Biotic interactions are often ignored in assessments of climate change impacts. However, climate-related changes in species interactions, often mediated through increased dominance of certain species or functional groups, may have important implications for how species respond to climate warming and altered precipitation patterns. We examined how a dominant plant functional group affected the population dynamics of four co-occurring forb species by experimentally removing graminoids in seminatural grasslands. Specifically, we explored how the interaction between dominants and subordinates varied with climate by replicating the removal experiment across a climate grid consisting of 12 field sites spanning broad-scale temperature and precipitation gradients in southern Norway. Biotic interactions affected population growth rates of all study species, and the net outcome of interactions between dominants and subordinates switched from facilitation to competition with increasing temperature along the temperature gradient. The impacts of competitive interactions on subordinates in the warmer sites could primarily be attributed to reduced plant survival. Whereas the response to dominant removal varied with temperature, there was no overall effect of precipitation on the balance between competition and facilitation. Our findings suggest that global warming may increase the relative importance of competitive interactions in seminatural grasslands across a wide range of precipitation levels, thereby favouring highly competitive dominant species over subordinate species. As a result, seminatural grasslands may become increasingly dependent on disturbance (i.e. traditional management such as grazing and mowing) to maintain viable populations of subordinate species and thereby biodiversity under future climates. Our study highlights the importance of population-level studies replicated under different climatic conditions for understanding the underlying mechanisms of climate change impacts on plants.
When plants establish outside their native range, their ability to adapt to the new environment is influenced by both demography and dispersal. However, the relative importance of these two factors is poorly understood. To quantify the influence of demography and dispersal on patterns of genetic diversity underlying adaptation, we used data from a globally distributed demographic research network comprising 35 native and 18 nonnative populations of Plantago lanceolata. Species-specific simulation experiments showed that dispersal would dilute demographic influences on genetic diversity at local scales. Populations in the native European range had strong spatial genetic structure associated with geographic distance and precipitation seasonality. In contrast, nonnative populations had weaker spatial genetic structure that was not associated with environmental gradients but with higher within-population genetic diversity. Our findings show that dispersal caused by repeated, long-distance, human-mediated introductions has allowed invasive plant populations to overcome environmental constraints on genetic diversity, even without strong demographic changes. The impact of invasive plants may, therefore, increase with repeated introductions, highlighting the need to constrain future introductions of species even if they already exist in an area.
Soil seed banks offer plants the possibility to disperse through time. This has implications for population and community dynamics, as recognised by ecological and evolutionary theory. In contrast, the conservation and restoration literature often find seed banks to be depauperate, weedy and without much conservation value or restoration potential. One explanation for these contrasting views might lie in a systematic bias in the sampling of seed banks versus established plant communities. We use the species-area relationship as a tool to assess and compare the per-area species richness and spatial structuring of the diversity of the established plant community versus soil seed banks. To allow this direct comparison we extensively survey the species-area relationship of the vegetation and underlying seed bank of a grassland community across twelve sites spanning regional bioclimatic gradients. We also compile a global dataset of established vegetation and seed banks from published sources. We find that seed banks have consistently higher intercepts and slopes of the relationship, and hence higher diversity at any given spatial scale, than the vegetation both in the field and literature study. This is consistent across habitat types, climate gradients, and biomes. Similarity indices are commonly used to compare vegetation and seed bank, and we find that sampling effort (% of the vegetation area sampled for seed bank) was the strongest predictor of vegetation-seed bank similarity for both the Sørensen (R 2 0.70) and the Raup-Crick (R 2 0.25) index. Our study suggests that the perception that seed banks are intrinsically less diverse than established plant communities has been based more on inadequate sampling than on biological reality. Across a range of ecosystems and climatic settings, we find high diversity in seed banks relative to the established community, suggesting potentially important roles of seed banks in population dynamics and diversity maintenance.
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Questions Is there a shift from positive to negative biotic interaction effects on seedling recruitment along two different stress gradients, temperature and precipitation (the stress‐gradient hypothesis); do such interaction effects differ between species with different bioclimatic affinities? Location Boreal, sub‐alpine and alpine grassland in southern Norway. Methods We tested the stress‐gradient hypothesis by comparing seedling recruitment in bare‐ground gaps where vegetation has been removed vs in extant grassland vegetation in 12 boreal, sub‐alpine and alpine grassland sites along a precipitation gradient. This was tested in (1) a seed‐sowing experiment and (2) in naturally occurring recruitment of alpine, generalist and boreal species. Results Emergence of the sown alpine species was higher in the cold alpine than in the warmer sub‐alpine sites, with no effects of precipitation or vegetation removal. The sown generalists also decreased in emergence towards warmer sites, whereas there was no effect of temperature on the sown boreal species. Vegetation removal, interacting with precipitation, increased the emergence of the generalist and boreal species sown at intermediate precipitation levels. In contrast, interactions between temperature and vegetation removal regulated the emergence of all groups of naturally occurring seedlings. Alpine and generalist species emerged at the highest rate in alpine sites, whereas boreal species had highest emergence in the lowlands. Conclusion For all species groups, strong effects of vegetation removal show that competition from the extant vegetation dominates in controlling seedling emergence across all study sites and species. In generalist and boreal species, positive interactions between vegetation removal and temperature show that competitive interactions affect seedling emergence more strongly towards warmer climates, in line with the stress‐gradient hypothesis. In contrast, alpine species show no such interactions. This suggests that species’ adaptations to climate, in combination with environmental forcing, control seedling emergence along the bioclimatic gradients. Our results have implications for nature conservation, as we propose that disturbance from grazing animals can be useful to release competition and thereby increase seedling recruitment and biodiversity in boreal and alpine grasslands in a warmer future.
Questions What are the most important factors explaining present‐day variation in species composition in a glacier foreland? Does the rate of species compositional change in glacier forelands decelerate through primary succession? How do data set properties and analytic methods influence our understanding of glacier foreland successional dynamics? Location Nigardsbreen glacier foreland, western Norway. Methods We sampled the species composition and recorded 21 explanatory variables in 74 plots, distributed on eight classes of terrain age (dated moraines). Gradients in species composition found using global non‐metric multidimensional scaling (GNMDS) ordination were interpreted with split‐plot generalized linear models. Yearly succession rates were calculated from plot positions along a vector of maximum compositional change related to terrain age in the interpreted GNMDS ordination. Results We interpreted the main gradient in species composition as being related to a complex gradient with soil moisture and soil nutrients as primary constituents. Terrain age contributed to the second most important gradient. Succession rates were nonlinear with time. Terrain age explained variation in species composition only when plots from the two youngest terrain age classes were retained in the data set. Conclusions In contrast to the majority of studies of glacier foreland successions performed so far, we find that terrain age is not the principal factor that explains present‐day variation in species composition. Instead, local environmental variables are the main determinants of species composition. This result emphasizes the importance of taking environmental gradients into account when variation in glacier foreland vegetation is studied. The limited importance of terrain age in our study is interpreted as likely due to this glacier foreland being situated below the tree line, the relatively long distance between the bulk of the studied foreland and the glacier snout, and inclusion of few plots from young terrain in our data set. The non‐linearity of succession rates with time implies that a linear time‐since‐deglaciation variable is inappropriate for constrained ordination of glacier foreland vegetation.
Abstract1. Species composition is a vital attribute of any ecosystem. Accordingly, ecological restoration often has the original, or "natural," species composition as its target.However, we still lack adequate methods for predicting the expected time to compositional recovery in restoration studies.2. We describe and explore a new, ordination regression-based approach (ORBA) for predicting time to recovery that allows both linear and asymptotic (logarithmic) relationships of compositional change with time. The approach uses distances between restored plots and reference plots along the successional gradient, represented by a vector in ordination space, to predict time to recovery. Thus, the approach rests on three requirements: (a) the general form of the relationship between compositional change and time must be known; (b) a sufficiently strong successional gradient must be present and adequately represented in a species compositional dataset; and (c) a restoration target must be specified. We tested the approach using data from a boreal old-growth forest that was followed for 18 years after experimental disturbance. Data from the first 9 years after disturbance were used to develop models, the subsequent 9 years for validation.3. Rates of compositional recovery in the example dataset followed the general pattern of decrease with time since disturbance. Accordingly, linear models were too optimistic about the time to recovery, whereas the asymptotic models provided more precise predictions. Synthesis and applications.Our results demonstrate that the new approach opens for reliable prediction of recovery rates and time to recovery using species compositional data. Moreover, it allows us to assess whether recovery proceeds in the desired direction and to quantitatively compare restoration speed, and hence effectiveness, between alternative management options.
Millennia of human land-use have resulted in the widespread occurrence of what have been coined ‘domesticated ecosystems’. The anthropogenic imprints on diversity, composition, structure and functioning of such systems are well documented. However, evolutionary consequences of human activities in these ecosystems are enigmatic. Calluna vulgaris (L.) is a keystone species of coastal heathlands in northwest Europe, an ancient semi-natural landscape of considerable conservation interest. Like many species from naturally fire-prone ecosystems, Calluna shows smoke-adapted germination, but it is unclear whether this trait arose prior to the development of these semi-natural landscapes or is an evolutionary response to the anthropogenic fire regime. We show that smoke-induced germination in Calluna is found in populations from traditionally burnt coastal heathlands but is lacking in naturally occurring populations from other habitats with infrequent natural fires. Our study thus demonstrates evolutionary imprints of human land-use in semi-natural ecosystems. Evolutionary consequences of historic anthropogenic impacts on wildlife have been understudied, but understanding these consequences is necessary for informed conservation and ecosystem management.
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