Selection against early flowering in geothermally heated soils is associated with pollen but not prey availability in a carnivorous plant.

PREMISE: In high-latitude environments, plastic responses of phenology to increasing spring temperatures allow plants to extend growing seasons while avoiding late frosts. However, evolved plasticity might become maladaptive if climatic conditions change and spring temperatures no longer provide reliable cues for conditions important for fitness. Maladaptative phenological responses might be related to both abiotic factors and mismatches with interacting species. When mismatches arise, we expect selection to favor changes in phenology. METHODS: We combined observations along a soil temperature gradient in a geothermally heated area with pollen and prey supplementation experiments and examined how phenotypic selection on flowering time in the carnivorous plant Pinguicula vulgaris depends on soil temperature, and pollen and prey availability. RESULTS: Flowering advanced and fitness decreased with increasing soil temperature. However, in pollen-supplemented plants, fitness instead increased with soil temperature. In heated soils, there was selection favoring later flowering, while earlier flowering was favored in unheated soils. This pattern remained also after artificially increasing pollen and prey availability. CONCLUSIONS: Plant-pollinator mismatches can be an important reason why evolved plastic responses of flowering time to increasing spring temperatures become maladaptive under novel environmental conditions, and why there is selection to delay flowering. In our study, selection for later flowering remained after artificially increasing pollen availability, suggesting that abiotic factors also contribute to the observed selection. Identifying the factors that make evolved phenological responses maladaptive under novel conditions is fundamental for understanding and predicting evolutionary responses to climate warming.

A natural heating experiment: Phenotypic and genotypic responses of plant phenology to geothermal soil warming.

Under global warming, the survival of many populations of sedentary organisms in seasonal environments will largely depend on their ability to cope with warming in situ by means of phenotypic plasticity or adaptive evolution. This is particularly true in high-latitude environments, where current growing seasons are short, and expected temperature increases large. In such short-growing season environments, the timing of growth and reproduction is critical to survival. Here, we use the unique setting provided by a natural geothermal soil warming gradient (Hengill geothermal area, Iceland) to study the response of Cerastium fontanum flowering phenology to temperature. We hypothesized that trait expression and phenotypic selection on flowering phenology are related to soil temperature, and tested the hypothesis that temperature-driven differences in selection on phenology have resulted in genetic differentiation using a common garden experiment. In the field, phenology was related to soil temperature, with plants in warmer microsites flowering earlier than plants at colder microsites. In the common garden, plants responded to spring warming in a counter-gradient fashion; plants originating from warmer microsites flowered relatively later than those originating from colder microsites. A likely explanation for this pattern is that plants from colder microsites have been selected to compensate for the shorter growing season by starting development at lower temperatures. However, in our study we did not find evidence of variation in phenotypic selection on phenology in relation to temperature, but selection consistently favoured early flowering. Our results show that soil temperature influences trait expression and suggest the existence of genetically based variation in flowering phenology leading to counter-gradient local adaptation along a gradient of soil temperatures. An important implication of our results is that observed phenotypic responses of phenology to global warming might often be a combination of short-term plastic responses and long-term evolutionary responses, acting in different directions.

Multiple mechanisms of early plant community assembly with stochasticity driving the process.

Initial plant establishment is one of the most critical phases in ecosystem development, where an early suite of physical (environmental filtering), biological (seed limitation, species interactions) and stochastic factors may affect successional trajectories and rates. While functional traits are commonly used to study processes that influence plant community assembly in late successional communities, few studies have applied them to primary succession. The objective here was to determine the importance of these factors in shaping early plant community assembly on a glacial outwash plain, Skeiðarársandur, in SE Iceland using a trait based approach. We used data on vascular plant assemblages at two different spatial scales (community and neighborhood) sampled in 2005 and 2012, and compiled a dataset on seven functional traits linked to species dispersal abilities, establishment, and persistence for all species within these assemblages. Trait-based null model analyses were used to determine the processes that influenced plant community assembly from the regional species pool into local communities, and to determine if the importance of these processes in community assembly was dependent on local environment or changed with time. On the community scale, for most traits, random processes dominated the assembly from the regional species pool. However, in some communities, there was evidence of non-random assembly in relation to traits linked to species dispersal abilities, persistence, and establishment. On the neighborhood scale, assembly was mostly random. The relative importance of different processes varied spatially and temporally and the variation was linked to local soil conditions. While stochasticity dominated assembly patterns of our early successional communities, there was evidence of both seed limitation and environmental filtering. Our results indicated that as soil conditions improved, environmental constraints on assembly became weaker and the assembly became more dependent on species availability.

Soil temperature effects on the structure and diversity of plant and invertebrate communities in a natural warming experiment.

Global warming is predicted to significantly alter species physiology, biotic interactions and thus ecosystem functioning, as a consequence of coexisting species exhibiting a wide range of thermal sensitivities. There is, however, a dearth of research examining warming impacts on natural communities. Here, we used a natural warming experiment in Iceland to investigate the changes in above-ground terrestrial plant and invertebrate communities along a soil temperature gradient (10°C-30°C). The α-diversity of plants and invertebrates decreased with increasing soil temperature, driven by decreasing plant species richness and increasing dominance of certain invertebrate species in warmer habitats. There was also greater species turnover in both plant and invertebrate communities with increasing pairwise temperature difference between sites. There was no effect of temperature on percentage cover of vegetation at the community level, driven by contrasting effects at the population level. There was a reduction in the mean body mass and an increase in the total abundance of the invertebrate community, resulting in no overall change in community biomass. There were contrasting effects of temperature on the population abundance of various invertebrate species, which could be explained by differential thermal tolerances and metabolic requirements, or may have been mediated by changes in plant community composition. Our study provides an important baseline from which the effect of changing environmental conditions on terrestrial communities can be tracked. It also contributes to our understanding of why community-level studies of warming impacts are imperative if we are to disentangle the contrasting thermal responses of individual populations.

A framework model for current land condition in Iceland.

Iceland border the Arctic with cold maritime climate and a large proportion of the land placed at highland plateaus. About 1100 years of human disturbance, such as grazing and wood harvesting, has left much of the island's ecosystems in a poor state, ranging from barren deserts to areas with altered vegetative composition and degraded soils. We constructed a novel resilience-based model (RBC-model) for current land condition in Iceland to test which and how factors, including elevation, slope characteristics, drainage, and proximity to volcanic activity, influence the resilience and stability of ecosystems to human disturbances. We tested the model by randomly placing 500 sample areas (250 x 250 m) all over the country and obtaining values for each factor and current land conditions for each area from existing databases and satellite images. Elevation and drainage explained the largest portions of variability in land condition in Iceland, while both proximity to volcanic activity and the presence of scree slopes also yielded significant relationships. Overall, the model explained about 65% of the variability. The model was improved (R2 from 0.65 to 0.68) when the country was divided into four broadly defined regions. Land condition at the colder northern peninsulas was poorer at lower elevations compared to inland positions. This novel RBC model was successful in explaining differences in present land condition in Iceland. The results have implication for current land use management, especially grazing, suggesting that management should consider elevation, drainage, slopes and location within the country in addition to current land condition.

Maladaptive plastic responses of flowering time to geothermal heating.

Phenotypic plasticity might increase fitness if the conditions under which it evolved remain unaltered, but becomes maladaptive if the environment no longer provides reliable cues for subsequent conditions. In seasonal environments, timing of reproduction can respond plastically to spring temperature, maximizing the benefits of a long season while minimizing the exposure to unfavorable cold temperatures. However, if the relationship between early spring temperatures and later conditions changes, the optimal response might change. In geothermally heated ecosystems, the plastic response of flowering time to springtime soil temperature that has evolved in unheated areas is likely to be non-optimal, because soil temperatures are higher and decoupled from air temperatures in heated areas. We therefore expect natural selection to favor a lower plasticity and a delayed flowering in these areas. Using observational data along a natural geothermal warming gradient, we tested the hypothesis that selection on flowering time depends on soil temperature and favors later flowering on warmer soils in the perennial Cerastium fontanum. In both study years, plants growing in warmer soils began flowering earlier than plants growing in colder soils, suggesting that first flowering date (FFD) responds plastically to soil temperature. In one of the two study years, selection favored earlier flowering in colder soils but later flowering in warmer soils, suggesting that the current level of plastic advance of FFD on warmer soils may be maladaptive in some years. Our results illustrate the advantages of using natural experiments, such as geothermal ecosystems, to examine selection in environments that recently have undergone major changes. Such knowledge is essential to understand and predict both ecological and evolutionary responses to climate warming.

Herbivore species coexistence in changing rangeland ecosystems: First high resolution national open-source and open-access ensemble models for Iceland.

Rangeland ecosystems are changing worldwide with the abandonment of extensive pastoralism practices and greater interest for species coexistence. However, the lack of compiled data on current changes in the abundance and distribution of herbivores challenges rangeland management decisions. Here we gathered and made available for the first time the most extensive set of occurrence data for rangeland herbivores in Iceland in an Open Access framework for transparent and repeatable science-based decisions. We mapped fine scale species distribution overlap to identify areas at risk for wildlife-livestock conflict and overgrazing. Nationwide and long term (1861-2021) occurrence data from 8 independent datasets were used alongside 11 predictor raster layers ("Big Data") to data mine and map the distribution of the domestic sheep (Ovis aries), feral reindeer (Rangifer tarandus tarandus), pink-footed geese (Anser brachyrhynchus), and rock ptarmigan (Lagopus muta islandorum) over the country during the summer. Using algorithms of Maxent in R, RandomForest, TreeNet (stochastic gradient boosting) and MARS (Splines) in Minitab-SPM 8.3, we computed 1 km pixel predictions from machine learning-based ensemble models. Our high-resolution models were tested with alternative datasets, and Area Under the Curve (AUC) values that indicated good (reindeer: 0.8817 and rock ptarmigan: 0.8844) to high model accuracy (sheep: 0.9708 and pink-footed goose: 0.9143). Whenever possible, source data and models are made available online and described with ISO-compliant metadata. Our results illustrate that sheep and pink-footed geese have the greatest overlap in distribution with potential implication for wildlife-livestock conflicts and continued ecosystem degradation even under diminishing livestock abundance at higher elevation. These nationwide models and data are a global asset and a first step in making available the best data for science-based sustainable decision-making about national herbivores affecting species coexistence and environmental management.

Author Correction: A global database for metacommunity ecology, integrating species, traits, environment and space.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

A global database for metacommunity ecology, integrating species, traits, environment and space.

The use of functional information in the form of species traits plays an important role in explaining biodiversity patterns and responses to environmental changes. Although relationships between species composition, their traits, and the environment have been extensively studied on a case-by-case basis, results are variable, and it remains unclear how generalizable these relationships are across ecosystems, taxa and spatial scales. To address this gap, we collated 80 datasets from trait-based studies into a global database for metaCommunity Ecology: Species, Traits, Environment and Space; "CESTES". Each dataset includes four matrices: species community abundances or presences/absences across multiple sites, species trait information, environmental variables and spatial coordinates of the sampling sites. The CESTES database is a live database: it will be maintained and expanded in the future as new datasets become available. By its harmonized structure, and the diversity of ecosystem types, taxonomic groups, and spatial scales it covers, the CESTES database provides an important opportunity for synthetic trait-based research in community ecology.

Seed rain and seed bank reveal that seed limitation strongly influences plant community assembly in grasslands.

Dispersal is an important factor in plant community assembly, but assembly studies seldom include information on actual dispersal into communities, i.e. the local propagule pool. The aim of this study was to determine which factors influence plant community assembly by focusing on two phases of the assembly process: the dispersal phase and the establishment phase. At 12 study sites in grazed ex-arable fields in Sweden the local plant community was determined and in a 100-m radius around the centre of each site, the regional species pool was measured. The local seed bank and the seed rain was explored to estimate the local propagule pool. Trait-based models were then applied to investigate if species traits (height, seed mass, clonal abilities, specific leaf area and dispersal method) and regional abundance influenced which species from the regional species pool, dispersed to the local community (dispersal phase) and which established (establishment phase). Filtering of species during the dispersal phase indicates the effect of seed limitation while filtering during the establishment phase indicates microsite limitation. On average 36% of the regional species pool dispersed to the local sites and of those 78% did establish. Species with enhanced dispersal abilities, e.g. higher regional abundance, smaller seeds and dispersed by cattle, were more likely to disperse to the sites than other species. At half the sites, dispersal was influenced by species height. Species establishment was however mainly unlinked to the traits included in this study. This study underlines the importance of seed limitation in local plant community assembly. It also suggests that without information on species dispersal into a site, it is difficult to distinguish between the influence of dispersal and establishment abilities, and thus seed and microsite limitation, as both can be linked to the same trait.

Underdispersion and overdispersion of traits in terrestrial snail communities on islands.

Understanding and disentangling different processes underlying the assembly and diversity of communities remains a key challenge in ecology. Species can assemble into communities either randomly or due to deterministic processes. Deterministic assembly leads to species being more similar (underdispersed) or more different (overdispersed) in certain traits than would be expected by chance. However, the relative importance of those processes is not well understood for many organisms, including terrestrial invertebrates. Based on knowledge of a broad range of species traits, we tested for the presence of trait underdispersion (indicating dispersal or environmental filtering) and trait overdispersion (indicating niche partitioning) and their relative importance in explaining land snail community composition on lake islands. The analysis of community assembly was performed using a functional diversity index (Rao's quadratic entropy) in combination with a null model approach. Regression analysis with the effect sizes of the assembly tests and environmental variables gave information on the strength of under- and overdispersion along environmental gradients. Additionally, we examined the link between community weighted mean trait values and environmental variables using a CWM-RDA. We found both trait underdispersion and trait overdispersion, but underdispersion (eight traits) was more frequently detected than overdispersion (two traits). Underdispersion was related to four environmental variables (tree cover, habitat diversity, productivity of ground vegetation, and location on an esker ridge). Our results show clear evidence for underdispersion in traits driven by environmental filtering, but no clear evidence for dispersal filtering. We did not find evidence for overdispersion of traits due to diet or body size, but overdispersion in shell shape may indicate niche differentiation between snail species driven by small-scale habitat heterogeneity. The use of species traits enabled us to identify key traits involved in snail community assembly and to detect the simultaneous occurrence of trait underdispersion and overdispersion.