or individual stress type, and the scale of interest (pairwise interactions or community-level response). Here, we introduce a theoretical framework to do this, supported by multiple lines of empirical evidence. We also discuss current gaps in our knowledge regarding how plant-plant interactions change along environmental gradients. These include the existence of thresholds in the amount of species-specific stress that a benefactor can alleviate, the linearity or non-linearity of the response of pairwise interactions across distance from the ecological optimum of the beneficiary, and the need to explore further how frequent interactions among multiple species are and how they change across different environments. We review the latest advances in these topics and provide new approaches to fill current gaps in our knowledge. We also apply our theoretical framework to advance our knowledge on the evolutionary aspects of plant facilitation, and the relative importance of facilitation, in comparison with other ecological processes, for maintaining ecosystem structure, functioning and dynamics. We build links between these topics and related fields, such as ecological restoration, woody encroachment, invasion ecology, ecological modelling and biodiversity-ecosystem-functioning relationships. By identifying commonalities and insights from alternative lines of research, we further advance our understanding of facilitation and provide testable hypotheses regarding the role of (positive) biotic interactions in the maintenance of biodiversity and the response of ecological communities to ongoing environmental changes. Europe PMC Funders Group

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 Weld experiment in which tree saplings of four diVerent 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 diVering in grazing pressure (low and high) and in exclosures. During one grazing season we followed the survival of the diVerent 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 diVer between species. The relative facilitation of sapling browsing and sapling survival -i.e. the diVerence between saplings inside and outside the shrub canopy -decreased at high grazing pressure as the facilitative species became less protective. Interestingly, these Wndings do not agree with current conceptual facilitationstress 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.

The stability of ecological communities is critical for the stable provisioning of ecosystem services, such as food and forage production, carbon sequestration, and soil fertility. Greater biodiversity is expected to enhance stability across years by decreasing synchrony among species, but the drivers of stability in nature remain poorly resolved. Our analysis of time series from 79 datasets across the world showed that stability was associated more strongly with the degree of synchrony among dominant species than with species richness. The relatively weak influence of species richness is consistent with theory predicting that the effect of richness on stability weakens when synchrony is higher than expected under random fluctuations, which was the case in most communities. Land management, nutrient addition, and climate change treatments had relatively weak and varying effects on stability, modifying how species richness, synchrony, and stability interact. Our results demonstrate the prevalence of biotic drivers on ecosystem stability, with the potential for environmental drivers to alter the intricate relationship among richness, synchrony, and stability.

Summary1. The stress gradient hypothesis (SGH) predicts a shift from net negative interactions in benign environments towards net positive in harsh environments in ecological communities. While several studies found support for the SGH, others found evidence against it, leading to a debate on how nature and strength of species interactions change along stress gradients, and to calls for new empirical and theoretical work. 2. In the latest attempt in this journal, it is successfully argued how the SGH should be expanded by considering different life strategies of species (stress tolerance versus competitive ability) and characteristics of abiotic stress (resource versus non-resource based) over wider stress gradients (opposed to low-high contrasts), but the crucial role of biotic stress by consumers is largely ignored in this refinement. 3. We point out that consumers strongly alter the outcome of species interactions in benign and harsh environments, and show how inclusion of consumer-incurred biotic stress alters the predicted outcome of interactions along resource-and non-resource-based stress gradients for stress-tolerant and competitive benefactors and beneficiaries. 4. Synthesis. New studies should include stress gradients consisting of both abiotic and biotic components to disentangle their impacts, and to improve our understanding of how species interactions change along environmental gradients.

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Question: Insufficient tree regeneration threatens the long‐term persistence of biodiverse Mediterranean open oak woodlands. Could shrubs, scarce due to decades of management (clearing and ploughing), facilitate holm oak recruitment at both acorn and seedling stages? Location: Open oak woodlands in Central Spain. Methods: Plots with four acorns were planted: (1) under the canopy of the spiny shrub Genista hirsuta, (2) in a small cage, protecting against ungulates, (3) in a shaded cage, protecting against ungulates and sun, and (4) in open grassland. Sets of these four treatments were spatially grouped according to a randomised block design, with 16 blocks near (< 10 m) and 16 away from (> 20 m) parent trees to test for distance‐related survival. Plots were regularly checked for seed removal. After emergence one seedling per plot (97 in total) was selected and its survival monitored. Results: Three months after sowing, 199 of 512 acorns were removed, predominantly by rodents. Acorn removal occurred at each treatment but was highest under shrubs. Eight months after sowing, seedling survival was highest under shrubs (50%), followed by shaded cages (16%), open grassland (4%) and cages (0%). Main mortality cause was drought (90%), killing most seedlings between June and July. No seedlings died from ungulate browsing. Conclusion: Shrubs demonstrated clear net facilitative effects for Quercus ilex recruitment, despite higher seed removal. Shading appears the crucial factor facilitating seedling survival. We therefore propose that lack of shrubs contributes largely to tree recruitment failure in Mediterranean open woodlands; management should aim at conserving shrubs.

Summary 1.In endangered wooded pasture ecosystems established tree saplings are frequently found in spatial association with protective structures, suggesting nurse effects. This associational resistance is thought to be a driving force behind tree regeneration in wooded pastures. Experimental evidence for associational resistance is, however, scarce. We studied the effects of unpalatable plants on tree sapling performance in a wooded pasture ecosystem in Switzerland and tested the associational resistance hypothesis experimentally. 2. We transplanted 600 Picea abies saplings of two size classes (5 cm and 10 cm) in plots: (i) with unpalatable plants (either Gentiana lutea or Cirsium acaule , representing a chemically and mechanically defended species, respectively); (ii) with one of both unpalatable plants clipped to the ground; and (iii) without any unpalatable plants. Grazing intensity was measured once in each plot and tree sapling performance was followed for 2 consecutive years. 3. Grazing intensity was significantly higher in plots without unpalatable plants, and significantly higher in plots with Cirsium than with Gentiana . Correspondingly, sapling survival (21·5% and 10·6% after 1 and 2 grazing years, respectively) was significantly higher near unpalatable plants, and significantly higher in plots with Gentiana than with Cirsium . Large tree saplings survived significantly better than small ones, and depended less on the unpalatable plants for survival. 4. Except for saplings planted near Gentiana , sapling removal by cows and horses was the major cause of death, the extent varying between unpalatable species and treatments. 5. The growth in height of the saplings was unaffected by the treatments. 6. Synthesis and applications. This study shows that unpalatable plants can enhance tree regeneration in wooded pastures. Sapling survival was significantly higher near unpalatable plants where grazing intensity was significantly lower. Protection against cattle is particularly important for small saplings. These results have important management implications for the endangered and disappearing wooded pastures in western Europe. Transplanting tree saplings near unpalatable plants could be an alternative reforestation technique in intensively grazed wooded pastures. On the other hand, removal of unpalatable plants and shrubs might prevent or slow down undesired tree encroachment in less intensively grazed areas.

Anti-predator responses by ungulates can be based on habitat features or on the near-imminent threat of predators. In dense forest, cues that ungulates use to assess predation risk likely differ from half-open landscapes, as scent relative to sight is predicted to be more important. We studied, in the Białowieża Primeval Forest (Poland), whether perceived predation risk in red deer (Cervus elaphus) and wild boar (Sus scrofa) is related to habitat visibility or olfactory cues of a predator. We used camera traps in two different set-ups to record undisturbed ungulate behavior and fresh wolf (Canis lupus) scats as olfactory cue. Habitat visibility at fixed locations in deciduous old growth forest affected neither vigilance levels nor visitation rate and cumulative visitation time of both ungulate species. However, red deer showed a more than two-fold increase of vigilance level from 22% of the time present on control plots to 46% on experimental plots containing one wolf scat. Higher vigilance came at the expense of time spent foraging, which decreased from 32% to 12% while exposed to the wolf scat. These behavioral changes were most pronounced during the first week of the experiment but continuous monitoring of the plots suggested that they might last for several weeks. Wild boar did not show behavioral responses indicating higher perceived predation risk. Visitation rate and cumulative visitation time were not affected by the presence of a wolf scat in both ungulate species. The current study showed that perceived predation risk in red deer and wild boar is not related to habitat visibility in a dense forest ecosystem. However, olfactory cues of wolves affected foraging behavior of their preferred prey species red deer. We showed that odor of wolves in an ecologically equivalent dose is sufficient to create fine-scale risk factors for red deer.