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Clonality of Thymus longicaulis

Clonality of Thymus longicaulis

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Empirical studies that link plants intraspecific variation to environmental conditions are almost lacking, despite their relevance in understanding mechanisms of plant adaptation, in predicting the outcome of environmental change and in conservation. Here, we investigate intraspecific trait variation of four grassland species along with abiotic env...

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... Ongoing global change (including climate warming, rising atmospheric nitrogen deposition, soil acidification, land-use change) does not only affect the species composition of grasslands , Duprč et al. 2010, Wesche et al. 2012, Diekmann et al. 2014, but also their functional trait composition , Wellstein et al. 2013, Lewis et al. 2014, Helm et al. 2019, Pichon et al. 2022. Within the last decades, an increasingly dramatic loss of species (Bruelheide et al. 2020, Jandt et al. 2022, Meier et al. 2022b) and changes in functional diversity (Bernhardt-Römermann et al. 2011, Socher et al. 2012) have been found for species-rich, often endangered and protected xerothermic (dry or semidry, respectively) grasslands . ...
... To predict the outcome of environmental changes on xerothermic grasslands, the underlying mechanisms of plant adaptation based on functional traits are an important prerequisite (Wellstein et al. 2013). In particular, soil properties (e.g. ...
... To characterize the ITV of the five dominant grasses, the coefficient of variation (CV) was calculated (Everitt 1998). Higher CV values indicate that species have a high ITV and thus their traits are more strongly influenced by environmental factors (according to Wellstein et al. 2013 a very high variability if CV > 0.51). We applied one-way analysis of variance (ANOVA) with subsequent Tukey's post-hoc test using the multcomp package (Hothorn et al. 2008) to determine differences in both the unweighted functional traits (normally distributed data) and their CV between the five dominant grass species and additionally for B. erectus between dry and semidry grasslands. ...
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Functional traits mediate how species and communities respond to (or affect) environmental gradients. These are impacted by global change, which has led to e.g. climate change and land-use change, affecting soil conditions, species richness and functional diversity in, among others, xerothermic (respectively dry or semidry) grasslands. Within the last decades, the dominance of grass species like Bromus erectus has increased in such grasslands in Central Germany, but factors driving their intraspecific trait variability (ITV) are not yet well understood. The aim of our study was, on the one hand, to compare the functional trait composition of two grassland types, namely dry and semidry, using multi-and single-trait approaches and to assess the effects of soil properties on these traits and, on the other hand, to reveal differences in functional traits and their ITV between the five dominant grasses B. erectus, Brachypodium pinnatum, Festuca rupicola, Helictotrichon pratense and Stipa capillata. Based on vegetation relevés, functional traits (VPH-vegetative plant height, LDM-leaf dry mass, LA-leaf area, SLA-specific leaf area, LDMC-leaf dry matter content, LNC-leaf nitrogen concentration, LCC-leaf carbon concentration, leaf C/N ratio) were measured and soil factors (soil depth, pH value, CaCO3 content, soil N and C content, soil C/N ratio) analysed. For each plot, the community weighted mean (CWM) of all functional traits was calculated to determine differences between the two grassland types and the coefficient of variation was used for interpreting differences in the ITV between the five grasses. There were minor differences between dry and semidry grasslands in the CWM of the functional traits LDM, LA, LNC and LCC, while other traits did not differ between the two grassland types. Soil factors had little effects on the trait composition of dry and semidry grasslands, although soil depth, CaCO3 content and C/N ratio had the greatest influence on CWM and were potentially the strongest drivers for differentiation. The five grasses had species-specific trait distributions but showed relatively similar ITV, so we conclude that B. erectus was not more adapted to changing environmental conditions than the other grasses. Generally, we only found minor changes in the functional trait composition of dry and semidry grasslands in Central Germany and thus the environmental gradient was too small to derive clear differences in the ecosystem function between both grassland types, although the functional structure was largely determined by the dominant grasses.
... Such combined environmental effects may become more problematic in the future. To fully understand the resilience of the dry grasslands to climate change impacts, further research is needed to determine the adaptive capacity of dry grassland species, as for example expressed in functional trait variation (Midolo et al., 2021;Wellstein et al., 2013). ...
... Such combined environmental effects may become more problematic in the future. To fully understand the resilience of the dry grasslands to climate change impacts, further research is needed to determine the adaptive capacity of dry grassland species, as for example expressed in functional trait variation (Midolo et al., 2021;Wellstein et al., 2013). ...
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European inner-alpine dry grasslands face substantial threats within the increasingly human-altered landscape, endangering their persistence. To understand changes in dry grassland communities, we revisited historical vegetation plots of Josias Braun-Blanquet after 70 years in Val Venosta, Italy, hosting rare steppe-like grassland vegetation. By disentangling the key environmental factors encompassing climate, land use and human management , and ecological site preferences, we aimed at explaining changes in dry grassland communities with implications for future conservation. By extending our analysis beyond conventional dissimilarity metrics and adopting a landscape-ecological perspective accounting for species-environment interactions, we assessed how environmental changes affect dissimilarity patterns among historical sites, recent non-protected sites, and recent protected sites with generalized additive modelling. Moreover, we examined ecologically significant species changes to evaluate their contribution to community variation within and between sites, discerning their consequences at the landscape scale. Our results revealed significant changes in dry grassland sites, both on non-protected and protected sites. The encroachment of shrubs was associated with a significant increase in generalist species, including various woody species on sites where grazing had ceased. Furthermore, we observed a higher abundance of nutrient-demanding species on sites next to intensive agriculture. These trends were consistent regardless of the protection status, implying that current conservation measures may be insufficient to guarantee their future persistence. To ensure the long-term conservation of typical inner-alpine dry grasslands, interdisciplinary conservation efforts are essential to address adverse environmental impacts across the entire landscape.
... Such combined environmental effects may become more problematic in the future. To fully understand the resilience of the dry grasslands to climate change impacts, further research is needed to determine the adaptive capacity of dry grassland species, as for example expressed in functional trait variation (Midolo et al., 2021;Wellstein et al., 2013). ...
... However, plants are susceptible to changing environmental conditions because of their sessile lifestyle (Anderson et al., 2011). From the short-term point of view, most plants can adjust adaptive strategies (resource acquisition and investment) quickly by phenotypic plasticity and show significant trait differences facing different environmental conditions (Wellstein et al., 2013). Moreover, trait responses to changes in environmental conditions are species-specific, population-specific, and trait-specific (Roybal & Butterfield, 2019;Wang et al., 2015). ...
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Understanding patterns of intraspecific trait variation can help us understand plant adaptability to environmental changes. To explore the underlying adaptation mechanisms of zonal plant species, we selected seven populations of Stipa krylovii , a dominant species in the Inner Mongolia Steppe of China, and evaluated the effects of phenotypic plasticity and genetic differentiation, the effects of climate variables on population trait differentiation, and traits coordinated patterns under each soil moisture treatment. We selected seeds from seven populations of S. krylovii in the Inner Mongolia Steppe, China, and carried out a soil moisture (2) × population origin (7) common garden experiment at Tianjin City, China, and measured ten plant traits of S. krylovii . General linear analyses were used to analyze how soil moisture and population origin affected each trait variation, Mantel tests were used to analyze population trait differentiation—geographic distance (or climatic difference) relationships, regression analyses were used to evaluate trait‐climatic variable relationships, and plant trait networks (PTNs) were used to evaluate traits coordinated patterns. Both soil moisture and population origin showed significant effects on most of traits. Aboveground biomass, root‐shoot ratio, leaf width, specific leaf area, and leaf nitrogen (N) content were significantly correlated with climate variables under the control condition. Specific leaf area and leaf N content were significantly correlated with climate variables under the drought condition. By PTNs, the hub trait(s) was plant height under the control condition and were aboveground biomass, root length, and specific leaf area under the drought condition. This study indicates that both phenotypic plasticity and genetic differentiation can significantly affect the adaptability of S. krylovii . In addition, soil moisture treatments show significant effects on trait‐climate relationships and traits coordinated patterns. These findings provide new insights into the adaptive mechanisms of zonal species in the semiarid grassland region.
... The fact that high LDMC values are inversely related to relative growth rate 34,35 indicates that ice hole individuals follow a more conservative strategy, with low growth rates and producing more resistant and longer-lived leaves compared to subalpine individuals 36 . This may confirm the adoption of an ecological strategy in ice holes related to the survival of individuals for a long time investing mainly in the capacity to conserve resources and repair cellular components decreasing leaf turnover 37,38 . Increased LA would explain lower SD values in ice hole populations than subalpine ones. ...
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In the mountain terrain, ice holes are little depressions between rock boulders that are characterized by the exit of cold air able to cool down the rock surface even in summer. This cold air creates cold microrefugia in warmer surroundings that preserve plant species probably over thousands of years under extra-zonal climatic conditions. We hypothesized that ice hole populations of the model species Vaccinium vitis-idaea (Ericaceae) show genetic differentiation from nearby zonal subalpine populations, and high functional trait distinctiveness, in agreement with genetic patterns. We genotyped almost 30,000 single nucleotide polymorphisms using restriction site-associated DNA sequencing and measured eight functional traits indicative of individual performance and ecological strategies. Genetic results showed high differentiation among the six populations suggesting isolation. On siliceous bedrock, ice hole individuals exhibited higher levels of admixture than those from subalpine populations which could have experienced more bottlenecks during demographic fluctuations related to glacial cycles. Ice hole and subalpine calcareous populations clearly separated from siliceous populations, indicating a possible effect of bedrock in shaping genetic patterns. Trait analysis reflected the bedrock effect on populations’ differentiation. The significant correlation between trait and genetic distances suggests the genetic contribution in shaping intraspecific functional differentiation. In conclusion, extra-zonal populations reveal a prominent genetic and phenotypic differentiation determined by history and ecological contingency. Therefore, microrefugia populations can contribute to the overall variability of the species and lead to intraspecific-driven responses to upcoming environmental changes.
... In marshy and agricultural habitats', plants had a higher SLA and lower LMA than roadside habitat, indicating a higher photosynthetic rate, growth rate, shorter leaf span, and weaker water-use efficiency, with more resources allocated to biomass production and reproductive output (Karbstein et al., 2020). In agricultural land habitat, longer shoots indicate a plant's competitive advantage and increased vigour (Wellstein et al., 2013). Higher specific root length on farmland indicates greater plant nutrient absorption relative to carbon investment (Ordoñez et al., 2009). ...
... According to Bazzaz (1996), adaptable species should have greater phenotypic plasticity. Phenotypic plasticity has been linked to successful plant invasion Richards et al., 2006), as it allows the species to adapt to different environmental conditions (Wellstein et al., 2013). Other reasons for trait variations could be genetic differentiations (Wellstein et al., 2013) or epigenetic changes (Bossdorf et al., 2008), but testing these effects was beyond the scope of this study. ...
... Phenotypic plasticity has been linked to successful plant invasion Richards et al., 2006), as it allows the species to adapt to different environmental conditions (Wellstein et al., 2013). Other reasons for trait variations could be genetic differentiations (Wellstein et al., 2013) or epigenetic changes (Bossdorf et al., 2008), but testing these effects was beyond the scope of this study. In support of our findings, Grassein et al. (2010) found phenotypic plasticity rather than genetic differentiation in two subalpine species (Dactylis glomerata and Festuca paniculata). ...
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Climate change and anthropogenic pressures have resulted in a significant shift in the invasion susceptibility and frequency of non-native species in mountain ecosystems. Cirsium arvense (L.) Scop. (Family: Asteraceae) is an invasive species that spreads quickly in mountains, especially in the trans-Himalayan region of Ladakh. The current study used a trait-based approach to evaluate the impact of local habitat heterogeneity (soil physico-chemical properties) on C. arvense. Thirteen plant functional traits (root, shoot, leaf, and reproductive traits) of C. arvense were studied in three different habitat types (agricultural, marshy, and roadside). Functional trait variability in C. arvense was higher between, than within habitats (between different populations). All the functional traits interacted with habitat change, except for leaf count and seed mass. Soil properties strongly affect C. arvense's resource-use strategies across habitats. The plant adapted to a resource-poor environment (roadside habitat) by conserving resources and to a resource-rich environment (agricultural and marshy land habitat) by acquiring them. The ability of C. arvense to use resources differently reflects its persistence in introduced habitats. In summary, our study shows that C. arvense invades different habitats in introduced regions through trait adaptations and resource-use strategies in the trans-Himalayan region.
... Grazing species effects were also modulated by slope in the forb and grass biomass proportion models. Despite the high importance of topographical factors for plant composition (Sebastià, 2004;Marini et al., 2007;Wellstein et al., 2013;Niu et al., 2019) few studies have addressed the relationship between forbs and grasses with slope (but see Sanaei et al., 2018;Averett et al., 2022). Under cattle and mixed grazing, forb proportion seemed to be reduced in steep terrains, whereas grass biomass became more dominant. ...
Article
Questions Does grazing management shape the proportion of plant guilds (grasses, legumes, forbs and sedges) in mountain grasslands? Which properties of grazing management have the greatest effect on the proportion of plant guilds: grazer diversity/identity or grazing intensity? Are the effects of grazing management on guild proportion modified by other environmental variables that explain plant guild distribution at broad spatial scales? Location Mountains in the Pyrenees. Methods We modelled the proportion of grasses, legumes, forbs and sedges using data from the PASTUS database ( n = 96), which contains a wide range of environmental and management conditions due to the high variety of environmental conditions in mountain grasslands in the Pyrenees. We used a machine‐learning algorithm to find those variables that best explained the proportions of each plant guild. We focussed on the differences between the levels of grazing intensity and the grazing species included in the model, and on detecting interactions between grazing variables and climate, topography and soil conditions. Results The proportion of forbs and grasses strongly depended on the grazing livestock species at broad spatial scales. Only soil pH showed a higher overall explanatory power on guild distribution. In general, forbs were favoured in cattle‐ and grasses in sheep‐grazed grasslands, the latter also being favoured on acidic soils, while forbs were favoured in more alkaline soil conditions. However, the effects of those factors (grazing species and soil pH) were modulated through interactions with several other environmental variables, including soil Mg, K and P, and terrain slope. In contrast, grazing intensity was a minor driver of guild distribution. Conclusions Our results provide information about the relationship between plant functional diversity, indicated by the different plant guild proportions, and grazing management in the Pyrenean grasslands. This information could be useful for developing hypotheses for future experimental studies and for designing policies to improve the management of mountain grasslands.
... Depending on surface moisture levels, we would expect subsurface traits (e.g., root traits) to respond accordingly, leading to trade-offs with above-ground traits adapted for above-ground water acquisition (e.g., height), and consequently increased functional richness in areas more heterogeneous in surface moisture. However, research on functional richness in temperate climates does not corroborate this theory, at least at local scales [25,77,78]. Other abiotic resources may be more limiting, being more pervasive in defining trait responses within temperate regions. ...
... Spatial heterogeneity in soil chemistry (Mg, K, Ca, P, nitrate, ammonium, and C/N) represents heterogeneity in the abiotic resource pool, thus driving richness in resource-acquisition traits for plants. This trend has been recognised in tropical [15] and montane systems [78], as well as in greenhouse experiments [38], though findings have been mixed for multivariate measures of trait diversity, including functional richness and divergence [15,50,81]. Notably, resource acquisition strategies in plants are linked to both aboveand below-ground traits; yet, measures of multivariate trait diversity have focused almost exclusively on above-ground traits, overlooking below-ground traits (e.g., rooting depth), which likely vary, depending on plant capacity for below-ground resource uptake. ...
... However, soil type may be too crude to capture ecologically relevant information on trait responses. Openly available databases (e.g., SoilGrids [84]) allow global characterisation of heterogeneity across multiple soil properties, such as nitrogen and organic carbon content [15,78]. However, as with many geospatial datasets covering broad extents, scaling issues, static data for dynamic features, and errors and uncertainty associated with widespread data interpolation for missing values mean that validation with local variables may be necessary for correct interpretation of such analyses [85]. ...
Article
Geodiversity - the abiotic heterogeneity of Earth's (sub)surface - is gaining recognition for its ecological links to biodiversity. However, theoretical and conceptual knowledge of geodiversity-trait diversity relationships is currently lacking and can improve understanding of abiotic drivers of community assembly. Here we synthesise the state of knowledge of these relationships. We find that some components of geodiversity (e.g., topographic heterogeneity) elicit strong trait responses, whereas other components (e.g., substrate heterogeneity) have marginal effects in driving trait distributions. However, current knowledge is lacking in key aspects, including geodiversity's effect on trait-specific diversity and intraspecific variation. We call for the explicit inclusion of geodiversity when relating environmental drivers to trait diversity, taking advantage of the increasing availability of trait and geodiversity data.
... The composition of plant traits, on the other hand, has a significant impact on soil carbon sequestration within biomes. Plant characteristics, such as flammability and combustibility, have a significant impact on fire regimes, resulting in sudden and massive carbon losses (Wellstein et al., 2013). While the relative abundances and productivities of the major plant functional categories, as well as their characteristics, are most likely the most important factors influencing soil carbon dynamics, interactions between plant species, as well as their avoidance, may also be important (Faucon et al., 2017). ...
Chapter
Climate change has had an impact on the ecosystem, reducing biodiversity, and threatening its stability. Ecologists have recently begun to focus on plant functional features (such as leaf traits, stem traits, regenerative traits, root traits, etc.). Functional characteristics have been shown to be an effective method for investigating ecological issues at all scales, including individual plants, populations, communities, and ecosystems. Changes in plant functional features will indicate climate change and soil health because plant functional traits respond to and adapt to environmental changes effectively. Soil carbon sequestration and dynamics, soil nutrient availability and fluctuation, soil microbiota diversity and quantity, and plant functional traits are all closely linked, and understanding this relationship is critical to addressing climate change issues. Plant functional traits are widely used in ecosystem ecology to address and comprehend the role of species in ecosystem function and their response to changing environmental conditions. This book chapter discusses plant functional traits and their relationship to soil functions, particularly mountainous soil. The chapter includes extensive research definitions of plant traits and soil function, as well as its ecosystem services, by delving into the interaction between plant features and carbon and nutrient dynamics, biodiversity pool, and other aspects of land as well as mountainous soil ecosystem.