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Nurse plants drive the assembly of facilitated communities and commonly promote plant–soil feedbacks, and are thus recognized as key engineers in abiotically stressful ecosystems. The literature neglects; however, the role of the communities which benefit from the presence of the nurse as contributors to soil ecosystem functions. We hypothesized that the nurse and its beneficiaries synergistically enhance essential ecosystem functions mediated by soil microbiota. To track how plant–plant facilitation impacts plant–soil feedbacks, we selected three nurse species in semi-arid mine tailings and defined three microsites (open space, nurse canopy and nurse + facilitated canopy). In each microsite, we quantified 18 abiotic and biotic variables associated with four functions: reduction in climatic stress, reduction in edaphic stress, soil fertility and soil microbial productivity (decomposition and nutrient cycling). Litter biomass increased from open spaces to the microsite beneath the nurses, and further beneath the nurses and their beneficiaries. Litter biomass was a good predictor of both the reduction in climatic stress and increase in edaphic stress (likely owing to metal bioaccumulation). We attributed increments in soil organics and heterotrophic respiration beneath the nurses and their beneficiaries, compared to nurses alone, to biomass effects through increased litter deposition. Variation in fertility and microbial productivity among microsites shaped by the nurses and their facilitated communities was attributed to both diversity and biomass effects. In particular, fertility was promoted beneath phenotypically diverse facilitated communities, as inferred from ten above- and below-ground traits. However, microbial productivity increased at low levels of root biomass likely due to reduced plant–microbe competition for nutrients. Synthesis. Our results show that facilitated plant communities sheltered by nurse species relieve local abiotic stress and promote plant–microbe interactions, both through biomass and biodiversity effects. These observations shift the conception of facilitated species from simple beneficiaries of the nurse's effects to co-drivers of essential ecosystem functions., This work was supported by the I Convocatoria de ayudas de la Fundación BBVA a proyectos de Investigación (project Mintegra) and the Spanish Ministerio de Ciencia, Innovación y Universidades (projects CGL2014‐58333‐P, CGL2017‐89751‐R and MG's Ramón y Cajal contract).

Summary 1. Ecological communities are assembled as complex networks of both positive (i.e. facilitation) and negative (i.e. competition) interactions. In networks established among plant species, many facilitative interactions occurring between a benefactor – a nurse – and a beneficiary – a facilitated seedling - turn into competition over time as the facilitated seedling grows and outcompetes the nurse. The facilitative associations that disappear over time are mainly restricted to closely related taxa, because close relatives tend to share niche requirements and compete more strongly for the same resources. In consequence, the phylogenetic structure of a network might change as positive associations become negative. 2. This study is aimed to characterize how the overall (i.e. nestedness and connectance) and the phylogenetic structure of facilitation networks in semi-arid communities change when facilitation turns into competition and some of the early species associations established by facilitation disappear. 3. We show that the initial facilitation networks retain the overall, but not the phylogenetic, structure. Phylogenetic analyses show that as seedlings, facilitated species tend to associate with the same subset of nurses while, on the contrary, nurses are indifferent to the identity of their facilitated seedlings. But when competition becomes important, closely related nurse species appear associated with only a subset of facilitated species in the community. 4. Synthesis. Temporal rearrangements in the phylogenetic structure of the facilitation networks provide evidence that plant–plant species interactions lead to highly species-specific networks in which the phylogenetic history has a pervasive influence not only on recruitment but also on adult community composition. The use of phylogenetic methods combined with complex network approaches opens the possibility to understand the complexity of ecological interactions occurring in nature as for example those occurring when biological invasions take place or those producing coextinction cascades following species removal from ecosystems.

10 páginas, 5 figuras, 1 tabla., 1. The balance between facilitation and competition through time is at the core of models for successional dynamics. However, since the 1980s, the studies of facilitation have shifted away from successional processes. In a return to the traditional roots of the study of facilitation, we assessed the phylogenetic signatures of competition and facilitation in Mediterranean successional communities and compared them with those recently quantified in non-successional communities of the Mexican desert. 2. Based on previous work, we hypothesized that facilitation between distantly related lineages is an important driver of successional dynamics, as has been shown in non-successional systems. However, we also predicted that the balance between facilitation and competition will be different in successional systems because many species disappear from communities during the process of succession, which does not occur in non-successional systems. 3. We sampled plant species composition over a chronosequence of post-fire succession and predicted phylogenetic clustering in communities at the early stages caused by species that reproduce by seed ('seeders', belonging to a few specific families) being favoured by fire; overdispersion in intermediate stages driven by facilitation interactions among distantly related species; and randomness in the final stage caused by the competitive exclusion of pioneer species belonging to a few families and the survival of species in many other families. 4. In the pioneer stage, we found a random phylogenetic pattern because seeders and many resprouter species were present at this stage, indicating that autosuccession was also occurring. In intermediate phases, once pioneers had recruited into open spaces and facilitated late-successional species, most of which were ancient taxa originating during the Tertiary, phylogenetic overdispersion predominated. Finally, in the later stages competitive exclusion of pioneer species reduced phylogenetic diversity, leading to a random phylogenetic structure. 5.Synthesis. As previously found for non-successional communities, facilitation among distantly related lineages appeared to drive successional dynamics. However, subsequent competition reduced phylogenetic diversity during succession in this Mediterranean system, and some species disappeared from the community., We thank Néctor Siles for field assistance and the Consejería de Medio Ambiente (CMA) and Junta Rectora of Sierra de Cazorla, Segura y Las Villas Natural Park for constant support and facilities. Housing facilities at Sierra deCazorla were provided by Estación Biológica de Doñana (CSIC). J. G. Pausas provided valuable comments on the manuscript. This study was funded by ‘Convenio de asesoramiento y seguimiento de las actuaciones de restauración del incendio del Puerto de las Palomas’ between CMA, EGMASA and University of Jaé (UJA) and projects CGL2008-05289-C02-01/BOS and CGL2006-02848 (Spanish Ministry of Science and Innovation). Projects 409AC0369 (CYTED) and A/017475/08 (AECID) allowed the cooperation between Spanish and Mexican research groups. G.S. was supported by a PhD grant from UJA.

Summary 1. The relative contribution of positive and negative interactions to the organization of ecological communities is an important area, though poorly understood because of the complexity inherent to long-term interactions. For example, positive interactions like plant facilitation turn into negative interactions (competition) along the ontogeny of a plant or in response to temporal fluctuations in the environment. Furthermore, when many plants grow together indirect effects are usually positive and alleviate direct competitive effects. 2. The large number of direct and indirect interactions potentially occurring in natural communities and the temporal scale necessary to account for the ontogenetic shifts in the sign of the interaction makes a fully experimental approach prohibitive. Here, we propose that the phylogenetic distance among co-occurring species is a good proxy to detect competition as a long-term force assembling community composition. Our proposal is based on the observation that closely related species tend to be phenotypically similar and therefore compete for the same niche. 3. We test the relationship between phylogenetic relatedness and the persistence of pairwise (nursefacilitated) interactions occurring between 102 woody species in three Mexican semi-arid communities in order to quantify the balance between competition and facilitation at the community level. 4. Our results indicate that facilitation turns into competition with increasing taxa relatedness. After validating the association between competition and phylogenetic relatedness, we estimate that 57% of the interactions remain with time while 43% become competitive. [Correction added after publication, 4 March 2008: in the preceding sentence, values corrected from 53% and 47%, respectively.] The preponderance of positive interactions may be explained if facilitation is considered as a mutualism in which both species benefit leading to vegetation clumps in communities. 5. Synthesis . We provide a new perspective on the balance between positive and negative interactions based on a phylogenetically structured network of interactions. This approach promises to contribute to our understanding of long standing issues in plant ecology and to reveal new areas of future research by testing the existence and the nature of the mutualisms as well as their complexity‐ stability properties on communities as a whole.