Competitive Species Research Articles

Competition for water and species coexistence in phenologically structured annual plant communities.

Both competition for water and phenological variation are important determinants of plant community structure, but ecologists lack a synthetic theory for how they affect coexistence outcomes. We developed an analytically tractable model of water competition for Mediterranean annual communities and demonstrated that variation in phenology alone can maintain high diversity in spatially homogenous assemblages of water-limited plants. We modelled a system where all water arrives early in the season and species vary in their ability to grow under drying conditions. As a consequence, species differ in growing season length and compete by shortening the growing season of their competitors. This model replicates and offers mechanistic explanations for patterns observed in empirical studies of how phenology influences coexistence among Mediterranean annuals. Additionally, we found that a decreasing, concave-up trade-off between growth rate and access to water can maintain high diversity under simple but realistic assumptions. High diversity is possible because: (1) later plants escape competition after their earlier season competitors have gone to seed and (2) early-season species are more than compensated for their shortened growing season by a growth rate advantage. Together, these mechanisms provide an explanation for how phenologically variable annual plant species might coexist when competing only for water.

A simple “turn-on” fluorescent probe capable of recognition cysteine with rapid response and high sensing in living cells and zebrafish

Cysteine (Cys), an essential biological amino acid, participates several crucial functions in various physiological and pathological processes. The sensitive and specific detection of Cys is of great significance for understanding its biological function to disease diagnosis. Herein, we designed and synthesized a simple fluorescence sensor 2-(benzothiophen-2-yl)-4-oxo-4H-chromen-3-yl acrylate (BTCA) composed of a flavonol skeleton as the fluorophore and acrylic ester group as the recognition receptor. Probe BTCA displayed high selectivity and extremely fast response toward Cys in phosphate buffer solution in the presence of other competitive species even Homocysteine (Hcy) and Glutathione (GSH) owing to a specific conjugate addition-cyclization reaction between the acrylate moiety and Cys. The photoluminescence mechanism of probe BTCA toward Cys was modulated by excited state intramolecular proton transfer (ESIPT) process. The sensing property for Cys was studied by UV–Visible, fluorescence spectrophotometric analyses and time-dependent density functional theory (TD-DFT) calculations, those results indicated that probe BTCA possessed excellent sensitivity, higher specificity, dramatically “naked-eye” fluorescence enhancement (30-fold), high anti-interference ability, especially immediate response speed (within 40 s). Additionally, the practicability of sensor BTCA in exogenous and endogenous Cys imaging in living cells and zebrafish was elucidated as well, suggesting that it has remarkedly diagnostic significance in physiological and pathological process.

Long‐term and year‐to‐year stability and its drivers in a Mediterranean grassland

Abstract Understanding the mechanisms underlying community stability has become an urgent need to protect ecosystems from global change and resulting biodiversity loss. While community stability can be influenced by species richness, synchrony in annual fluctuations of species, species stability and functional traits, the relative contributions of these drivers to stability are still unclear. In semi‐natural grasslands, land‐use changes such as fertilization might affect stability by decreasing richness and influencing year‐to‐year fluctuations. In addition, they can promote long‐term directional trends, shifting community composition and influencing grassland maintenance. Thus, it is important to consider how species and community stability vary year‐to‐year but also in the long term. Using a 14‐year vegetation time series of a species‐rich semi‐natural Mediterranean grassland, we studied the relative importance of richness, synchrony, species stability and functional traits on community stability. To assess land‐use change effects on stability, we applied a fertilization treatment. To distinguish stability patterns produced by year‐to‐year fluctuations from those caused by long‐term trends, we compared the results obtained using a detrending approach from those without detrending. Independently of the treatment and approach applied, the most stable communities were those composed of asynchronous species with low specific leaf area. Fertilization decreased year‐to‐year and long‐term community stability by increasing community‐weighted mean of specific leaf area, decreasing species stability or also reducing richness in the case of year‐to‐year stability. Additionally, traits such as seed mass had an indirect effect on stability through synchrony. Long‐term trends appeared in control and fertilized plots (due to fertilization), decreasing community and species stability and leading to differences in the relationships found between community stability and some of its drivers. This reflects the importance of accounting for the effect of temporal trends on community and species stability using both a long‐term and a year‐to‐year approach. Synthesis. Stability is influenced by richness, synchrony and functional traits. Fertilization decreases species and community stability by promoting long‐term trends in species composition, favouring competitive species and decreasing richness. Studying stability at the community level and species level, and accounting for the effect of trends is essential to understand stability and its drivers more comprehensively.

The core root microbiome of Spartina alterniflora is predominated by sulfur-oxidizing and sulfate-reducing bacteria in Georgia salt marshes, USA

BackgroundSalt marshes are dominated by the smooth cordgrass Spartina alterniflora on the US Atlantic and Gulf of Mexico coastlines. Although soil microorganisms are well known to mediate important biogeochemical cycles in salt marshes, little is known about the role of root microbiomes in supporting the health and productivity of marsh plant hosts. Leveraging in situ gradients in aboveground plant biomass as a natural laboratory, we investigated the relationships between S. alterniflora primary productivity, sediment redox potential, and the physiological ecology of bulk sediment, rhizosphere, and root microbial communities at two Georgia barrier islands over two growing seasons.ResultsA marked decrease in prokaryotic alpha diversity with high abundance and increased phylogenetic dispersion was found in the S. alterniflora root microbiome. Significantly higher rates of enzymatic organic matter decomposition, as well as the relative abundances of putative sulfur (S)-oxidizing, sulfate-reducing, and nitrifying prokaryotes correlated with plant productivity. Moreover, these functional guilds were overrepresented in the S. alterniflora rhizosphere and root core microbiomes. Core microbiome bacteria from the Candidatus Thiodiazotropha genus, with the metabolic potential to couple S oxidation with C and N fixation, were shown to be highly abundant in the root and rhizosphere of S. alterniflora.ConclusionsThe S. alterniflora root microbiome is dominated by highly active and competitive species taking advantage of available carbon substrates in the oxidized root zone. Two microbially mediated mechanisms are proposed to stimulate S. alterniflora primary productivity: (i) enhanced microbial activity replenishes nutrients and terminal electron acceptors in higher biomass stands, and (ii) coupling of chemolithotrophic S oxidation with carbon (C) and nitrogen (N) fixation by root- and rhizosphere-associated prokaryotes detoxifies sulfide in the root zone while potentially transferring fixed C and N to the host plant.5gNBGD5xoDXmzcUFa4HG9gVideo

Resource recovery and freshwater ecosystem restoration — Prospecting for phytoremediation potential in wild macrophyte stands

The primary aim of this study was to understand the factors, e.g., harvest frequency and plant community type, that can facilitate optimising phytoextraction in wild macrophyte communities as part of a strategy for water-quality improvement and resource recovery. This was achieved by surveying wild macrophyte communities and quantifying standing stocks of key nutrient pollutants such as N and P, and a range of other recoverable macro and micro-nutrients (Ca, K, Mg, Cu, Fe, Mn, Mo, Zn, Na and Cr). By scaling-up the pollutant export potential over a decade, it was determined which harvest strategy and plant community types provide the greatest levels of nutrient export. Grime’s CSR plant strategy framework was used to categorise each surveyed community, where large-statured, higher biomass producing competitor and stress tolerator-type communities were compared with ruderal-type communities that have rapid growth and high nutrient acquisition but smaller standing biomass and statures. High biomass plant communities containing competitor or stress tolerator species, produce greater standing stocks of macronutrients (such as N and P) for harvesting, while yields of micronutrient-type pollutants are more likely to be influenced by specific physiological traits that determine leaf tissue concentration. Utilising a high frequency harvest regime over a multi-year time scale suggested that small fast-growing ruderals could yield 4–6 times the concentration of macronutrients and micronutrients for export compared to competitor or stress tolerator-dominated communities e.g., P yields from ruderals were 25 g/m2 versus 5 g/m2 from competitor/stress tolerator-dominated communities. These results emphasise the need to consider both the plant community and the harvesting regime when using phytoextraction as a management tool. We anticipate that these results will help guide environmental managers in their approach in developing circular economy schemes that improve water quality through nutrient export.

An experimental assessment of competitive interactions between sexual and apomictic fern gametophytes using Easy Leaf Area.

PremiseFew studies have explored competition in fern gametophyte populations. One limiting factor is the tedious measurement of gametophyte size as a proxy for biomass in these small plants. Here, an alternative approach of estimating the number of green pixels from photos was employed to measure the competitive interactions among apomictic and sexual Dryopteris gametophytes.MethodsWe cultivated the gametophytes of two apomictic (diploid and triploid) and one sexual (tetraploid) Dryopteris species in monocultures and in two‐species mixtures in the ratios 1 : 1 and 1 : 3. The total gametophyte cover of each population originating from 20 spores was assessed using Easy Leaf Area. Assessments were performed weekly between weeks 4 and 10 of cultivation. Additionally, during week 5, the cover of each species in each mixture was estimated separately.ResultsWe identified a positive correlation between gametophyte size and ploidy level as well as sexual reproduction. The performance of the tested species in mixtures was dependent on the competitor species identity, indicating the importance of competition between gametophytes.DiscussionThe methods outlined can be used for a rapid assessment of fern gametophyte cover in large gametophyte populations. Ploidy level and reproduction type seem to play a major role in the competitive abilities of fern gametophytes, but more research is needed on this topic.

Ecological impacts of photosynthetic light harvesting in changing aquatic environments: A systematic literature map.

Underwater light is spatially as well as temporally variable and directly affects phytoplankton growth and competition. Here we systematically (following the guidelines of PRISMA‐EcoEvo) searched and screened the published literature resulting in 640 individual articles. We mapped the conducted research for the objectives of (1) phytoplankton fundamental responses to light, (2) effects of light on the competition between phytoplankton species, and (3) effects of climate‐change‐induced changes in the light availability in aquatic ecosystems. Among the fundamental responses of phytoplankton to light, the effects of light intensity (quantity, as measure of total photon or energy flux) were investigated in most identified studies. The effects of the light spectrum (quality) that via species‐specific light absorbance result in direct consequences on species competition emerged more recently. Complexity in competition arises due to variability and fluctuations in light which effects are sparsely investigated on community level. Predictions regarding future climate change scenarios included changes in in stratification and mixing, lake and coastal ocean darkening, UV radiation, ice melting as well as light pollution which affect the underwater light‐climate. Generalization of consequences is difficult due to a high variability, interactions of consequences as well as a lack in sustained timeseries and holistic approaches. Nevertheless, our systematic literature map, and the identified articles within, provide a comprehensive overview and shall guide prospective research.

Biodiversity stabilizes primary productivity through compensatory effects under warming conditions

AbstractAimDiversity may increase the resistance of ecosystem productivity to environmental stress, such as warming, via compensatory processes associated with adjustments of species interactions. However, experimental evidence of compensatory processes that buffer productivity declines in relation to environmental stress is limited, especially in below‐ground settings. We asked whether species richness could stabilize productivity under warming via compensatory responses in root biomass and root functional traits.MethodsUsing three herbaceous species, we created plant communities composed of four individuals in either monocultures or two‐ and three‐species assemblages. We grew them at three temperatures, simulating current climate conditions, moderate warming and severe warming, respectively. We built mixed‐linear mixed models to model plant productivity by species richness and warming and we also analyzed the interactive roles of species richness and warming in species interaction and root functional traits.ResultsWe found that warming reduced both above‐ and below‐ground productivity and shifted the biodiversity–productivity relationship from negative to positive. Productivity reductions were weaker in richer species combinations. Warming ameliorated the strength of interspecific competition below‐ground in mixed‐species communities by reducing the root biomass of strong competitors, which benefitted root growth of weaker competitors.ConclusionsOur results suggest warming can facilitate compensatory responses in herbaceous root productivity across species competition hierarchies. These compensatory processes by which species richness stabilizes plant community functioning emphasize the importance that plant functional diversity has in maintaining ecosystem functioning with climate change.

Minimal-speed selection of traveling fronts to a three components lattice competition system

In this paper, we study the traveling wave problem for a three species lattice competition system with monostable nonlinearity. We are interested in the linear or nonlinear determinacy of the minimal wave speed. Some conditions on the linear determinacy has been obtained in [J.-S. Guo, Y. Wang, C.-H. Wu and C.-C. Wu, The minimal speed of traveling wave solutions for a diffusive three species competition system, Taiwanese J. Math. 19 (2015) 1805–1829]. In this paper, we first provided some general conditions for the linear determinacy. Then, we derive some novel results on the nonlinear determinacy. Finally, some numerical simulations are carried out to illustrate our analytic results. To our knowledge, this may be the first investigation on the nonlinear determinacy of the three species competition system.

Two-species competition model with chemotaxis: well-posedness, stability and dynamics

We study a system of PDEs modelling the population dynamics of two competitive species whose spatial movements are governed by both diffusion and mutually repulsive chemotaxis effects. We prove that solutions to this system are globally well-posed, without any smallness assumptions on the chemotactic coefficients. Moreover, in the weak competition regime, we prove that neither species can be driven to extinction as the time goes to infinity, regardless of how strong the chemotaxis coefficients are. Finally, long-time behaviours of the system are studied both analytically in the weakly nonlinear regime, and numerically in the fully nonlinear regime.

Dynamics of interspecific k species competition model

Herein, we examine the positive equilibrium of a k species competition model with population densities with a condition that each species’ population density is influenced by its sencondn prevnious population density and another species’ previous and second previous population densities i.e., depends on and , for j = 1, 2, … , k. However, for all n.

Temporal resource partitioning mediates vertebrate coexistence at carcasses: The role of competitive and facilitative interactions

Unravelling how biodiversity is maintained despite species competition for shared resources has been a central question in community ecology, and is gaining relevance amidst the current biodiversity crisis. Yet, we have still a poor understanding of the mechanisms that regulate species coexistence and shape the structure of assemblages in highly competitive environments such as carrion pulsed resources. Here, we study how large vertebrates coexist in scavenger assemblages by adapting their diel activity at large ungulate carcasses in NW Spain. We used camera traps to record vertebrate scavengers consuming 34 carcasses of livestock and hunted wild ungulates, which allowed us to assess also differences regarding carcass origin. To evaluate temporal resource partition among species, we estimated the overlap of diel activity patterns and the mean times of each scavenger at carcasses. We recorded 16 species of scavengers, 7 mammals and 9 birds, and found similar richness at both types of carcasses. Birds and mammals showed contrasting diel activity patterns, with birds using carcasses during daytime (mean= 11:38 h) and mammals mostly at night (23:09 h). The unimodal activity patterns of scavengers showed asynchronous peaks among species. Subordinate species modified their activity patterns at carcasses used by apex species to reduce temporal overlap. Also, diel activity patterns of vultures closely followed those of corvids, suggesting facilitation processes in which corvids would enhance carcass detection by vultures. Two mammal species (12.5%) increased nocturnality at carcasses of hunted ungulates, which could be a response to human disturbance. Our results suggest that both temporal segregation and coupling mediate the coexistence of large vertebrates at carcasses. These mechanisms might lead to richer scavenger assemblages and thereby more efficient ones in driving critical ecosystem functions related to carrion consumption, such as energy and nutrient recycling and biodiversity maintenance.

Competitive Interactions between Two Non-Native Species (Alliaria petiolata [M. Bieb.] Cavara & Grande and Hesperis matronalis L.) and a Native Species (Ageratina altissima [L.] R.M. King & H. Rob.).

Alliaria petiolata and Hesperis matronalis are wide-ranging non-native species in North America. Ageratina altissima is native to North America but has become a concern as an invasive species in Asia. A replacement series experiment was established to quantify the competitive interactions between these three species and to rank their relative competitiveness with each other. We assessed leaf count, chlorophyll content, and aboveground biomass with comparisons between replacement series mixtures and competition species. Overall leaf count and aboveground biomass were greatest in A. altissima and chlorophyll content was lowest in A. petiolata. Chlorophyll content and aboveground biomass were lower for A. altissima in competition with A. petiolata compared to H. matronalis. Leaf count for A. petiolata was lower in competition with A. altissima compared to H. matronalis. Aboveground biomass for H. matronalis was lower in competition regardless of the species compared to monoculture. There were also negative trends in biomass for A. petiolata in competition with increasing neighbors. However, for A. altissima, the negative trend in biomass was with A. petiolata, H. matronalis did not negatively affect A. altissima biomass. Our rank order of competitiveness was A. altissima > A. petiolata >> H. matronalis.

Long-term fertilization lowers the alkaline phosphatase activity by impacting the phoD-harboring bacterial community in rice-winter wheat rotation system

PhoD-harboring bacteria and the secreted alkaline phosphatases (ALP) are crucial in the regulation of soil phosphorus (P) cycling. However, the influential factors of these crucial indicators and their internal interactions remain controversial. Here, a long-term field experiment containing different fertilization regimes was conducted (chemical, organic, and no fertilizer applied). The results indicated that the richness and diversity of phoD-harboring bacterial community were significantly decreased after long-term fertilization. The applied fertilizer promoted the growth of competitive species, while phoD-harboring bacteria lost the advantage to outcompete other microorganisms after long-term fertilization. The decreased ALP activity was caused by the declined phoD gene abundance, which is attributed to the comprehensive effects of soil organic C (SOC), total nitrogen (TN), and various forms of P. The random forest models identified SOC, TN, and available P (AP) to be the dominant environmental factors in shaping the phoD-harboring bacterial community. In addition, some other forms of P such as organic P (Po), inorganic P (Pi) or total P (TP) also exerted significant effects. Different fertilization regimes changed the keystone genera that contributed significantly to soil ALP activities, while Pseudolabrys and Pseudomonas were predicted to be the most important genera regardless of different fertilization regimes. This study extends the understanding of the main process and mechanisms of P mobilization in response to different fertilization regimes.

Successional adaptive strategies revealed by correlating arbuscular mycorrhizal fungal abundance with host plant gene expression.

The shifts in adaptive strategies revealed by ecological succession and the mechanisms that facilitate these shifts are fundamental to ecology. These adaptive strategies could be particularly important in communities of arbuscular mycorrhizal fungi (AMF) mutualistic with sorghum, where strong AMF succession replaces initially ruderal species with competitive ones and where the strongest plant response to drought is to manage these AMF. Although most studies of agriculturally important fungi focus on parasites, the mutualistic symbionts, AMF, constitute a research system of human-associated fungi whose relative simplicity and synchrony are conducive to experimental ecology. First, we hypothesize that, when irrigation is stopped to mimic drought, competitive AMF species should be replaced by AMF species tolerant to drought stress. We then, for the first time, correlate AMF abundance and host plant transcription to test two novel hypotheses about the mechanisms behind the shift from ruderal to competitive AMF. Surprisingly, despite imposing drought stress, we found no stress-tolerant AMF, probably due to our agricultural system having been irrigated for nearly six decades. Remarkably, we found strong and differential correlation between the successional shift from ruderal to competitive AMF and sorghum genes whose products (i) produce and release strigolactone signals, (ii) perceive mycorrhizal-lipochitinoligosaccharide (Myc-LCO) signals, (iii) provide plant lipid and sugar to AMF, and (iv) import minerals and water provided by AMF. These novel insights frame new hypotheses about AMF adaptive evolution and suggest a rationale for selecting AMF to reduce inputs and maximize yields in commercial agriculture.

Not all trees can make a forest: Tree species composition and competition control forest encroachment in a tropical savanna

Abstract Forest encroachment into savannas is a widespread phenomenon, the rate of which may depend on soil conditions, species composition or changes in stand structure. As savanna specialist trees are replaced by generalist species, rates of stand development may increase. Because generalists can persist in forests, they are likely to grow more quickly and survive longer in dense stands, compared to savanna specialists. Furthermore, the faster growth rates of generalists may allow them to overtop and outcompete savanna specialists, causing rapid species turnover. We measured growth and survival of 6,147 individuals of 112 species of savanna and generalist tree species over a period of 10 years in an ecological reserve in Assis, São Paulo State, Brazil. We modelled growth and mortality as a function of soil texture and nutrients, tree size, competitive neighbourhood, and membership in savanna or generalist (species which can persist in forests and savannas) functional groups. Tree growth and survival was strongly influenced by competition, as estimated by the basal area of trees taller than a focal tree. At the stand level, savanna species are unable to contribute basal area growth in closed stands, while generalist species continue to increase in basal area even at high stand basal area. This phenomenon is driven by differences in growth and mortality. Generalists grew faster than savanna species, both in height and diameter. This difference in growth rates led to savanna species becoming suppressed more rapidly than generalists. When suppressed, savanna species were more than twice as likely to die than were generalists. Soils had inconsistent and mostly weak effects which were difficult to separate from gradients of stand structure. Synthesis. We demonstrate that the presence of generalist trees accelerates the rates of basal area accumulation due to their greater growth rates and tolerance of shading. Generalists outcompete savanna trees by growing faster in the open and overtopping savanna specialists. Due to the slow growth and high mortality of savanna species in the shade, they are unable to form closed‐canopy stands. Accounting for differences among functional types and development of vegetation structure is critical for modelling forest encroachment.

Drivers of change in mountain and upland bird populations in Europe

Mountain ecosystems have special significance for biodiversity and are vulnerable to climate and other environmental changes. However, few assessments of drivers of change have been conducted in these areas in comparison to other more accessible biomes. In this study, we developed an objective and broad definition of a mountain bird, and systematically reviewed the existing literature for the 34 European mountain and upland species identified in order to quantify which drivers of environmental change have been most consistently associated with positive or negative demographic responses. Raptors and gamebirds were the most‐studied species, whereas mountain specialists and arctic–alpine species were little studied. Using a standardized threat classification, the analyses reported significant negative impacts of hunting, collision with energy‐generation infrastructures, predation by competitive species, human disturbance and poisoning. There were significant positive effects for management of non‐target species, but less support for effects of drivers operating over longer timescales, such as climate and land‐use changes. Analysis of trends in drivers found evidence of increasing temperatures and vegetation encroachment, whereas agricultural impacts decreased, which was largely related to abandonment of grazing. A lack of fine‐grained studies investigating ecological mechanisms and population responses to widespread impacts were highlighted. We have found that some drivers are well studied, but we need to collect more detailed and longer‐term data on species requirements and on the impacts that bird populations face to devise priorities for conservation action and research for mountain and upland birds.

Maintenance of Intraspecific Diversity in Response to Species Competition and Nutrient Fluctuations.

Intraspecific diversity is a substantial part of biodiversity, yet little is known about its maintenance. Understanding mechanisms of intraspecific diversity shifts provides realistic detail about how phytoplankton communities evolve to new environmental conditions, a process especially important in times of climate change. Here, we aimed to identify factors that maintain genotype diversity and link the observed diversity change to measured phytoplankton morpho-functional traits Vmax and cell size of the species and genotypes. In an experimental setup, the two phytoplankton species Emiliania huxleyi and Chaetoceros affinis, each consisting of nine genotypes, were cultivated separately and together under different fluctuation and nutrient regimes. Their genotype composition was assessed after 49 and 91 days, and Shannon’s diversity index was calculated on the genotype level. We found that a higher intraspecific diversity can be maintained in the presence of a competitor, provided it has a substantial proportion to total biovolume. Both fluctuation and nutrient regime showed species-specific effects and especially structured genotype sorting of C. affinis. While we could relate species sorting with the measured traits, genotype diversity shifts could only be partly explained. The observed context dependency of genotype maintenance suggests that the evolutionary potential could be better understood, if studied in more natural settings including fluctuations and competition.

Competitive Adsorption Processes at Clay Mineral Surfaces: A Coupled Experimental and Modeling Approach

Quantification of adsorption processes on clay mineral surfaces is often necessary to predict the extent and the evolution of contaminants’ migration in surficial and underground environments. Many studies have been dedicated to retention measurement as a function of pH and ionic strength in relation with the two main identified adsorption processes for clay minerals, that is, cation exchange on their basal surfaces and surface complexation on their edge surfaces. The latest process has been repeatedly assessed as an effective retention mechanism at circumneutral pH conditions, which often prevail in natural environments. This assessment must however be tempered by the lack of information about competitive processes that can take place with the numerous chemical species present in natural settings, compared to simplified systems investigated in laboratory experiments. In this study, we quantified experimentally the competition between Pb2+, Co2+, Zn2+, and Mg2+ for specific adsorption on montmorillonite edge surfaces. Zn2+ was an effective competitor with Pb2+ and Co2+, and our results showed also unambiguously the influence of Mg2+ concentration levels on the specific adsorption of Pb2+ and Co2+. Because of the high ionic strength used in the experiments, cation exchange with Mg2+ was dismissed as a possible reason for such competition process, leaving specific competitive adsorption on edge surfaces as a unique explanation for our observations. Modeling of Pb2+ adsorption data with a state-of-art electrostatic complexation model for montmorillonite edge surfaces, supported by state-of-the-art, made it possible to distinguish two types of possible competition driving forces: Zn2+ competition for adsorption site occupancy but also detrimental changes in surface electrostatic potential following Mg2+ adsorption on sites neighboring those of Pb2+ adsorption. Mg2+ competition observed in our experiment should apply in most of clayey environments. Consequently, adsorption data obtained on pure clay mineral phases, and the associated models that have been built based on these data without considering the geochemical background in competitive species, may overestimate the retention properties of clay minerals when applied to natural settings.

Invertebrate herbivory rather than competition with tussocks will increasingly delay highland forest regeneration in degraded areas under active restoration

Determining barriers to tree regeneration along elevational gradients is important to predict shifts in regeneration patterns under climate change scenarios. The stress-gradient hypothesis predicts that facilitation predominates at high elevations and competition at low elevations. Invertebrate herbivory may also play an important, yet hardly recognized role, in low elevation and degraded areas. Our objective was to understand the relative changes in facilitation, competition and invertebrate herbivory along an elevational gradient degraded by long-term livestock rearing and repeated wildfires. Our study area was a seasonally dry ecosystem in central Argentina subjected to forest restoration activities. We planted 3000 saplings of the dominant early and late successional tree species in a full factorial design that included elevation (3 levels: low, 1300; intermediate, 1800; and high, 2300 m a.s.l.), microsite treatment, where we manipulated facilitation and competition (4 levels: tussock grasses mowed to the ground, tussock grasses mowed to 15 cm above the ground, unmowed tussock grasses with an average height of 70 cm, and tussock grasses mowed to the ground near rock outcrops) and 5 blocks per elevation. We monitored sapling survival, change in height, and damage by invertebrate herbivores during one year. For both species, sapling survival significantly increased with elevation, while sapling change in height decreased. Survival and change in height for microsite treatments suggest weak competition for the early successional species and facilitation for the late successional species. Notably, we did not find the elevation and microsite treatment interaction predicted by the stress-gradient hypothesis. The main invertebrate herbivores were leaf-cutting ants, which damaged 42, 33 and 0%, and 25, 27 and 0% of the saplings according to species, for the low, intermediate and high elevations, respectively. Damage did not differ significantly between microsite treatments for either species. Survival for saplings with evidence of damage by leaf-cutting ants was 2.5 and 3.4 times lower as compared saplings with no evidences, according to species. Our findings imply that under a climate warming scenario, future facilitation and competition effects will be similar to current effects, whereas leaf-cutting ants rather than competition may increasingly limit tree regeneration.