Importance, but not intensity of plant interactions relates to species diversity under the interplay of stress and disturbance

The lack of clarity on how the intensity and importance of plant interactions change under the co-occurrence of stress and disturbance strongly impedes assessing the relative importance of plant interactions for species diversity. We addressed this issue in subalpine grasslands of the French Pyrenees. A natural soil moisture gradient further experimentally stretched at both ends was used and a mowing disturbance treatment was applied at each position along the soil moisture gradient. Changes in intensity and importance of plant interactions were assessed by a neighbour removal experiment using four target ecotypes. A structural equation modelling approach was used to assess the relative impact of stress, disturbance, the intensity and importance of plant interactions on diversity at both the neighbourhood and community scales. Without mowing, changes in intensity and importance of plant interactions only diverged in the dry part of the soil moisture gradient. The intensity of plant interactions linearly shifted from competition to facilitation with increasing stress, while the importance followed a hump-shaped relationship. Species diversity components were tightly related to the importance of plant interactions only, both the neighbourhood and community scales. Mowing disturbance strongly reduced the importance of facilitation along the soil moisture gradient, and suppressed the relationship between the importance of plant interactions and diversity components. Together, our results highlight that 1) the importance is the best predictor of variations in species diversity in this subalpine herbaceous system, and 2) that fine-scale processes such as plant interactions can affect the entire plant communities. Finally, our results suggest that high level of constraints due to co-occurring stress and disturbance can inhibit the effects of plant interactions on species diversity, highlighting their potential role in regulating diversity and the maintenance/extinction of plant communities. The co-occurrence of stress (i.e. factors such as drought limiting plant growth, sensu Grime 1973) and disturbance (drastic events such as mowing removing plant biomass) can lead to a rapid loss of diversity. Co-occurring negative effects of stress and disturbance on diversity and ecosystem functioning are specific to severe environments such as alpine grasslands or dry steppes OIKOS How plant interactions change along environmental gradients is an unsolved debate, particularly when both stress and disturbance interact. This lack of clarity explains why the relative impact of plant interactions (intensity and importance) on species diversity has been rarely assessed. Using an experimental approach, we found that the importance of plant interactions highly contributed to variation in species diversity, confirming that neighbourhood scale processes such as plant interactions can affect the entire plant communities. The co-occurrence of stress and disturbance inhibited the effects of plant interactions, highlighting that plant interactions may regulate drops of diversity and the maintenance/ extinction of plant communities. Synthesi

Phylogenetic, functional, and taxonomic richness have both positive and negative effects on ecosystem multifunctionality

Biodiversity encompasses multiple attributes such as the richness and abundance of species (taxonomic diversity), the presence of different evolutionary lineages (phylogenetic diversity), and the variety of growth forms and resource use strategies (functional diversity). These biodiversity attributes do not necessarily relate to each other and may have contrasting effects on ecosystem functioning. However, how they simultaneously influence the provision of multiple ecosystem functions related to carbon, nitrogen, and phosphorus cycling (multifunctionality) remains unknown. We evaluated the effects of the taxonomic, phylogenetic, and functional attributes of dominant (mass ratio effects) and subordinate (richness effect) plant species on the multifunctionality of 123 drylands from six continents. Our results highlight the importance of the phylogenetic and functional attributes of subordinate species as key drivers of multifunctionality. In addition to a higher taxonomic richness, we found that simultaneously increasing the richness of early diverging lineages and the functional redundancy between species increased multifunctionality. In contrast, the richness of most recent evolutionary lineages and the functional and phylogenetic attributes of dominant plant species (mass ratio effects) were weakly correlated with multifunctionality. However, they were important drivers of individual nutrient cycles. By identifying which biodiversity attributes contribute the most to multifunctionality, our results can guide restoration efforts aiming to maximize either multifunctionality or particular nutrient cycles, a critical step to combat dryland desertification worldwide.This work was funded by the European Research Council [ERC Grant Agreements 242658 (BIOCOM) and 647038 (BIODESERT)]. F.T.M., M.B., and Y.L.B.-P. are supported by the ERC (BIODESERT). Y.L.B.-P. was also supported by a Marie Sklodowska-Curie Actions Individual Fellowship within the European Program Horizon 2020 (DRYFUN Project 656035). S.S. was supported by the Spanish Government under a Ramón y Cajal Contract (RYC-2016-20604). N.G. was supported by the AgreenSkills+fellowship programme, which has received funding from the European Union’s Seventh Framework Programme under Grant Agreement FP7-609398 (AgreenSkills+ contract). This work was supported by the French government Initiatives d’Excellence–Initiatives Science/Innovation/Territoires/Économie (IDEX-ISITE) initiative 16-IDEX-0001 (CAP 20-25)

The BIODESERT survey: assessing the impacts of grazing on the structure and functioning of global drylands

Grazing by domestic livestock is both the main land use across drylands worldwide and a major desertification and global change driver. The ecological consequences of this key human activity have been studied for decades, and there is a wealth of information on its impacts on biodiversity and ecosystem processes. However, most field assessments of the ecological impacts of grazing on drylands conducted to date have been carried out at local or regional scales and have focused on single ecosystem attributes (e.g., plant productivity) or particular taxa (mainly aboveground, e.g., plants). Here we introduce the BIODESERT survey, the first systematic field survey devoted to evaluating the joint impacts of grazing by domestic livestock and climate on the structure and functioning of dryland ecosystems worldwide. This collaborative global survey was carried out between 2016 and 2019 and has involved the collection of field data and plant, biocrust, and soil samples from a total of 326 45 m × 45 m plots from 98 sites located in 25 countries from 6 continents. Here we describe the major characteristics and the field protocols used in this survey. We also introduce the organizational aspects followed, as these can be helpful to everyone wishing to establish a global collaborative network of researchers. The BIODESERT survey provides baseline data to assess the current status of dryland rangelands worldwide and the impacts of grazing on these key ecosystems, and it constitutes a good example of the power of collaborative research networks to study the ecology of our planet using much-needed field data.This research has been supported by the European Research Council (ERC grant agreement no. 647038 – BIODESERT) and the Conselleria de Innovación, Universidades, Ciencia y Sociedad Digital, Generalitat Valenciana (grant no. CIDEGENT/2018/041). Nicolas Gross was supported by CAP 20-25 (16-IDEX-0001) and the AgreenSkills+ fellowship program which has received funding from the EU's Seventh Framework Programme under grant agreement no. 996 FP7-609398 (AgreenSkills+ contract). Hugo Saiz is supported by a María Zambrano fellowship funded by the Ministry of Universities and European Union Next Generation plan

Functional rarity and evenness are key facets of biodiversity to boost multifunctionality

The functional traits of organisms within multispecies assemblages regulate biodiversity effects on ecosystem functioning. Yet how traits should assemble to boost multiple ecosystem functions simultaneously (multifunctionality) remains poorly explored. In a multibiome litter experiment covering most of the global variation in leaf trait spectra, we showed that three dimensions of functional diversity (dispersion, rarity, and evenness) explained up to 66% of variations in multifunctionality, although the dominant species and their traits remained an important predictor. While high dispersion impeded multifunctionality, increasing the evenness among functionally dissimilar species was a key dimension to promote higher multifunctionality and to reduce the abundance of plant pathogens. Because too-dissimilar species could have negative effects on ecosystems, our results highlight the need for not only diverse but also functionally even assemblages to promote multifunctionality. The effect of functionally rare species strongly shifted from positive to negative depending on their trait differences with the dominant species. Simultaneously managing the dispersion, evenness, and rarity in multispecies assemblages could be used to design assemblages aimed at maximizing multifunctionality independently of the biome, the identity of dominant species, or the range of trait values considered. Functional evenness and rarity offer promise to improve the management of terrestrial ecosystems and to limit plant disease risks.This work was funded by the British Ecological Society (SR17\1297 grant, PI: P.G.-P.) and by the European Research Council (ERC Grant Agreement #647038, BIODESERT, PI: F.T.M.). Y.L.B.-P. was supported by a Marie Sklodowska-Curie Actions Individual Fellowship within the European Program Horizon 2020 (DRYFUN Project #656035). H.S. was supported by a Juan de la Cierva-Formación grant from the Spanish Ministry of Economy and Competitiveness (FJCI-2015-26782). F.T.M. and S.A. were supported from the Generalitat Valenciana (CIDEGENT/2018/041). M.D. was supported by a Formación del Profesorado Universitario (FPU) fellowship from the Spanish Ministry of Education, Culture and Sports (FPU-15/00392). S.A. was supported by the Spanish MINECO for financial support via the DIGGING_DEEPER project through the 2015 to 2016 BiodivERsA3/FACCE‐JPI joint call for research proposals. B.K.S. research on biodiversity-ecosystem functions was supported by the Australian Research Council (DP170104634 and DP190103714). P.G.-P. was supported by a Ramón y Cajal grant from the Spanish Ministry of Science and Innovation (RYC2018-024766-I). R.M. was supported by MINECO (Grants CGL2014-56567-R and CGL2017-83855-R)

Soil fungal abundance and plant functional traits drive fertile island formation in global drylands

Dryland vegetation is characterized by discrete plant patches that accumulate and capture soil resources under their canopies. These “fertile islands” are major drivers of dryland ecosystem structure and functioning, yet we lack an integrated understanding of the factors controlling their magnitude and variability at the global scale.EEA BarilocheFil: Ochoa-Hueso, Raúl. Universidad Autónoma de Madrid. Department of Ecology; EspañaFil: Eldridge, David J. University of New South Wales. School of Biological, Earth and Environmental Sciences; AustraliaFil: Delgado-Baquerizo, Manuel. University of Colorado. Cooperative Institute for Research in Environmental Sciences; Estados Unidos. Universidad Rey Juan Carlos. Escuela Superior de Ciencias Experimentales y Tecnología. Departamento de Biología y Geología, Física y Química Inorgánica; EspañaFil: Soliveres, Santiago. University of Bern. Institute of Plant Sciences; SuizaFil: Bowker, Matthew A. Northern Arizona University. School of Forestry; Estados UnidosFil: Gross, Nicolás. Universidad Rey Juan Carlos. Escuela Superior de Ciencias Experimentales y Tecnología. Departamento de Biología y Geología, Física y Química Inorgánica; España. Institut Nationale de la Recherche Agronomique; Francia. Université La Rochelle. Centre d’étude biologique de Chizé; FranciaFil: Le Bagousse-Pinguet, Yoann. Universidad Rey Juan Carlos. Escuela Superior de Ciencias Experimentales y Tecnología. Departamento de Biología y Geología, Física y Química Inorgánica; EspañaFil: Quero, José L. Universidad de Córdoba. Escuela Técnica Superior de Ingeniería Agronómica y de Montes. Departamento de Ingeniería Forestal: EspañaFil: García-Gómez, Miguel. Universidad Rey Juan Carlos. Escuela Superior de Ciencias Experimentales y Tecnología. Departamento de Biología y Geología, Física y Química Inorgánica; EspañaFil: Valencia, Enrique. Universidad Rey Juan Carlos. Escuela Superior de Ciencias Experimentales y Tecnología. Departamento de Biología y Geología, Física y Química Inorgánica; EspañaFil: Arredondo, Tulio. Instituto Potosino de Investigación Científica y Tecnológica. División de Ciencias Ambientales; MéxicoFil: Beinticinco, Laura. Universidad Nacional de La Pampa. Facultad de Agronomía; ArgentinaFil: Bran, Donaldo Eduardo. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Bariloche; ArgentinaFil: Cea, Alex. Universidad de La Serena. Departamento de Biología; ChileFil: Coaguila, Daniel. Instituto de Ensino Superior de Rio Verde; BrasilFil: Dougill, Andrew J. University of Leeds. School of Earth and Environment; Gran BretañaFil: Espinosa, Carlos I. Universidad Técnica Particular de Loja. Departamento de Ciencias Naturales; EcuadorFil: Gaitan, Juan Jose. Instituto Nacional de Tecnología Agropecuaria (INTA). Instituto de Suelos; ArgentinaFil: Guuroh, Reginald T. University of Cologne. Botanical Institute. Range Ecology and Range Management Group; Alemania. CSIR-Forestry Research Institute of Ghana; GhanaFil: Guzmán, Elizabeth. Universidad Técnica Particular de Loja. Departamento de Ciencias Naturales; EcuadorFil: Gutiérrez, Julio R.. Universidad de La Serena. Departamento de Biología; Chile. Centro de Estudios Avanzados en Zonas Áridas (CEAZA); Chile. Instituto de Ecología y Biodiversidad; ChileFil: Hernández, Rosa M. Universidad Experimental Simón Rodríguez. Centro de Agroecología Tropical. Laboratorio de Biogeoquímica; VenezuelaFil: Huber-Sannwald, Elisabeth. Instituto Potosino de Investigación Científica y Tecnológica. División de Ciencias Ambientales; MéxicoFil: Jeffries, Thomas. Western Sydney University. Hawkesbury Institute for the Environment; AustraliaFil: Linstädter, Anja. University of Cologne. Botanical Institute. Range Ecology and Range Management Group; AlemaniaFil: Mau, Rebecca L. Northern Arizona University. Center for Ecosystem Science and Society: Estados UnidosFil: Monerris, Jorge. Université du Québec à Montréal. Pavillon des Sciences Biologiques. Département des Sciences Biologiques; CanadáFil: Prina, Anibal. Universidad Nacional de La Pampa. Facultad de Agronomía; ArgentinaFil: Pucheta, Eduardo. Universidad Nacional de San Juan. Facultad de Ciencias Exactas, Físicas y Naturales. Departamento de Biología; ArgentinaFil: Stavi, Ilan. Dead Sea and Arava Science Center, IsraelFil: Thomas, Andrew. Aberystwyth University. Department of Geography and Earth Sciences; Gran BretañaFil: Zaady, Eli. Agricultural Research Organization. Gilat Research Center. Natural Resources; IsraelFil: Singh, Brajesh K. Western Sydney University. Hawkesbury Institute for the Environment; Australia. Western Sydney University. Global Centre for Land-Based Innovation; AustraliaFil: Maestre, Fernando T. Universidad Rey Juan Carlos. Escuela Superior de Ciencias Experimentales y Tecnología. Departamento de Biología y Geología, Física y Química Inorgánica; Españ

Soil fungal abundance and plant functional traits drive fertile island formation in global drylands

International audience1.Dryland vegetation is characterised by discrete plant patches that accumulate and capture soil resources under their canopies. These “fertile islands” are major drivers of dryland ecosystem structure and functioning, yet we lack an integrated understanding of the factors controlling their magnitude and variability at the global scale.2.We conducted a standardized field survey across two hundred and thirty-six drylands from five continents. At each site, we measured the composition, diversity and cover of perennial plants. Fertile island effects were estimated at each site by comparing composite soil samples obtained under the canopy of the dominant plants and in open areas devoid of perennial vegetation. For each sample, we measured fifteen soil variables (functions) associated with carbon, nitrogen and phosphorus cycling and used the Relative Interaction Index to quantify the magnitude of the fertile island effect for each function. In eighty sites, we also measured fungal and bacterial abundance (quantitative PCR) and diversity (Illumina MiSeq).3.The most fertile islands, i.e. those where a higher number of functions were simultaneously enhanced, were found at lower-elevation sites with greater soil pH values and sand content under semiarid climates, particularly at locations where the presence of tall woody species with a low specific leaf area increased fungal abundance beneath plant canopies, the main direct biotic controller of the fertile island effect in the drylands studied. Positive effects of fungal abundance were particularly associated with greater nutrient contents and microbial activity (soil extracellular enzymes) under plant canopies.4.Synthesis. Our results show that the formation of fertile islands in global drylands largely depends on: (i) local climatic, topographic and edaphic characteristics, (ii) the structure and traits of local plant communities and (iii) soil microbial communities. Our study also has broad implications for the management and restoration of dryland ecosystems worldwide, where woody plants are commonly used as nurse plants to enhance the establishment and survival of beneficiary species. Finally, our results suggest that forecasted increases in aridity may enhance the formation of fertile islands in drylands worldwide

Indirekten positiven Interaktionen als ein potentieller strukturierender Mechanismus für Pflanzengemeinschaften (Makrophyten)

Untersuchungen zu Interaktionen zwischen Pflanzen sind fundamental wichtig für das Verständnis der Prozesse, die die Struktur und Funktion von Pflanzengemeinschaften und Ökosystemen steuern. Das Auftreten von positiven Interaktionen ("Facilitation") sowie der Einfluss, den die Antwort der Pflanzengemeinschaft auf Veränderungen in der Umwelt haben ist aus terrestrischen und marinen Ökosystemen bekannt, allerdings ist nur wenig darüber bekannt, wie sie submerse Pflanzengemeinschaften (Makrophyten) steuern. Aufgrund ihrer strukturierenden Funktion in Süßwasser-Ökosystemen (z.B. bei der Sedimentation, der Nährstoffdynamik oder der Stabilität von eutrophierten Ökosystemen) ist das Verständnis, wie positive Interaktionen diese Gemeinschaften beeinflussen von grundlegender Bedeutung. In meiner Dissertation untersuchte ich als erstes im Rahmen eines Review-Artikels das Zusammenspiel von direkten und indirekten positiven Interaktionen entlang von Umwelt- und Grazing-Gradienten bzw. auf die Struktur und Stabilität von Pflanzengemeinschaften in vielen verschiedenen Ökosystemen. Im Folgenden untersuchte ich experimentell die Funktion von indirekten positiven Interaktionen als ein potentieller Mechanismus zur Regulierung von Makrophyten-Gemeinschaften in drei verschiedenen Zusammenhängen: Ich konzentrierte mich auf 1) den potentiellen indirekten positiven Effekt von Fischen auf Makrophyten durch Verminderung der Herbivoren (indirekte positive Interaktion über zwei trophische Ebenen), 2) die mögliche indirekte positive Interaktion zwischen Makrophyten bei steigendem Fraßdruck und 3) mit steigender Eutrophierung. Als Fischart, Herbivore und Makrophyten wählte ich für diese Untersuchungen dominante und weit verbreitete Arten.Der Review-Artikel legt die Vermutung nahe, dass indirekte und direkte positive Interaktionen insgesamt ein Maximum in Umgebungen mit mittlerem Stress und Störungen verursachen, was unter diesen Umweltbedingungen zu einer sehr diversen Pflanzengesellschaft führt. Ein Zusammenbruch der positiven Interaktionen in Gemeinschaften mit hohem Stress und vielen Störungen wiederum kann auf verminderte Schutzwirkung der Nachbarn zurückgeführt werden. Dies kann zu einer lokalen Abnahme der Biodiversität, zum Aussterben von Pflanzengemeinschaften und zu Veränderungen im Zustand des Ökosystems führen. Die experimentelle Herangehensweise wiederum hebt das Auftreten von indirekten positiven Effekten in Süßwasser-Ökosystemen, welche den Zustand von Makrophyten (Überleben und Biomasse) positiv beeinflussten, hervor. Die Anwesenheit von Arten mit positivem Einfluss (benefactors) wie Fischen (höhere trophische Ebene) oder Makrophyten (gleiche trophische Ebene) fängt direkte negative Interaktionen (z.B. Pflanzen-Herbivoren oder Pflanzen-Phytoplankton Interaktionen) auf.Die Erhaltung des Überlebens- und Wachstums-Erfolgs der Pflanzen durch positive Effekte der Nachbarn zeigt die Bedeutung der indirekten positiven Interaktionen für aquatische Pflanzengemeinschaften und wie sie negative biotische (z.B. Fraßdruck) und abiotische Faktoren (z.B. erhöhte Eutrophierung) auffangen können. Darüber hinaus zeigte ich auch, dass der schützende Effekt durch die Nachbarpflanzen unter starken Fraßdruck abnehmen kann und durch den Zusammenbruch der indirekten positiven Interaktionen einen Erfolg der Pflanzen vermindern kann. Die Folgen eines solchen Kollapses sind von großer Bedeutung für die Reaktion der Pflanzengemeinschaften und Ökosysteme auf Umweltveränderungen. Eine Abnahme des schützenden Effekts der Nachbarpflanzen könnte zu einem Zusammenbruch der gesamten Süßwasser-Vegetation führen, wie es aus anderen Ökosystemen bekannt ist. Demzufolge könnte dies in Süßwasser-Ökosystemen teilweise den Verlust der Klarwasser-Phase, d.h. den Wechsel zu einem trüben, von Phytoplankton dominierten Stadium, erklären

TRY plant trait database – enhanced coverage and open access

Plant traits - the morphological, anatomical, physiological, biochemical and phenological characteristics of plants - determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait‐based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits - almost complete coverage for ‘plant growth form’. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait–environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives

Hotspots of biogeochemical activity linked to aridity and plant traits across global drylands

14 páginas.- 4 figuras.- 67 referencias.- The online version contains supplementary material available at https://doi.org/10.1038/s41477-024-01670-7Perennial plants create productive and biodiverse hotspots, known as fertile islands, beneath their canopies. These hotspots largely determine the structure and functioning of drylands worldwide. Despite their ubiquity, the factors controlling fertile islands under conditions of contrasting grazing by livestock, the most prevalent land use in drylands, remain virtually unknown. Here we evaluated the relative importance of grazing pressure and herbivore type, climate and plant functional traits on 24 soil physical and chemical attributes that represent proxies of key ecosystem services related to decomposition, soil fertility, and soil and water conservation. To do this, we conducted a standardized global survey of 288 plots at 88 sites in 25 countries worldwide. We show that aridity and plant traits are the major factors associated with the magnitude of plant effects on fertile islands in grazed drylands worldwide. Grazing pressure had little influence on the capacity of plants to support fertile islands. Taller and wider shrubs and grasses supported stronger island effects. Stable and functional soils tended to be linked to species-rich sites with taller plants. Together, our findings dispel the notion that grazing pressure or herbivore type are linked to the formation or intensification of fertile islands in drylands. Rather, our study suggests that changes in aridity, and processes that alter island identity and therefore plant traits, will have marked effects on how perennial plants support and maintain the functioning of drylands in a more arid and grazed world.This research was supported by the European Research Council (ERC grant 647038 (BIODESERT) awarded to F.T.M.) and Generalitat Valenciana (CIDEGENT/2018/041). D.J.E. was supported by the Hermon Slade Foundation (HSF21040). J. Ding was supported by the National Natural Science Foundation of China Project (41991232) and the Fundamental Research Funds for the Central Universities of China. M.D.-B. acknowledges support from TED2021-130908B-C41/AEI/10.13039/501100011033/Unión Europea Next Generation EU/PRTR and the Spanish Ministry of Science and Innovation for the I + D + i project PID2020-115813RA-I00 funded by MCIN/AEI/10.13039/501100011033. O.S. was supported by US National Science Foundation (Grants DEB 1754106, 20-25166), and Y.L.B.-P. by a Marie Sklodowska-Curie Actions Individual Fellowship (MSCA-1018 IF) within the European Program Horizon 2020 (DRYFUN Project 656035). K.G. and N.B. acknowledge support from the German Federal Ministry of Education and Research (BMBF) SPACES projects OPTIMASS (FKZ: 01LL1302A) and ORYCS (FKZ: FKZ01LL1804A). B.B. was supported by the Taylor Family-Asia Foundation Endowed Chair in Ecology and Conservation Biology, and M. Bowker by funding from the School of Forestry, Northern Arizona University. C.B. acknowledges funding from the National Natural Science Foundation of China (41971131). D.B. acknowledges support from the Hungarian Research, Development and Innovation Office (NKFI KKP 144096), and A. Fajardo support from ANID PIA/BASAL FB 210006 and the Millennium Science Initiative Program NCN2021-050. M.F. and H.E. received funding from Ferdowsi University of Mashhad (grant 39843). A.N. and M.K. acknowledge support from FCT (CEECIND/02453/2018/CP1534/CT0001, SFRH/BD/130274/2017, PTDC/ASP-SIL/7743/2020, UIDB/00329/2020), EEA (10/CALL#5), AdaptForGrazing (PRR-C05-i03-I-000035) and LTsER Montado platform (LTER_EU_PT_001) grants. O.V. acknowledges support from the Hungarian Research, Development and Innovation Office (NKFI KKP 144096). L.W. was supported by the US National Science Foundation (EAR 1554894). Y.Z. and X.Z. were supported by the National Natural Science Foundation of China (U2003214). H.S. is supported by a María Zambrano fellowship funded by the Ministry of Universities and European Union-Next Generation plan. The use of any trade, firm or product names does not imply endorsement by any agency, institution or government. Finally, we thank the many people who assisted with field work and the landowners, corporations and national bodies that allowed us access to their land.Peer reviewe

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