Soil seed bank responses to edge effects in temperate European forests.

Aim: The amount of forest edges is increasing globally due to forest fragmentation and land-use changes. However, edge effects on the soil seed bank of temperate forests are still poorly understood. Here, we assessed edge effects at contrasting spatial scales across Europe and quantified the extent to which edges can preserve the seeds of forest specialist plants. Location: Temperate European deciduous forests along a 2,300-km latitudinal gradient. Time period: 2018-2021. Major taxa studied: Vascular plants. Methods: Through a greenhouse germination experiment, we studied how edge effects alter the density, diversity, composition and functionality of forest soil seed banks in 90 plots along different latitudes, elevations and forest management types. We also assessed which environmental conditions drive the seed bank responses at the forest edge versus interior and looked at the relationship between the seed bank and the herb layer species richness. Results: Overall, 10,108 seedlings of 250 species emerged from the soil seed bank. Seed density and species richness of generalists (species not only associated with forests) were higher at edges compared to interiors, with a negative influence of C : N ratio and litter quality. Conversely, forest specialist species richness did not decline from the interior to the edge. Also, edges were compositionally, but not functionally, different from interiors. The correlation between the seed bank and the herb layer species richness was positive and affected by microclimate. Main conclusions: Our results underpin how edge effects shape species diversity and composition of soil seed banks in ancient forests, especially increasing the proportion of generalist species and thus potentially favouring a shift in community composition. However, the presence of many forest specialists suggests that soil seed banks still play a key role in understorey species persistence and could support the resilience of our fragmented forests.

Lichens buffer tundra microclimate more than the expanding shrub Betula nana.

BACKGROUND AND AIMS: In tundra systems, soil-borne lichens are often the dominant groundcover organisms, and act to buffer microclimate extremes within or at the surface of the soil. However, shrubs are currently expanding across tundra systems, potentially causing major shifts in the microclimate landscape. METHODS: Here, we compared soil temperature and moisture underneath the dwarf birch Betula nana and seven abundant lichen species in sub-alpine Norway. We also examined mixtures of lichens and dwarf birch - an intermediate phase of shrubification - and measured several functional traits relating to microclimate. KEY RESULTS: We found that all lichen species strongly buffered the daily temperature range, on average reducing maximum temperatures by 6.9 °C (± 0.7 s.d.) and increasing minimum temperatures by 1.0 °C (± 0.2 s.d.) during summer. The dwarf birch had a much weaker effect (maximum reduced by 2.4 ±â€…5.0 °C and minimum raised by 0.2 ± 0.9 °C). In species mixtures, the lichen effect predominated, affecting temperature extremes by more than would be expected from their abundance. Lichens also tended to reduce soil moisture, which could be explained by their ability to intercept rainfall. Our trait measurements under laboratory conditions suggest that, on average, lichens can completely absorb a 4.09 mm (± 1.81 s.d.) rainfall event, which might be an underappreciated part of lichen-vascular plant competition in areas where summer rainfall events are small. CONCLUSIONS: In the context of shrubification across tundra systems, our findings suggest that lichens will continue to have a large effect on microclimate until they are fully excluded, at which point microclimate extremes will increase greatly.

No genetic erosion after five generations for Impatiens glandulifera populations across the invaded range in Europe.

BACKGROUND: The observation that many alien species become invasive despite low genetic diversity has long been considered the 'genetic paradox' in invasion biology. This paradox is often resolved through the temporal buildup genetic diversity through multiple introduction events. These temporal dynamics in genetic diversity are especially important for annual invasive plants that lack a persistent seed bank, for which population persistence is strongly dependent on consecutive seed 're-establishment' in each growing season. Theory predicts that the number of seeds during re-establishment, and the levels of among-population gene flow can strongly affect recolonization dynamics, resulting in either an erosion or build-up of population genetic diversity through time. This study focuses on temporal changes in the population genetic structure of the annual invasive plant Impatiens glandulifera across Europe. We resampled 13 populations in 6 regions along a 1600 km long latitudinal gradient from northern France to central Norway after 5 years, and assessed population genetic diversity with 9 microsatellite markers. RESULTS: Our study suggests sufficiently high numbers of genetically diverse founders during population re-establishment, which prevent the erosion of local genetic diversity. We furthermore observe that I. glandulifera experiences significant among-population gene flow, gradually resulting in higher genetic diversity and lower overall genetic differentiation through time. Nonetheless, moderate founder effects concerning population genetic composition (allele frequencies) were evident, especially for smaller populations. Despite the initially low genetic diversity, this species seems to be successful at persisting across its invaded range, and will likely continue to build up higher genetic diversity at the local scale.

Late Quaternary climate legacies in contemporary plant functional composition.

The functional composition of plant communities is commonly thought to be determined by contemporary climate. However, if rates of climate-driven immigration and/or exclusion of species are slow, then contemporary functional composition may be explained by paleoclimate as well as by contemporary climate. We tested this idea by coupling contemporary maps of plant functional trait composition across North and South America to paleoclimate means and temporal variation in temperature and precipitation from the Last Interglacial (120 ka) to the present. Paleoclimate predictors strongly improved prediction of contemporary functional composition compared to contemporary climate predictors, with a stronger influence of temperature in North America (especially during periods of ice melting) and of precipitation in South America (across all times). Thus, climate from tens of thousands of years ago influences contemporary functional composition via slow assemblage dynamics.

Predictability in community dynamics.

The coupling between community composition and climate change spans a gradient from no lags to strong lags. The no-lag hypothesis is the foundation of many ecophysiological models, correlative species distribution modelling and climate reconstruction approaches. Simple lag hypotheses have become prominent in disequilibrium ecology, proposing that communities track climate change following a fixed function or with a time delay. However, more complex dynamics are possible and may lead to memory effects and alternate unstable states. We develop graphical and analytic methods for assessing these scenarios and show that these dynamics can appear in even simple models. The overall implications are that (1) complex community dynamics may be common and (2) detailed knowledge of past climate change and community states will often be necessary yet sometimes insufficient to make predictions of a community's future state.

Biotic and abiotic drivers of intraspecific trait variation within plant populations of three herbaceous plant species along a latitudinal gradient.

BACKGROUND: The importance of intraspecific trait variation (ITV) is increasingly acknowledged among plant ecologists. However, our understanding of what drives ITV between individual plants (ITVBI) at the population level is still limited. Contrasting theoretical hypotheses state that ITVBI can be either suppressed (stress-reduced plasticity hypothesis) or enhanced (stress-induced variability hypothesis) under high abiotic stress. Similarly, other hypotheses predict either suppressed (niche packing hypothesis) or enhanced ITVBI (individual variation hypothesis) under high niche packing in species rich communities. In this study we assess the relative effects of both abiotic and biotic niche effects on ITVBI of four functional traits (leaf area, specific leaf area, plant height and seed mass), for three herbaceous plant species across a 2300 km long gradient in Europe. The study species were the slow colonizing Anemone nemorosa, a species with intermediate colonization rates, Milium effusum, and the fast colonizing, non-native Impatiens glandulifera. RESULTS: Climatic stress consistently increased ITVBI across species and traits. Soil nutrient stress, on the other hand, reduced ITVBI for A. nemorosa and I. glandulifera, but had a reversed effect for M. effusum. We furthermore observed a reversed effect of high niche packing on ITVBI for the fast colonizing non-native I. glandulifera (increased ITVBI), as compared to the slow colonizing native A. nemorosa and M. effusum (reduced ITVBI). Additionally, ITVBI in the fast colonizing species tended to be highest for the vegetative traits plant height and leaf area, but lowest for the measured generative trait seed mass. CONCLUSIONS: This study shows that stress can both reduce and increase ITVBI, seemingly supporting both the stress-reduced plasticity and stress-induced variability hypotheses. Similarly, niche packing effects on ITVBI supported both the niche packing hypothesis and the individual variation hypothesis. These results clearly illustrates the importance of simultaneously evaluating both abiotic and biotic factors on ITVBI. This study adds to the growing realization that within-population trait variation should not be ignored and can provide valuable ecological insights.

Plant movements and climate warming: intraspecific variation in growth responses to nonlocal soils.

Most range shift predictions focus on the dispersal phase of the colonization process. Because moving populations experience increasingly dissimilar nonclimatic environmental conditions as they track climate warming, it is also critical to test how individuals originating from contrasting thermal environments can establish in nonlocal sites. We assess the intraspecific variation in growth responses to nonlocal soils by planting a widespread grass of deciduous forests (Milium effusum) into an experimental common garden using combinations of seeds and soil sampled in 22 sites across its distributional range, and reflecting movement scenarios of up to 1600 km. Furthermore, to determine temperature and forest-structural effects, the plants and soils were experimentally warmed and shaded. We found significantly positive effects of the difference between the temperature of the sites of seed and soil collection on growth and seedling emergence rates. Migrant plants might thus encounter increasingly favourable soil conditions while tracking the isotherms towards currently 'colder' soils. These effects persisted under experimental warming. Rising temperatures and light availability generally enhanced plant performance. Our results suggest that abiotic and biotic soil characteristics can shape climate change-driven plant movements by affecting growth of nonlocal migrants, a mechanism which should be integrated into predictions of future range shifts.

Trade-offs in biodiversity and ecosystem services between edges and interiors in European forests.

Forest biodiversity and ecosystem services are hitherto predominantly quantified in forest interiors, well away from edges. However, these edges also represent a substantial proportion of the global forest cover. Here we quantified plant biodiversity and ecosystem service indicators in 225 plots along forest edge-to-interior transects across Europe. We found strong trade-offs: phylogenetic diversity (evolutionary measure of biodiversity), proportion of forest specialists, decomposition and heatwave buffering increased towards the interior, whereas species richness, nectar production potential, stemwood biomass and tree regeneration decreased. These trade-offs were mainly driven by edge-to-interior structural differences. As fragmentation continues, recognizing the role of forest edges is crucial for integrating biodiversity and ecosystem service considerations into sustainable forest management and policy.

Variation in insect herbivory across an urbanization gradient: The role of abiotic factors and leaf secondary metabolites.

Urbanization impacts plant-herbivore interactions, which are crucial for ecosystem functions such as carbon sequestration and nutrient cycling. While some studies have reported reductions in insect herbivory in urban areas (relative to rural or natural forests), this trend is not consistent and the underlying causes for such variation remain unclear. We conducted a continental-scale study on insect herbivory along urbanization gradients for three European tree species: Quercus robur, Tilia cordata, and Fraxinus excelsior, and further investigated their biotic and abiotic correlates to get at mechanisms. To this end, we quantified insect leaf herbivory and foliar secondary metabolites (phenolics, terpenoids, alkaloids) for 176 trees across eight European cities. Additionally, we collected data on microclimate (air temperature) and soil characteristics (pH, carbon, nutrients) to test for abiotic correlates of urbanization effects directly or indirectly (through changes in plant secondary chemistry) linked to herbivory. Our results showed that urbanization was negatively associated with herbivory for Q. robur and F. excelsior, but not for T. cordata. In addition, urbanization was positively associated with secondary metabolite concentrations, but only for Q. robur. Urbanization was positively associated with air temperature for Q. robur and F. excelsior, and negatively with soil nutrients (magnesium) in the case of F. excelsior, but these abiotic variables were not associated with herbivory. Contrary to expectations, we found no evidence for indirect effects of abiotic factors via plant defences on herbivory for either Q. robur or F. excelsior. Additional biotic or abiotic drivers must therefore be accounted for to explain observed urbanization gradients in herbivory and their interspecific variation.

Plant community type and small-scale disturbances, but not altitude, influence the invasibility in subarctic ecosystems.

Little of our knowledge about invasibility comes from arctic and alpine ecosystems, despite increasing plant migration and invasion in those regions. Here, we examine how community type, altitude, and small-scale disturbances affect invasibility in a subarctic ecosystem. Over a period of 4 yr, we studied seedling emergence and establishment in 17 species sown in gaps or undisturbed vegetation in four subarctic community types (Salix scrub, meadow, rich heath, poor heath) along an elevation gradient. Invasibility was lowest in rich heath and highest in Salix scrub. Small disturbances significantly increased the invasibility in most communities, thereby showing the importance of biotic resistance to invasion in subarctic regions. Unexpectedly, invasibility did not decrease with increasing elevation, and it was also not related to summer temperature. Our data suggest that biotic resistance might be more important than abiotic stress for invasibility in subarctic tundra and that low temperatures do not necessarily limit seedling establishment at high altitudes. High elevations are therefore potentially more vulnerable to invasion than was originally thought. Changes in community composition as a result of species migration or invasion are most likely to occur in Salix scrub and meadow, whereas Empetrum-dominated rich heath will largely remain unchanged.

Correlations between physical and chemical defences in plants: tradeoffs, syndromes, or just many different ways to skin a herbivorous cat?

Most plant species have a range of traits that deter herbivores. However, understanding of how different defences are related to one another is surprisingly weak. Many authors argue that defence traits trade off against one another, while others argue that they form coordinated defence syndromes. We collected a dataset of unprecedented taxonomic and geographic scope (261 species spanning 80 families, from 75 sites across the globe) to investigate relationships among four chemical and six physical defences. Five of the 45 pairwise correlations between defence traits were significant and three of these were tradeoffs. The relationship between species' overall chemical and physical defence levels was marginally nonsignificant (P = 0.08), and remained nonsignificant after accounting for phylogeny, growth form and abundance. Neither categorical principal component analysis (PCA) nor hierarchical cluster analysis supported the idea that species displayed defence syndromes. Our results do not support arguments for tradeoffs or for coordinated defence syndromes. Rather, plants display a range of combinations of defence traits. We suggest this lack of consistent defence syndromes may be adaptive, resulting from selective pressure to deploy a different combination of defences to coexisting species.

Decoupled phenotypic variation between floral and vegetative traits: distinguishing between developmental and environmental correlations.

BACKGROUND AND AIMS: In species with specialized pollination, floral traits are expected to be relatively invariant and decoupled from the phenotypic variation affecting vegetative traits. However, inferring the degree of decoupling between morphological characters from patterns of phenotypic correlations is difficult because phenotypic correlations result from the superimposition of several sources of covariance. In this study it is hypothesized that, in some cases, negative environmental correlations generated by non-congruent reaction norms across traits overshadow positive developmental correlations and generate a decoupling of the phenotypic variation between vegetative and floral traits. METHODS: To test this hypothesis, Campanula rotundifolia were grown from two distinct populations under two temperature treatments, and patterns of correlation were analysed between leaf size and flower size within and among treatments. KEY RESULTS: Flower size was less sensitive to temperature variation than leaf size. Furthermore, flower size and leaf size showed temperature-induced reaction norms in opposite directions. Flower size decreased with an increasing temperature, while leaf size increased. Consequently, among treatments, correlations between leaf size and flower size were negative or absent, while, within treatments, these correlations were positive or absent in the cold and warm environments, respectively. CONCLUSIONS: These results confirm that the decoupling of the phenotypic variation between vegetative and floral traits can be dependent on the environment. They also underline the importance of distinguishing sources of phenotypic covariance when testing hypotheses about phenotypic integration.

The response of forest plant regeneration to temperature variation along a latitudinal gradient.

BACKGROUND AND AIMS: The response of forest herb regeneration from seed to temperature variations across latitudes was experimentally assessed in order to forecast the likely response of understorey community dynamics to climate warming. METHODS: Seeds of two characteristic forest plants (Anemone nemorosa and Milium effusum) were collected in natural populations along a latitudinal gradient from northern France to northern Sweden and exposed to three temperature regimes in growth chambers (first experiment). To test the importance of local adaptation, reciprocal transplants were also made of adult individuals that originated from the same populations in three common gardens located in southern, central and northern sites along the same gradient, and the resulting seeds were germinated (second experiment). Seedling establishment was quantified by measuring the timing and percentage of seedling emergence, and seedling biomass in both experiments. KEY RESULTS: Spring warming increased emergence rates and seedling growth in the early-flowering forb A. nemorosa. Seedlings of the summer-flowering grass M. effusum originating from northern populations responded more strongly in terms of biomass growth to temperature than southern populations. The above-ground biomass of the seedlings of both species decreased with increasing latitude of origin, irrespective of whether seeds were collected from natural populations or from the common gardens. The emergence percentage decreased with increasing home-away distance in seeds from the transplant experiment, suggesting that the maternal plants were locally adapted. CONCLUSIONS: Decreasing seedling emergence and growth were found from the centre to the northern edge of the distribution range for both species. Stronger responses to temperature variation in seedling growth of the grass M. effusum in the north may offer a way to cope with environmental change. The results further suggest that climate warming might differentially affect seedling establishment of understorey plants across their distribution range and thus alter future understorey plant dynamics.

How do bryophytes govern generative recruitment of vascular plants?

Interactions between vascular plants and bryophytes determine plant community composition in many ecosystems. Yet, little is known about the importance of interspecific differences between bryophytes with respect to their effects on vascular plants. We compared the extent to which species-specific bryophyte effects on vascular plant generative recruitment depend on the following underlying mechanisms: allelopathy, mechanical obstruction, soil moisture and temperature control. We sowed 10 vascular plant species into monospecific mats of six chemically and structurally diverse bryophytes, and examined 1-yr seedling recruitment. Allelopathic effects were also assessed in a laboratory phyto-assay. Although all bryophytes suppressed vascular plant regeneration, there were significant differences between the bryophyte species. The lack of interactions indicated the absence of species-specific adaptations of vascular plants for recruitment in bryophyte mats. Differences between bryophyte species were best explained by alterations in temperature regime under bryophyte mats, mostly by reduced temperature amplitudes during germination. The temperature regime under bryophyte mats was well predicted by species-specific bryophyte cushion thickness. The fitness of established seedlings was not affected by the presence of bryophytes. Our results suggest that climatically or anthropogenically driven changes in the species' composition of bryophyte communities have knock-on effects on vascular plant populations via generative reproduction.

Putting plant resistance traits on the map: a test of the idea that plants are better defended at lower latitudes.

• It has long been believed that plant species from the tropics have higher levels of traits associated with resistance to herbivores than do species from higher latitudes. A meta-analysis recently showed that the published literature does not support this theory. However, the idea has never been tested using data gathered with consistent methods from a wide range of latitudes. • We quantified the relationship between latitude and a broad range of chemical and physical traits across 301 species from 75 sites world-wide. • Six putative resistance traits, including tannins, the concentration of lipids (an indicator of oils, waxes and resins), and leaf toughness were greater in high-latitude species. Six traits, including cyanide production and the presence of spines, were unrelated to latitude. Only ash content (an indicator of inorganic substances such as calcium oxalates and phytoliths) and the properties of species with delayed greening were higher in the tropics. • Our results do not support the hypothesis that tropical plants have higher levels of resistance traits than do plants from higher latitudes. If anything, plants have higher resistance toward the poles. The greater resistance traits of high-latitude species might be explained by the greater cost of losing a given amount of leaf tissue in low-productivity environments.

Strong microsite control of seedling recruitment in tundra.

The inclusion of environmental variation in studies of recruitment is a prerequisite for realistic predictions of the responses of vegetation to a changing environment. We investigated how seedling recruitment is affected by seed availability and microsite quality along a steep environmental gradient in dry tundra. A survey of natural seed rain and seedling density in vegetation was combined with observations of the establishment of 14 species after sowing into intact or disturbed vegetation. Although seed rain density was closely correlated with natural seedling establishment, the experimental seed addition showed that the microsite environment was even more important. For all species, seedling emergence peaked at the productive end of the gradient, irrespective of the adult niches realized. Disturbance promoted recruitment at all positions along the environmental gradient, not just at high productivity. Early seedling emergence constituted the main temporal bottleneck in recruitment for all species. Surprisingly, winter mortality was highest at what appeared to be the most benign end of the gradient. The results highlight that seedling recruitment patterns are largely determined by the earliest stages in seedling emergence, which again are closely linked to microsite quality. A fuller understanding of microsite effects on recruitment with implications for plant community assembly and vegetation change is provided.

Drivers of carbon stocks in forest edges across Europe.

Forests play a key role in global carbon cycling and sequestration. However, the potential for carbon drawdown is affected by forest fragmentation and resulting changes in microclimate, nutrient inputs, disturbance and productivity near edges. Up to 20% of the global forested area lies within 100 m of an edge and, even in temperate forests, knowledge on how edge conditions affect carbon stocks and how far this influence penetrates into forest interiors is scarce. Here we studied carbon stocks in the aboveground biomass, forest floor and the mineral topsoil in 225 plots in deciduous forest edges across Europe and tested the impact of macroclimate, nitrogen deposition and smaller-grained drivers (e.g. microclimate) on these stocks. Total carbon and carbon in the aboveground biomass stock were on average 39% and 95% higher at the forest edge than 100 m into the interior. The increase in the aboveground biomass stock close to the edge was mainly related to enhanced nitrogen deposition. No edge influence was found for stocks in the mineral topsoil. Edge-to-interior gradients in forest floor carbon changed across latitude: carbon stocks in the forest floor were higher near the edge in southern Europe. Forest floor carbon decreased with increasing litter quality (i.e. high decomposition rate) and decreasing plant area index, whereas higher soil temperatures negatively affected the mineral topsoil carbon. Based on high-resolution forest fragmentation maps, we estimate that the additional carbon stored in deciduous forest edges across Europe amounts to not less than 183 Tg carbon, which is equivalent to the storage capacity of 1 million ha of additional forest. This study underpins the importance of including edge influences when quantifying the carbon stocks in temperate forests and stresses the importance of preserving natural forest edges and small forest patches with a high edge-to-interior surface area.

Community assembly in experimental grasslands: suitable environment or timely arrival?

It is hard to defend the view that biotic communities represent a simple and predictable response to the abiotic environment. Biota and the abiotic environment interact, and the environment of an individual certainly includes its neighbors and visitors in the community. The complexity of community assembly calls forth a quest for general principles, yet current results and theories on assembly rules differ widely. Using a grassland microcosm as a model system, we manipulated fertility, disturbance by defoliation, soil/microclimate, and arrival order of species belonging to two groups differing in functional attributes. We analyzed the outcome of community assembly dynamics in terms of species richness, invasibility, and species composition. The analyses revealed strong environmental control over species richness and invasibility. Species composition was mainly determined by the arrival order of species, indicating that historical contingency may change the outcome of community assembly. The probability for multiple equilibria appeared to increase with productivity and environmental stability. The importance of arrival order offers an explanation of the difficulties in predicting local occurrences of species in the field. In our experiment, variation in fertility and disturbance was controlling colonization with predictable effects on emergent community properties such as species richness. The key mechanism is suggested to be asymmetric competition, and our results show that this mechanism is relatively insensitive to the species through which it works. While our analyses indicate a positive and significant correlation between richness and invasibility, the significance disappears after accounting for the effect of the environment. The importance of arrival order (historical contingency) and environmental control supports the assumption of the unified neutral theory that different species within a trophic level can be considered functionally equivalent when it comes to community assembly. However, our results indicate that variation in asymmetric competition is the key factor determining the richness of the resulting communities, and this is far from neutral.