Long-term shift towards shady and nutrient-rich habitats in Central European temperate forests.

Biodiversity world-wide has been under increasing anthropogenic pressure in the past century. The long-term response of biotic communities has been tackled primarily by focusing on species richness, community composition and functionality. Equally important are shifts between entire communities and habitat types, which remain an unexplored level of biodiversity change. We have resurveyed > 2000 vegetation plots in temperate forests in central Europe to capture changes over an average of five decades. The plots were assigned to eight broad forest habitat types using an algorithmic classification system. We analysed transitions between the habitat types and interpreted the trend in terms of changes in environmental conditions. We identified a directional shift along the combined gradients of canopy openness and soil nutrients. Nutrient-poor open-canopy forest habitats have declined strongly in favour of fertile closed-canopy habitats. However, the shift was not uniform across the whole gradients. We conclude that the shifts in habitat types represent a century-long successional trend with significant consequences for forest biodiversity. Open forest habitats should be urgently targeted for plant diversity restoration through the implementation of active management. The approach presented here can be applied to other habitat types and at different spatio-temporal scales.

Contrasting biomass allocations explain adaptations to cold and drought in the world's highest-growing angiosperms.

BACKGROUND AND AIMS: Understanding biomass allocation among plant organs is crucial for comprehending plant growth optimization, survival and responses to global change drivers. Yet, mechanisms governing mass allocation in vascular plants from extreme elevations exposed to cold and drought stresses remain poorly understood. METHODOLOGY: We analyzed organ mass weights and fractions in 258 Himalayan herbaceous species across diverse habitats (wetland, steppe, alpine), growth forms (annual, perennial taprooted, rhizomatous, cushiony), and climatic gradients (3500-6150 m elevation) to explore whether biomass distribution adhered to fixed allometric or optimal partitioning rules, and how variation in size, phylogeny, and ecological preferences influence their strategies for resource allocation. KEY FINDINGS: Following the optimal partitioning theory, Himalayan plants distribute more biomass to key organs vital for acquiring and preserving limited resources necessary for their growth and survival. Allocation strategies are mainly influenced by plant growth forms and habitat conditions, notably temperature, water availability, and evaporative demands. Alpine plants primarily invest in belowground stem bases for storage and regeneration, reducing aboveground stems while increasing leaf mass fraction to maximize carbon assimilation in their short growing season. Conversely, arid steppe plants prioritize deep roots over leaves to secure water and minimize transpiration. Wetland plants allocate resources to aboveground stems and belowground rhizomes, enabling them to resist competition and grazing in fertile environments. CONCLUSIONS: Himalayan plants from extreme elevations optimize their allocation strategies to acquire scarce resources under specific conditions, efficiently investing carbon from supportive to acquisitive and protective functions with increasing cold and drought. Intraspecific variation and shared ancestry did not significantly alter Himalayan plants' biomass allocation strategies. Despite diverse evolutionary histories, plants from similar habitats have developed comparable phenotypic structures to adapt to their specific environments. This study offers new insights into plant adaptations in diverse Himalayan environments and underscores the importance of efficient resource allocation for survival and growth in challenging conditions.

Combining multiple investigative approaches to unravel functional responses to global change in the understorey of temperate forests.

Plant communities are being exposed to changing environmental conditions all around the globe, leading to alterations in plant diversity, community composition, and ecosystem functioning. For herbaceous understorey communities in temperate forests, responses to global change are postulated to be complex, due to the presence of a tree layer that modulates understorey responses to external pressures such as climate change and changes in atmospheric nitrogen deposition rates. Multiple investigative approaches have been put forward as tools to detect, quantify and predict understorey responses to these global-change drivers, including, among others, distributed resurvey studies and manipulative experiments. These investigative approaches are generally designed and reported upon in isolation, while integration across investigative approaches is rarely considered. In this study, we integrate three investigative approaches (two complementary resurvey approaches and one experimental approach) to investigate how climate warming and changes in nitrogen deposition affect the functional composition of the understorey and how functional responses in the understorey are modulated by canopy disturbance, that is, changes in overstorey canopy openness over time. Our resurvey data reveal that most changes in understorey functional characteristics represent responses to changes in canopy openness with shifts in macroclimate temperature and aerial nitrogen deposition playing secondary roles. Contrary to expectations, we found little evidence that these drivers interact. In addition, experimental findings deviated from the observational findings, suggesting that the forces driving understorey change at the regional scale differ from those driving change at the forest floor (i.e., the experimental treatments). Our study demonstrates that different approaches need to be integrated to acquire a full picture of how understorey communities respond to global change.

Evaluating plant lineage losses and gains in temperate forest understories: a phylogenetic perspective on climate change and nitrogen deposition.

Global change has accelerated local species extinctions and colonizations, often resulting in losses and gains of evolutionary lineages with unique features. Do these losses and gains occur randomly across the phylogeny? We quantified: temporal changes in plant phylogenetic diversity (PD); and the phylogenetic relatedness (PR) of lost and gained species in 2672 semi-permanent vegetation plots in European temperate forest understories resurveyed over an average period of 40 yr. Controlling for differences in species richness, PD increased slightly over time and across plots. Moreover, lost species within plots exhibited a higher degree of PR than gained species. This implies that gained species originated from a more diverse set of evolutionary lineages than lost species. Certain lineages also lost and gained more species than expected by chance, with Ericaceae, Fabaceae, and Orchidaceae experiencing losses and Amaranthaceae, Cyperaceae, and Rosaceae showing gains. Species losses and gains displayed no significant phylogenetic signal in response to changes in macroclimatic conditions and nitrogen deposition. As anthropogenic global change intensifies, temperate forest understories experience losses and gains in specific phylogenetic branches and ecological strategies, while the overall mean PD remains relatively stable.

Les changements globaux accélèrent les processus de colonisation et d'extinction locales d'espèces, aboutissant à des gains ou à des pertes de lignées évolutives uniques. Ces gains et pertes se produisent-ils de manière aléatoire dans l'arbre phylogénétique ? Nous avons mesuré: les changements de diversité phylogénétique; et la parenté phylogénétique des espèces végétales gagnées ou perdues dans 2672 placettes semi-permanentes disposées dans le sous-bois de forêts tempérées d'Europe sur une période moyenne de 40 ans. Une fois corrigée par la richesse spécifique, la diversité phylogénétique a légèrement augmenté au cours du temps dans les différentes placettes. Les espèces perdues ont une plus grande parenté phylogénétique que les espèces gagnées. Les espèces gagnées sont donc issues d'un plus grand nombre de lignées évolutives que les espèces perdues. Certaines lignées ont gagné ou perdu davantage d'espèces que ce qui est prédit par le hasard : les Ericaceae, les Fabaceae et les Orchidaceae ayant davantage perdu, tandis que les Amaranthaceae, les Cyperaceae, et les Rosaceae ont plus gagné. Il n'y a pas de signal phylogénétique des gains ou pertes d'espèces en réponse aux changements de conditions macroclimatiques ou des dépôts atmosphériques d'azote. Alors que les changements globaux d'origine anthropique s'intensifient, les sous-bois des forêts tempérées connaissent des gains et des pertes de certaines lignées évolutives et de certaines stratégies écologiques, sans que la diversité phylogénétique moyenne ne s'en trouve véritablement affectée.

El cambio global ha acelerado las extinciones y colonizaciones a escala local, lo que a menudo ha supuesto pérdidas y ganancias de linajes evolutivos con características únicas. Ahora bien, ¿estas pérdidas y ganancias ocurren aleatoriamente a lo largo de la filogenia? Cuantificamos: los cambios temporales en la diversidad filogenética de las plantas; y la relación filogenética de las especies perdidas y ganadas en 2.672 parcelas de vegetación semipermanente en sotobosques templados europeos y re-muestreadas durante un período promedio de 40 años. Al controlar por las diferencias en la riqueza de especies, la diversidad filogenética aumentó ligeramente con el tiempo y entre parcelas. Además, las especies perdidas dentro de las parcelas exhibieron un mayor grado de relación filogenética que las especies ganadas. Esto implica que las especies ganadas se originaron en un conjunto de linajes evolutivos más diversos que las especies perdidas. Ciertos linajes también perdieron y ganaron más especies de las esperadas aleatoriamente: Ericaceae, Fabaceae y Orchidaceae experimentaron pérdidas y Amaranthaceae, Cyperaceae y Rosaceae mostraron ganancias. Las pérdidas y ganancias de especies no mostraron ninguna señal filogenética significativa en respuesta a los cambios en las condiciones macro-climáticas y la deposición de nitrógeno. A medida que se intensifica el cambio global antropogénico, los sotobosques temperados experimentan pérdidas y ganancias en ramas filogenéticas y estrategias ecológicas específicas, mientras que la diversidad filogenética media general permanece relativamente estable.

Microclimate reveals the true thermal niche of forest plant species.

Species distributions are conventionally modelled using coarse-grained macroclimate data measured in open areas, potentially leading to biased predictions since most terrestrial species reside in the shade of trees. For forest plant species across Europe, we compared conventional macroclimate-based species distribution models (SDMs) with models corrected for forest microclimate buffering. We show that microclimate-based SDMs at high spatial resolution outperformed models using macroclimate and microclimate data at coarser resolution. Additionally, macroclimate-based models introduced a systematic bias in modelled species response curves, which could result in erroneous range shift predictions. Critically important for conservation science, these models were unable to identify warm and cold refugia at the range edges of species distributions. Our study emphasizes the crucial role of microclimate data when SDMs are used to gain insights into biodiversity conservation in the face of climate change, particularly given the growing policy and management focus on the conservation of refugia worldwide.

Plant-symbiotic fungal diversity tracks variation in vegetation and the abiotic environment along an extended elevational gradient in the Himalayas.

Arbuscular mycorrhizal (AM) fungi can benefit plants under environmental stress, and influence plant adaptation to warmer climates. However, very little is known about the ecology of these fungi in alpine environments. We sampled plant roots along a large fraction (1941-6150 m asl (above sea level)) of the longest terrestrial elevational gradient on Earth and used DNA metabarcoding to identify AM fungi. We hypothesized that AM fungal alpha and beta diversity decreases with increasing elevation, and that different vegetation types comprise dissimilar communities, with cultured (putatively ruderal) taxa increasingly represented at high elevations. We found that the alpha diversity of AM fungal communities declined linearly with elevation, whereas within-site taxon turnover (beta diversity) was unimodally related to elevation. The composition of AM fungal communities differed between vegetation types and was influenced by elevation, mean annual temperature, and precipitation. In general, Glomeraceae taxa dominated at all elevations and vegetation types; however, higher elevations were associated with increased presence of Acaulosporaceae, Ambisporaceae, and Claroideoglomeraceae. Contrary to our expectation, the proportion of cultured AM fungal taxa in communities decreased with elevation. These results suggest that, in this system, climate-induced shifts in habitat conditions may facilitate more diverse AM fungal communities at higher elevations but could also favour ruderal taxa.

ForestClim-Bioclimatic variables for microclimate temperatures of European forests.

Microclimate research gained renewed interest over the last decade and its importance for many ecological processes is increasingly being recognized. Consequently, the call for high-resolution microclimatic temperature grids across broad spatial extents is becoming more pressing to improve ecological models. Here, we provide a new set of open-access bioclimatic variables for microclimate temperatures of European forests at 25 × 25 m2 resolution.

Divergent roles of herbivory in eutrophying forests.

Ungulate populations are increasing across Europe with important implications for forest plant communities. Concurrently, atmospheric nitrogen (N) deposition continues to eutrophicate forests, threatening many rare, often more nutrient-efficient, plant species. These pressures may critically interact to shape biodiversity as in grassland and tundra systems, yet any potential interactions in forests remain poorly understood. Here, we combined vegetation resurveys from 52 sites across 13 European countries to test how changes in ungulate herbivory and eutrophication drive long-term changes in forest understorey communities. Increases in herbivory were associated with elevated temporal species turnover, however, identities of winner and loser species depended on N levels. Under low levels of N-deposition, herbivory favored threatened and small-ranged species while reducing the proportion of non-native and nutrient-demanding species. Yet all these trends were reversed under high levels of N-deposition. Herbivores also reduced shrub cover, likely exacerbating N effects by increasing light levels in the understorey. Eutrophication levels may therefore determine whether herbivory acts as a catalyst for the "N time bomb" or as a conservation tool in temperate forests.

Can high-resolution topography and forest canopy structure substitute microclimate measurements? Bryophytes say no.

Increasingly available high-resolution digital elevation models (DEMs) facilitate the use of fine-scale topographic variables as proxies for microclimatic effects not captured by the coarse-grained macroclimate datasets. Species distributions and community assembly rules are, however directly shaped by microclimate and not by topography. DEM-derived topography, sometimes combined with vegetation structure, is thus widely used as a proxy for microclimatic effects in ecological research and conservation applications. However, the suitability of such a strategy has not been evaluated against in situ measured microclimate and species composition. Because bryophytes are highly sensitive to microclimate, they are ideal model organisms for such evaluation. To provide this much needed evaluation, we simultaneously recorded bryophyte species composition, microclimate, and forest vegetation structure at 218 sampling sites distributed across topographically complex sandstone landscape. Using a LiDAR-based DEM with a 1 m resolution, we calculated eleven topographic variables serving as a topographic proxy for microclimate. To characterize vegetation structure, we used hemispherical photographs and LiDAR canopy height models. Finally, we calculated eleven microclimatic variables from a continuous two-year time- series of air and soil temperature and soil moisture. To evaluate topography and vegetation structure as substitutes for the ecological effect of measured microclimate, we partitioned the variation in bryophyte species composition and richness explained by microclimate, topography, and vegetation structure. In situ measured microclimate was clearly the most important driver of bryophyte assemblages in temperate coniferous forests. The most bryophyte-relevant variables were growing degree days, maximum air temperature, and mean soil moisture. Our results thus showed that topographic variables, even when derived from high-resolution LiDAR data and combined with in situ sampled vegetation structure, cannot fully substitute effects of in situ measured microclimate on forest bryophytes.

Inadequacies of Physical Examination in Patients with Acute Lower Limb Ischemia Are Associated with Dreadful Consequences.

BACKGROUND: Acute lower limb ischemia (ALI) is limb and life-threatening. The aim of this study was to explore the association between adherence to guidelines on clinical diagnosis of ALI and outcome at 1 year. The hypothesis was that that better examination was associated with favorable outcome in ALI patients. METHODS: Retrospective cohort study between 2015 and 2018. In-hospital, operation, radiological and autopsy registries captured 161 citizens of Malmö with ALI. The initial bedside evaluation was performed by an emergency physician. Scoring was based on evaluation of the 6 "Ps" and 1 point was given for pain, pallor, pulselessness, perishing cold, paresthesia, paralysis or ankle-brachial pressure index (ABI). The performance was scored (range 0-7), and a score ≥5 was defined as a satisfactory vascular leg status. A multivariate logistic regression was performed to adjust for confounders and expressed in Odds Ratios (OR) with 95% confidence intervals (CI). RESULTS: A satisfactory first clinical examination was performed in 55.3% of the patients. Measurement of ABI (OR 0.25, 95% CI 0.11-0.55), performing complete pulse status (OR 0.41, 95% CI 0.20-0.85), evaluating paralysis (OR 0.43, 95% CI 0.20-0.89), and a bedside score ≥5 points (OR 0.48, 95% CI 0.23-0.97) were independently associated with reduced risk of major amputation/mortality at 1-year follow up. CONCLUSIONS: Quality of initial bedside evaluation in patients with ALI was unsatisfactory to a large extent and better clinical examinations were associated with favorable outcome at 1 year. Skills in clinical diagnostics in ALI needs to be much improved.

Directional turnover towards larger-ranged plants over time and across habitats.

Species turnover is ubiquitous. However, it remains unknown whether certain types of species are consistently gained or lost across different habitats. Here, we analysed the trajectories of 1827 plant species over time intervals of up to 78 years at 141 sites across mountain summits, forests, and lowland grasslands in Europe. We found, albeit with relatively small effect sizes, displacements of smaller- by larger-ranged species across habitats. Communities shifted in parallel towards more nutrient-demanding species, with species from nutrient-rich habitats having larger ranges. Because these species are typically strong competitors, declines of smaller-ranged species could reflect not only abiotic drivers of global change, but also biotic pressure from increased competition. The ubiquitous component of turnover based on species range size we found here may partially reconcile findings of no net loss in local diversity with global species loss, and link community-scale turnover to macroecological processes such as biotic homogenisation.

ForestTemp - Sub-canopy microclimate temperatures of European forests.

Ecological research heavily relies on coarse-gridded climate data based on standardized temperature measurements recorded at 2 m height in open landscapes. However, many organisms experience environmental conditions that differ substantially from those captured by these macroclimatic (i.e. free air) temperature grids. In forests, the tree canopy functions as a thermal insulator and buffers sub-canopy microclimatic conditions, thereby affecting biological and ecological processes. To improve the assessment of climatic conditions and climate-change-related impacts on forest-floor biodiversity and functioning, high-resolution temperature grids reflecting forest microclimates are thus urgently needed. Combining more than 1200 time series of in situ near-surface forest temperature with topographical, biological and macroclimatic variables in a machine learning model, we predicted the mean monthly offset between sub-canopy temperature at 15 cm above the surface and free-air temperature over the period 2000-2020 at a spatial resolution of 25 m across Europe. This offset was used to evaluate the difference between microclimate and macroclimate across space and seasons and finally enabled us to calculate mean annual and monthly temperatures for European forest understories. We found that sub-canopy air temperatures differ substantially from free-air temperatures, being on average 2.1°C (standard deviation ± 1.6°C) lower in summer and 2.0°C higher (±0.7°C) in winter across Europe. Additionally, our high-resolution maps expose considerable microclimatic variation within landscapes, not captured by the gridded macroclimatic products. The provided forest sub-canopy temperature maps will enable future research to model below-canopy biological processes and patterns, as well as species distributions more accurately.

Topographic Wetness Index calculation guidelines based on measured soil moisture and plant species composition.

Soil moisture controls environmental processes and species distributions, but it is difficult to measure and interpolate across space. Topographic Wetness Index (TWI) derived from digital elevation model is therefore often used as a proxy for soil moisture. However, different algorithms can be used to calculate TWI and this potentially affects TWI relationship with soil moisture and species assemblages. To disentangle insufficiently-known effects of different algorithms on TWI relation with soil moisture and plant species composition, we measured the root-zone soil moisture throughout a growing season and recorded vascular plants and bryophytes in 45 temperate forest plots. For each plot, we calculated 26 TWI variants from a LiDAR-based digital terrain model and related these TWI variants to the measured soil moisture and moisture-controlled species assemblages of vascular plants and bryophytes. A flow accumulation algorithm determined the ability of the TWI to predict soil moisture, while the flow width and slope algorithms had only a small effects. The TWI calculated with the most often used single-flow D8 algorithm explained less than half of the variation in soil moisture and species composition explained by the TWI calculated with the multiple-flow FD8 algorithm. Flow dispersion used in the FD8 algorithm strongly affected the TWI performance, and a flow dispersion close to 1.0 resulted in the TWI best related to the soil moisture and species assemblages. Using downslope gradient instead of the local slope gradient can strongly decrease TWI performance. Our results clearly showed that the method used to calculate TWI affects study conclusion. However, TWI calculation is often not specified and thus impossible to reproduce and compare among studies. We therefore provide guidelines for TWI calculation and recommend the FD8 flow algorithm with a flow dispersion close to 1.0, flow width equal to the raster cell size and local slope gradient for TWI calculation.

Response to Comment on "Forest microclimate dynamics drive plant responses to warming".

Schall and Heinrichs question our interpretation that the climatic debt in understory plant communities is locally modulated by canopy buffering. However, our results clearly show that the discrepancy between microclimate warming rates and thermophilization rates is highest in forests where canopy cover was reduced, which suggests that the need for communities to respond to warming is highest in those forests.

Response to Comment on "Forest microclimate dynamics drive plant responses to warming".

Bertrand et al question our interpretation about warming effects on the thermophilization in forest plant communities and propose an alternative way to analyze climatic debt. We show that microclimate warming is a better predictor than macroclimate warming for studying forest plant community responses to warming. Their additional analyses do not affect or change our interpretations and conclusions.

Forest microclimate dynamics drive plant responses to warming.

Climate warming is causing a shift in biological communities in favor of warm-affinity species (i.e., thermophilization). Species responses often lag behind climate warming, but the reasons for such lags remain largely unknown. Here, we analyzed multidecadal understory microclimate dynamics in European forests and show that thermophilization and the climatic lag in forest plant communities are primarily controlled by microclimate. Increasing tree canopy cover reduces warming rates inside forests, but loss of canopy cover leads to increased local heat that exacerbates the disequilibrium between community responses and climate change. Reciprocal effects between plants and microclimates are key to understanding the response of forest biodiversity and functioning to climate and land-use changes.

Plant's-eye view of temperature governs elevational distributions.

Explaining species geographic distributions by macroclimate variables is the most common approach for getting mechanistic insights into large-scale diversity patterns and range shifts. However, species' traits influencing biophysical processes can produce a large decoupling from ambient air temperature, which can seriously undermine biogeographical inference. We combined stable oxygen isotope theory with a trait-based approach to assess leaf temperature during carbon assimilation (TL ) and its departure (ΔT) from daytime free air temperature during the growing season (Tgs ) for 158 plant species occurring from 3,400 to 6,150 m a.s.l. in Western Himalayas. We uncovered a general extent of temperature decoupling in the region. The interspecific variation in ΔT was best explained by the combination of plant height and δ13 C, and leaf dry matter content partly captured the variation in TL . The combination of TL and ΔT, with ΔT contributing most, explained the interspecific difference in elevational distributions. Stable oxygen isotope theory appears promising for investigating how plants perceive temperatures, a pivotal information to species biogeographic distributions.

Replacements of small- by large-ranged species scale up to diversity loss in Europe's temperate forest biome.

Biodiversity time series reveal global losses and accelerated redistributions of species, but no net loss in local species richness. To better understand how these patterns are linked, we quantify how individual species trajectories scale up to diversity changes using data from 68 vegetation resurvey studies of seminatural forests in Europe. Herb-layer species with small geographic ranges are being replaced by more widely distributed species, and our results suggest that this is due less to species abundances than to species nitrogen niches. Nitrogen deposition accelerates the extinctions of small-ranged, nitrogen-efficient plants and colonization by broadly distributed, nitrogen-demanding plants (including non-natives). Despite no net change in species richness at the spatial scale of a study site, the losses of small-ranged species reduce biome-scale (gamma) diversity. These results provide one mechanism to explain the directional replacement of small-ranged species within sites and thus explain patterns of biodiversity change across spatial scales.

Temporal changes in the spatial distribution of carabid beetles around arable field-woodlot boundaries.

Carabids are considered beneficial arthropods in agroecosystems, where they prey on crop pests or consume weed seeds. Therefore, knowledge of the spatial distribution of carabids in agricultural landscapes is crucial to efficiently manage the ecosystem services that they provide. In the present study, we investigated the spatial distribution of carabids around arable field-woodlot boundaries in different seasons: (1) early spring, (2) late spring, (3) summer and (4) late autumn. The spatial distribution of carabid abundance (activity-density) and species richness varied seasonally, and the total abundance was highest within arable fields, except in early spring when it peaked at the boundaries. The observed pattern was mainly driven by the spatial distribution of the open-habitat species, which aggregated near the field boundaries during winter and early spring. The open-habitat species penetrated into woodlots during the summer season but occurred almost exclusively outside woodlots in the other sampling periods. The abundance of the forest species was highest within woodlots with the exception of the early spring season, when their abundance peaked at the boundaries. Carabid species richness was highest within arable fields in close proximity to woodlot boundaries with the exception of the summer season, when the total species richness was similar across habitats.

Functionally distinct assembly of vascular plants colonizing alpine cushions suggests their vulnerability to climate change.

BACKGROUND AND AIMS: Alpine cushion plants can initially facilitate other species during ecological succession, but later on can be negatively affected by their development, especially when beneficiaries possess traits allowing them to overrun their host. This can be reinforced by accelerated warming favouring competitively strong species over cold-adapted cushion specialists. However, little empirical research has addressed the trait-based mechanisms of these interactions. The ecological strategies of plants colonizing the cushion plant Thylacospermum caespitosum (Caryophyllaceae), a dominant pioneer of subnival zones, were studied in the Western Himalayas. METHODS: To assess whether the cushion colonizers are phylogenetically and functionally distinct, 1668 vegetation samples were collected, both in open ground outside the cushions and inside their live and dead canopies, in two mountain ranges, Karakoram and Little Tibet. More than 50 plant traits related to growth, biomass allocation and resource acquisition were measured for target species, and the phylogenetic relationships of these species were studied [or determined]. KEY RESULTS: Species-based trait-environment analysis with phylogenetic correction showed that in both mountain ranges Thylacospermum colonizers are phylogenetically diverse but functionally similar and are functionally different from species preferring bare soil outside cushions. Successful colonizers are fast-growing, clonal graminoids and forbs, penetrating the cushion by rhizomes and stolons. They have higher root-to-shoot ratios, leaf nitrogen and phosphorus concentrations, and soil moisture and nutrient demands, sharing the syndrome of competitive species with broad elevation ranges typical of the late stages of primary succession. In contrast, the species from open ground have traits typical of stress-tolerant specialists from high and dry environments. CONCLUSION: Species colonizing tight cushions of T. caespitosum are competitively strong graminoids and herbaceous perennials from alpine grasslands. Since climate change in the Himalayas favours these species, highly specialized subnival cushion plants may face intense competition and a greater risk of decline in the future.