Microhabitat amelioration and reduced competition among understorey plants as drivers of facilitation across environmental gradients: Towards a unifying framework

Studies of facilitative interactions as drivers of plant richness along environmental gradients often assume the existence of an overarching stress gradient that equally affects the performance of all the species in a given community. However, co-existing species differ in their ecophysiological adaptations, and do not experience the same stress level under particular environmental conditions. Moreover, these studies assume a unimodal relationship between richness and biomass, which is not as general as previously thought. We ignored these assumptions to assess changes in plant–plant interactions and their effect on local species richness across environmental gradients in semi-arid areas of Spain and Australia. We aimed to understand the relative importance of direct (microhabitat amelioration) and indirect (changes in the competitive relationships among the understorey species: niche segregation, competitive exclusion or intransitivity) mechanisms that might underlie the effects of nurse plants on local species richness. By jointly studying these direct and indirect mechanisms using a unifying framework, we found that nurse plants (trees, shrubs and tussock grasses) increased local richness not only by expanding the niche of neighbouring species but also by increasing niche segregation among them, though the latter was not important in all cases. The outcome of the competition-facilitation continuum varied depending on the study area, likely because the different types of stress gradient considered. When driven by both rainfall and temperature, or rainfall alone, the community-wide importance of nurse plants remained constant (Spanish sites), or showed a unimodal relationship along the gradient (Australian sites). This study expands our understanding of the relative roles of plant–plant interactions and environmental conditions as drivers of local species richness in semi-arid environments. The results can also be used to refine predictions about the response of plant communities to environmental change, and to clarify the relative importance of biotic interactions as drivers of such responses.     

Host plant availability potentially limits butterfly distributions under cold environmental conditions

Species ranges are shaped by both climatic factors and interactions with other species. The stress gradient hypothesis predicts that under physiologically stressful environmental conditions abiotic factors shape range edges while in less stressful environments negative biotic interactions are more important. Butterflies provide a suitable system to test this hypothesis since larvae of most species depend on biotic interactions with a specific set of host plants, which in turn can shape patterns of occurrence and distribution. Here we modelled the distribution of 92 butterfly and 136 host plant species with three different modelling algorithms, using distribution data from the Swiss biodiversity monitoring scheme at a 1 × 1 km spatial resolution. By comparing the ensemble prediction for each butterfly species and the corresponding host plant(s), we assessed potential constraints imposed by host plant availability on distribution of butterflies at their distributional limits along the main environmental gradient, which closely parallels an elevational gradient. Our results indicate that host limitation does not play a role at the lower limit. At the upper limit 50% of butterfly species have a higher elevational limit than their primary host plant, and 33% have upper elevational limits that exceed the limits of both primary and secondary hosts. We conclude that host plant limitation was not relevant to butterfly distributional limits in less stressful environments and that distributions are more likely limited by climate, land use or antagonistic biotic interactions. Obligatory dependency of butterflies on their host plants, however, seems to represent an important limiting factor for the distribution of some species towards the cold, upper end of the environmental gradient, suggesting that biotic factors can shape ranges in stressful environments. Thus, predictions by the stress gradient hypothesis were not always applicable.     

Performance of germinating tree seedlings below and above treeline in the Swiss Alps

The germination and early survival of tree seedlings is a critical process for the understanding of treeline dynamics with ongoing climate change. Here we analyzed the performance of 0–4 year-old seedlings of seven tree species at three sites above and below the current altitudinal treeline in the Swiss Central Alps near Davos. Starting from sown seeds, we monitored the seedling performance as proportions of living seedlings, seedling shoot height growth, and biomass allocation over 4 years to examine changes along an elevational gradient. We evaluated the relative importance of the environmental factors soil temperature, light conditions, water use efficiency, and nitrogen availability on seedling performance. During the 4 years, the proportions of living seedlings differed only slightly along the elevational gradient even in species currently occurring at lower elevations. Microsite-specific soil temperature and light availability had only little effect on the proportion of living seedlings and seedling biomass across the elevational gradient. Conversely, seedling biomass and biomass allocation correlated well with the foliar stable nitrogen isotope abundance (δ ¹⁵N) that was used as an indicator for nitrogen availability. Collectively, our results suggested that the early establishment of seedlings of a variety of tree species in the treeline ecotone was not limited by current climatic conditions even beyond the species’ actual upper distribution limit. Nitrogen dynamics appeared to be an important environmental co-driver for biomass production and allocation in very young tree seedlings.     

Adaptive phenotypic divergence in an annual grass differs across biotic contexts*

Climate is a powerful force shaping adaptation within species, yet adaptation to climate occurs against a biotic background: species interactions can filter fitness consequences of genetic variation by altering phenotypic expression of genotypes. We investigated this process using populations of teosinte, a wild annual grass related to maize (Zea mays ssp. mexicana), sampling plants from 10 sites along an elevational gradient as well as rhizosphere biota from three of those sites. We grew half‐sibling teosinte families in each biota to test whether trait divergence among teosinte populations reflects adaptation or drift, and whether rhizosphere biota affect expression of diverged traits. We further assayed the influence of rhizosphere biota on contemporary additive genetic variation. We found that adaptation across environment shaped divergence of some traits, particularly flowering time and root biomass. We also observed that different rhizosphere biota shifted expressed values of these traits within teosinte populations and families and altered within‐population genetic variance and covariance. In sum, our results imply that changes in trait expression and covariance elicited by rhizosphere communities could have played a historical role in teosinte adaptation to environments and that they are likely to play a role in the response to future selection.     

Geomorphological disturbance affects ecological driving forces and plant turnover along an altitudinal stress gradient on alpine slopes

The altitudinal gradient is considered as a stress gradient for plant species because the development and fitness of plant communities tend to decrease as a result of the extreme environmental conditions present at high elevations. Abiotic factors are predicted to be the primary filter for species assemblage in high alpine areas, influencing biotic interactions through both competition for resources and positive interactions among species. We hypothesised that the relative importance of the ecological driving forces that affect the biotic interactions within plant communities changes along an elevation gradient on alpine debris slopes. We used multiple gradient analyses of 180 vegetation plots along an altitudinal range from ~1,600 to 2,600 m and single 100 m-bands in the Adamello-Presanella Group (Central Alps) to investigate our hypothesis; we measured multiple environmental variables related to different ecological driving forces. Our results illustrate that resource limitations at higher elevations affect not only the shift from competition to facilitation among species. A geomorphological disturbance regime along alpine slopes favours the resilience of the high-altitude species within topographic/geomorphological traps. An understanding of the ecological driving forces and positive interactions as a function of altitude may clarify the mechanisms underlying plant responses to present and future environmental changes.     

Elevational Gradients in β-Diversity Reflect Variation in the Strength of Local Community Assembly Mechanisms across Spatial Scales

Despite long-standing interest in elevational-diversity gradients, little is known about the processes that cause changes in the compositional variation of communities (β-diversity) across elevations. Recent studies have suggested that β-diversity gradients are driven by variation in species pools, rather than by variation in the strength of local community assembly mechanisms such as dispersal limitation, environmental filtering, or local biotic interactions. However, tests of this hypothesis have been limited to very small spatial scales that limit inferences about how the relative importance of assembly mechanisms may change across spatial scales. Here, we test the hypothesis that scale-dependent community assembly mechanisms shape biogeographic β-diversity gradients using one of the most well-characterized elevational gradients of tropical plant diversity. Using an extensive dataset on woody plant distributions along a 4,000-m elevational gradient in the Bolivian Andes, we compared observed patterns of β-diversity to null-model expectations. β-deviations (standardized differences from null values) were used to measure the relative effects of local community assembly mechanisms after removing sampling effects caused by variation in species pools. To test for scale-dependency, we compared elevational gradients at two contrasting spatial scales that differed in the size of local assemblages and regions by at least an order of magnitude. Elevational gradients in β-diversity persisted after accounting for regional variation in species pools. Moreover, the elevational gradient in β-deviations changed with spatial scale. At small scales, local assembly mechanisms were detectable, but variation in species pools accounted for most of the elevational gradient in β-diversity. At large spatial scales, in contrast, local assembly mechanisms were a dominant force driving changes in β-diversity. In contrast to the hypothesis that variation in species pools alone drives β-diversity gradients, we show that local community assembly mechanisms contribute strongly to systematic changes in β-diversity across elevations. We conclude that scale-dependent variation in community assembly mechanisms underlies these iconic gradients in global biodiversity.     

Elevational turnover in the composition of leaf miners and their interactions with host plants in Australian subtropical rainforest

Leaf miners are specialist herbivorous insects that are potentially vulnerable to environmental change because of their dependency on particular host plants. Little, however, is known about how climate affects the distribution of leaf miner communities and their interactions with host plants. Elevational gradients are useful tools for understanding how ecological communities respond to local clines in climate. Given that plant communities are known to undergo elevational turnover in response to changes in climatic conditions, we expect that leaf miner species will also change with elevation. We repeatedly hand collected leaf miners along three elevational gradients in subtropical rainforest in eastern Australia. Individual leaf miners were counted and identified to species, and their host plants were recorded. We tested if leaf miner species richness and the number of unique interactions among leaf miner and host plant species were affected by elevation. We also tested if the composition of leaf miner species and the composition of interactions between leaf miners and host plants showed a relationship with elevation. The rarefied number of unique leaf miner–host plant interactions significantly decreased with elevation, with a slight peak at approx. 700 m a.s.l., while neither rarefied or observed species richness (species density) of leaf miners nor observed numbers of unique interactions (interaction density) were significantly affected by elevation. The composition of leaf miner species and the composition of leaf miner–host plant interactions (occurrence of pairwise interactions) were significantly related to elevation. Elevational turnover in leaf miner species composition indicated that different species varied in their response to changes in biotic and/or abiotic conditions imposed by increasing elevation. Through our analyses, we identified four leaf miner species that may be locally vulnerable to climate change, as a result of their restricted elevational distribution and level of host specificity.     

Lags in the response of mountain plant communities to climate change

Rapid climatic changes and increasing human influence at high elevations around the world will have profound impacts on mountain biodiversity. However, forecasts from statistical models (e.g. species distribution models) rarely consider that plant community changes could substantially lag behind climatic changes, hindering our ability to make temporally realistic projections for the coming century. Indeed, the magnitudes of lags, and the relative importance of the different factors giving rise to them, remain poorly understood. We review evidence for three types of lag: “dispersal lags” affecting plant species’ spread along elevational gradients, “establishment lags” following their arrival in recipient communities, and “extinction lags” of resident species. Variation in lags is explained by variation among species in physiological and demographic responses, by effects of altered biotic interactions, and by aspects of the physical environment. Of these, altered biotic interactions could contribute substantially to establishment and extinction lags, yet impacts of biotic interactions on range dynamics are poorly understood. We develop a mechanistic community model to illustrate how species turnover in future communities might lag behind simple expectations based on species’ range shifts with unlimited dispersal. The model shows a combined contribution of altered biotic interactions and dispersal lags to plant community turnover along an elevational gradient following climate warming. Our review and simulation support the view that accounting for disequilibrium range dynamics will be essential for realistic forecasts of patterns of biodiversity under climate change, with implications for the conservation of mountain species and the ecosystem functions they provide.     

Evidence that climate sets the lower elevation range limit in a high‐elevation endemic salamander

A frequent assumption in ecology is that biotic interactions are more important than abiotic factors in determining lower elevational range limits (i.e., the “warm edge” of a species distribution). However, for species with narrow environmental tolerances, theory suggests the presence of a strong environmental gradient can lead to persistence, even in the presence of competition. The relative importance of biotic and abiotic factors is rarely considered together, although understanding when one exerts a dominant influence on controlling range limits may be crucial to predicting extinction risk under future climate conditions. We sampled multiple transects spanning the elevational range limit of Plethodon shenandoah and site and climate covariates were recorded. A two‐species conditional occupancy model, accommodating heterogeneity in detection probability, was used to relate variation in occupancy with environmental and habitat conditions. Regional climate data were combined with datalogger observations to estimate the cloud base heights and to project future climate change impacts on cloud elevations across the survey area. By simultaneously accounting for species’ interactions and habitat variables, we find that elevation, not competition, is strongly correlated with the lower elevation range boundary, which had been presumed to be restricted mainly as a result of competitive interactions with a congener. Because the lower elevational range limit is sensitive to climate variables, projected climate change across its high‐elevation habitats will directly affect the species’ distribution. Testing assumptions of factors that set species range limits should use models which accommodate detection biases.     

Elevational gradient in the cyclicity of a forest-defoliating insect

Observed changes in the cyclicity of herbivore populations along latitudinal gradients and the hypothesis that shifts in the importance of generalist versus specialist predators explain such gradients has long been a matter of intense interest. In contrast, elevational gradients in population cyclicity are largely unexplored. We quantified the cyclicity of gypsy moth populations along an elevational gradient by applying wavelet analysis to spatially referenced 31-year records (1975–2005) of defoliation. Based on geographically weighted regression and nonlinear regression, we found either a hump-shaped or plateauing relationship between elevation and the cyclicity of gypsy moth populations and a positive relationship between cyclicity and the density of the gypsy moth’s preferred host-tree species. The potential effects of elevational gradients in the density of generalist predators and preferred host-tree species on the cyclicity of gypsy moth populations were evaluated with mechanistic simulation models. The models suggested that an elevational gradient in the densities of preferred host tree species could partially explain elevational patterns of gypsy moth cyclicity. Results from a model assuming a type-III functional response of generalist predators to changes in gypsy moth density were inconsistent with the observed elevational gradient in gypsy moth cyclicity. However, a model with a more realistic type-II functional response gave results roughly consistent with the empirical findings. In contrast to classical studies on the effects of generalist predators on prey population cycles, our model with a type-II functional response predicts a unimodal relationship between generalist-predator density and the cyclicity of gypsy moth populations.     

Competition and facilitation may lead to asymmetric range shift dynamics with climate change

Forecasts of widespread range shifts with climate change stem from assumptions that climate drives species' distributions. However, local adaptation and biotic interactions also influence range limits and thus may impact range shifts. Despite the potential importance of these factors, few studies have directly tested their effects on performance at range limits. We address how population‐level variation and biotic interactions may affect range shifts by transplanting seeds and seedlings of western North American conifers of different origin populations into different competitive neighborhoods within and beyond their elevational ranges and monitoring their performance. We find evidence that competition with neighboring trees limits performance within current ranges, but that interactions between adults and juveniles switch from competitive to facilitative at upper range limits. Local adaptation had weaker effects on performance that did not predictably vary with range position or seed origin. Our findings suggest that competitive interactions may slow species turnover within forests at lower range limits, whereas facilitative interactions may accelerate the pace of tree expansions upward near timberline.     

Global patterns of intraspecific leaf trait responses to elevation

Elevational gradients are often used to quantify how traits of plant species respond to abiotic and biotic environmental variations. Yet, such analyses are frequently restricted spatially and applied along single slopes or mountain ranges. Since we know little on the response of intraspecific leaf traits to elevation across the globe, we here perform a global meta‐analysis of leaf traits in 109 plant species located in 4 continents and reported in 71 studies published between 1983 and 2018. We quantified the intraspecific change in seven morpho‐ecophysiological leaf traits along global elevational gradients: specific leaf area (SLA), leaf mass per area (LMA), leaf area (LA), nitrogen concentration per unit of area (Narea), nitrogen concentration per unit mass (Nmass), phosphorous concentration per unit mass (Pmass) and carbon isotope composition (δ¹³C). We found LMA, Narea, Nmass and δ¹³C to significantly increase and SLA to decrease with increasing elevation. Conversely, LA and Pmass showed no significant pattern with elevation worldwide. We found significantly larger increase in Narea, Nmass, Pmass and δ¹³C with elevation in warmer regions. Larger responses to increasing elevation were apparent for SLA of herbaceous compared to woody species, but not for the other traits. Finally, we also detected evidences of covariation across morphological and physiological traits within the same elevational gradient. In sum, we demonstrate that there are common cross‐species patterns of intraspecific leaf trait variation across elevational gradients worldwide. Irrespective of whether such variation is genetically determined via local adaptation or attributed to phenotypic plasticity, the leaf trait patterns quantified here suggest that plant species are adapted to live on a range of temperature conditions. Since the distribution of mountain biota is predominantly shifting upslope in response to changes in environmental conditions, our results are important to further our understanding of how plants species of mountain ecosystems adapt to global environmental change.     

Strain and vegetation effects on local limiting resources explain the outcomes of biotic interactions

Positive interactions are hypothesized to increase with stress (stress-gradient hypothesis, “SGH”), which is defined in terms of standing biomass at the community level. However, recent evidence suggests that facilitation may decrease or remain constant as stress increases. Several reasons for this discrepancy are possible: (i) the outcomes of biotic interactions depend on the component of the fitness considered; (ii) they are influenced by how vegetation affects local limiting resources; (iii) within a particular community, only species that are deviated from their physiological optima are likely to be facilitated. In a removal experiment, we quantified the deviations of species in subalpine grassland from their physiological optima, defined here as species-level “strain”, and examined whether strain and vegetation effects on local resources can explain the outcome of biotic interactions. The experiment was performed along a gradient of standing biomass driven by contrasting land use and resource availability, and used five grass species with contrasting traits and ecological optima. Strain for each species was estimated by comparing growth without vegetation (target species only submitted to local abiotic factors) to growth in optimal conditions (under controlled conditions in an experimental garden). The outcomes of biotic interactions, recorded in terms of survival and growth, could be predicted from the data about strain and vegetation effects on local limiting resources (light and water). Only highly strained species were affected by facilitation, which occurred when the surrounding vegetation alleviated the constraining factors. On the other hand, standing biomass was poorly related with the outcomes of biotic interactions. The “SGH” was only partially validated with growth data when strain and vegetation effects co-varied with standing biomass. As a consequence, strain (at species level) represents a mechanistic basis which could improve the prediction of the outcomes of biotic interactions along ecological gradients.     

Exploring the role of physiology and biotic interactions in determining elevational ranges of tropical animals

Tropical mountains contain some of the world’s richest animal communities as a result of high turnover of species along elevational gradients. We describe an approach to study the roles of biotic and abiotic factors in establishing elevational ranges, and to improve our ability to predict the effects of climate change on these communities. As a framework we use Hutchinson’s concept of the fundamental niche (determined by the match between the physical environment and the organism’s physiological and biophysical characteristics) and realized niche (the subset of the fundamental niche determined by biotic interactions). Using tropical birds as an example, we propose a method for estimating fundamental niches and discuss five biotic interactions that we expect to influence distributions of tropical montane animals: predation, competition, parasites and pathogens, mutualisms, and habitat associations. The effects of biotic factors on elevational ranges have been studied to some extent, but there is little information on physiological responses of tropical montane animals. It will be necessary to understand all of these ecological constraints in concert to predict current and future elevational ranges and potential threats to montane species. Given the importance of tropical mountains as global biodiversity hotspots, we argue that this area of research requires urgent attention.     

Speciation in a keystone plant genus is driven by elevation: a case study in New Guinean Ficus

Much of the world's insect and plant biodiversity is found in tropical and subtropical ‘hotspots’, which often include long elevational gradients. These gradients may function as ‘diversity pumps’ and contribute to both regional and local species richness. Climactic conditions on such gradients often change rapidly along short vertical distances and may result in local adaptation and high levels of population genetic structure in plants and insects. We investigated the population genetic structure of two species of Ficus (Moraceae) along a continuously forested elevational gradient in Papua New Guinea. This speciose plant genus is pollinated by tiny, species‐specific and highly coevolved chalcid wasps (Agaonidae) and represented by at least 73 species at our study gradient. We present results from two species of Ficus sampled from six elevations between 200 m and 2700 m a.s.l. (almost the entire elevational range of the genus) and 10 polymorphic microsatellite loci. These results show that strong barriers to gene flow exist between 1200 m and 1700 m a.s.l. Whereas lowland populations are panmictic across distances over 70 km, montane populations can be disjunct over 4 km, despite continuous forest cover. We suggest that the limited gene flow between populations of these two species of montane Ficus may be driven by environmental limitations on pollinator or seed dispersal in combination with local adaptation of Ficus populations. Such a mechanism may have wider implications for plant and pollinator speciation across long and continuously forested elevational gradients if generalist insect pollinators and vertebrate seed dispersers also form populations based on elevation.     

Do biotic interactions shape both sides of the humped‐back model of species richness in plant communities?

A humped-back relationship between species richness and community biomass has frequently been observed in plant communities, at both local and regional scales, although often improperly called a productivity-diversity relationship. Explanations for this relationship have emphasized the role of competitive exclusion, probably because at the time when the relationship was first examined, competition was considered to be the significant biotic filter structuring plant communities. However, over the last 15 years there has been a renewed interest in facilitation and this research has shown a clear link between the role of facilitation in structuring communities and both community biomass and the severity of the environment. Although facilitation may enlarge the realized niche of species and increase community richness in stressful environments, there has only been one previous attempt to revisit the humped-back model of species richness and to include facilitative processes. However, to date, no model has explored whether biotic interactions can potentially shape both sides of the humped-back model for species richness commonly detected in plant communities. Here, we propose a revision of Grime's original model that incorporates a new understanding of the role of facilitative interactions in plant communities. In this revised model, facilitation promotes diversity at medium to high environmental severity levels, by expanding the realized niche of stress-intolerant competitive species into harsh physical conditions. However, when environmental conditions become extremely severe the positive effects of the benefactors wane (as supported by recent research on facilitative interactions in extremely severe environments) and diversity is reduced. Conversely, with decreasing stress along the biomass gradient, facilitation decreases because stress-intolerant species become able to exist away from the canopy of the stress-tolerant species (as proposed by facilitation theory). At the same time competition increases for stress-tolerant species, reducing diversity in the most benign conditions (as proposed by models of competition theory). In this way our inclusion of facilitation into the classic model of plant species diversity and community biomass generates a more powerful and richer predictive framework for understanding the role of plant interactions in changing diversity. We then use our revised model to explain both the observed discrepancies between natural patterns of species richness and community biomass and the results of experimental studies of the impact of biodiversity on the productivity of herbaceous communities. It is clear that explicit consideration of concurrent changes in stress-tolerant and competitive species enhances our capacity to explain and interpret patterns in plant community diversity with respect to environmental severity.     

The role of selection and historical factors in driving population differentiation along an elevational gradient in an island bird

Adaptation to local environmental conditions and the range dynamics of populations can influence evolutionary divergence along environmental gradients. Thus, it is important to investigate patterns of both phenotypic and genetic variations among populations to reveal the respective roles of these two types of factors in driving population differentiation. Here, we test for evidence of phenotypic and genetic structure across populations of a passerine bird (Zosterops borbonicus) distributed along a steep elevational gradient on the island of Réunion. Using 11 microsatellite loci screened in 401 individuals from 18 localities distributed along the gradient, we found that genetic differentiation occurred at two spatial levels: (i) between two main population groups corresponding to highland and lowland areas, respectively, and (ii) within each of these two groups. In contrast, several morphological traits varied gradually along the gradient. Comparison of neutral genetic differentiation (F_ST) and phenotypic differentiation (P_ST) showed that P_ST largely exceeds F_ST at several morphological traits, which is consistent with a role for local adaptation in driving morphological divergence along the gradient. Overall, our results revealed an area of secondary contact midway up the gradient between two major, cryptic, population groups likely diverged in allopatry. Remarkably, local adaptation has shaped phenotypic differentiation irrespective of population history, resulting in different patterns of variation along the elevational gradient. Our findings underscore the importance of understanding both historical and selective factors when trying to explain variation along environmental gradients.     

Local adaptation across a fertility gradient is influenced by soil biota in the invasive grass, <Emphasis Type="Italic">Bromus inermis</Emphasis>

Biotic soil factors, such as fungi, bacteria and herbivores affect resource acquisition and fitness in plants, yet little is known of their role as agents of selection. Evolutionary responses to these selective agents could be an important mechanism that explains the success of invasive species. In this study, we tested whether populations of the invasive grass Bromus inermis are adapted to their home soil environment, and whether biotic factors influence the magnitude of this adaptation. We selected three populations growing at sites that differed in soil fertility and grew individuals from each population in each soil. To assess whether biotic factors influence the magnitude of adaptation, we also grew the same populations in sterilized field soil. To further examine the role of one element of the soil biota (fungi) in local adaptation, we measured colonization by arbuscular mycorrhizal (AM) and septate fungi, and tested whether the extent of colonization differed between local and foreign plants. In non-sterilized (living) soil, there was evidence of a home site advantage because local plants produced significantly more biomass than at least one of the two populations of foreign plants in all three soil origins. By contrast, there was no evidence of a home site advantage in sterilized soil because local plants never produced significantly more biomass than either population of foreign plants. Fungal colonization differed between local and foreign plants in the living soil and this variation corresponded with biomass differences. When local plants produced more biomass than foreign plants, they were also less intensively colonized by AM fungi. Colonization by septate fungi did not vary between local and foreign plants. Our results suggest that biotic soil factors are important causes of plant adaptation, and that selection for reduced interactions with mycorrhizae could be one mechanism through which adaptation to a novel environment occurs.     

The Importance of Biotic vs. Abiotic Drivers of Local Plant Community Composition Along Regional Bioclimatic Gradients

We assessed if the relative importance of biotic and abiotic factors for plant community composition differs along environmental gradients and between functional groups, and asked which implications this may have in a warmer and wetter future. The study location is a unique grid of sites spanning regional-scale temperature and precipitation gradients in boreal and alpine grasslands in southern Norway. Within each site we sampled vegetation and associated biotic and abiotic factors, and combined broad- and fine-scale ordination analyses to assess the relative explanatory power of these factors for species composition. Although the community responses to biotic and abiotic factors did not consistently change as predicted along the bioclimatic gradients, abiotic variables tended to explain a larger proportion of the variation in species composition towards colder sites, whereas biotic variables explained more towards warmer sites, supporting the stress gradient hypothesis. Significant interactions with precipitation suggest that biotic variables explained more towards wetter climates in the sub alpine and boreal sites, but more towards drier climates in the colder alpine. Thus, we predict that biotic interactions may become more important in alpine and boreal grasslands in a warmer future, although more winter precipitation may counteract this trend in oceanic alpine climates. Our results show that both local and regional scales analyses are needed to disentangle the local vegetation-environment relationships and their regional-scale drivers, and biotic interactions and precipitation must be included when predicting future species assemblages.