Light and warming drive forest understorey community development in different environments

Plant community composition and functional traits respond to chronic drivers such as climate change and nitrogen (N) deposition. In contrast, pulse disturbances from ecosystem management can additionally change resources and conditions. Community responses to combined environmental changes may further depend on land‐use legacies. Disentangling the relative importance of these global change drivers is necessary to improve predictions of future plant communities. We performed a multifactor global change experiment to disentangle drivers of herbaceous plant community trajectories in a temperate deciduous forest. Communities of five species, assembled from a pool of 15 forest herb species with varying ecological strategies, were grown in 384 mesocosms on soils from ancient forest (forested at least since 1850) and postagricultural forest (forested since 1950) collected across Europe. Mesocosms were exposed to two‐level full‐factorial treatments of warming, light addition (representing changing forest management) and N enrichment. We measured plant height, specific leaf area (SLA) and species cover over the course of three growing seasons. Increasing light availability followed by warming reordered the species towards a taller herb community, with limited effects of N enrichment or the forest land‐use history. Two‐way interactions between treatments and incorporating intraspecific trait variation (ITV) did not yield additional inference on community height change. Contrastingly, community SLA differed when considering ITV along with species reordering, which highlights ITV’s importance for understanding leaf morphology responses to nutrient enrichment in dark conditions. Contrary to our expectations, we found limited evidence of land‐use legacies affecting community responses to environmental changes, perhaps because dispersal limitation was removed in the experimental design. These findings can improve predictions of community functional trait responses to global changes by acknowledging ITV, and subtle changes in light availability. Adaptive forest management to impending global change could benefit the restoration and conservation of understorey plant communities by reducing the light availability.     

Plant functional trait response to environmental drivers across European temperate forest understorey communities

• Functional traits respond to environmental drivers, hence evaluating trait‐environment relationships across spatial environmental gradients can help to understand how multiple drivers influence plant communities. Global‐change drivers such as changes in atmospheric nitrogen deposition occur worldwide, but affect community trait distributions at the local scale, where resources (e.g. light availability) and conditions (e.g. soil pH) also influence plant communities.
• We investigate how multiple environmental drivers affect community trait responses related to resource acquisition (plant height, specific leaf area (SLA), woodiness, and mycorrhizal status) and regeneration (seed mass, lateral spread) of European temperate deciduous forest understoreys. We sampled understorey communities and derived trait responses across spatial gradients of global‐change drivers (temperature, precipitation, nitrogen deposition, and past land use), while integrating in‐situ plot measurements on resources and conditions (soil type, Olsen phosphorus (P), Ellenberg soil moisture, light, litter mass, and litter quality).
• Among the global‐change drivers, mean annual temperature strongly influenced traits related to resource acquisition. Higher temperatures were associated with taller understoreys producing leaves with lower SLA, and a higher proportional cover of woody and obligate mycorrhizal (OM) species. Communities in plots with higher Ellenberg soil moisture content had smaller seeds and lower proportional cover of woody and OM species. Finally, plots with thicker litter layers hosted taller understoreys with larger seeds and a higher proportional cover of OM species.
• Our findings suggest potential community shifts in temperate forest understoreys with global warming, and highlight the importance of local resources and conditions as well as global‐change drivers for community trait variation.

     

Taller and larger: shifts in Arctic tundra leaf traits after 16 years of experimental warming

Understanding plant trait responses to elevated temperatures in the Arctic is critical in light of recent and continuing climate change, especially because these traits act as key mechanisms in climate‐vegetation feedbacks. Since 1992, we have artificially warmed three plant communities at Alexandra Fiord, Nunavut, Canada (79°N). In each of the communities, we used open‐top chambers (OTCs) to passively warm vegetation by 1–2 °C. In the summer of 2008, we investigated the intraspecific trait responses of five key species to 16 years of continuous warming. We examined eight traits that quantify different aspects of plant performance: leaf size, specific leaf area (SLA), leaf dry matter content (LDMC), plant height, leaf carbon concentration, leaf nitrogen concentration, leaf carbon isotope discrimination (LCID), and leaf δ¹⁵N. Long‐term artificial warming affected five traits, including at least one trait in every species studied. The evergreen shrub Cassiope tetragona responded most frequently (increased leaf size and plant height/decreased SLA, leaf carbon concentration, and LCID), followed by the deciduous shrub Salix arctica (increased leaf size and plant height/decreased SLA) and the evergreen shrub Dryas integrifolia (increased leaf size and plant height/decreased LCID), the forb Oxyria digyna (increased leaf size and plant height), and the sedge Eriophorum angustifolium spp. triste (decreased leaf carbon concentration). Warming did not affect δ¹⁵N, leaf nitrogen concentration, or LDMC. Overall, growth traits were more sensitive to warming than leaf chemistry traits. Notably, we found that responses to warming were sustained, even after many years of treatment. Our work suggests that tundra plants in the High Arctic will show a multifaceted response to warming, often including taller shoots with larger leaves.     

Seasonality,species richness and poor dispersion mediate intraspecific trait variability in stonefly community responses along an elevational gradient

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Intraspecific variability of specific leaf area fosters the persistence of understorey specialists across a light availability gradient

• Forest understorey plants are sensitive to light availability, and different species groups can respond differently to changing light conditions. A plant trait tightly linked to light capture is specific leaf area (SLA). Studies considering the relative role of within‐ and among‐species SLA variation across different species groups (e.g. specialists and generalists) are rarely implemented in temperate forest understories varying in their maturity.
• We examined community‐level SLA patterns of beech forest understories along a light availability gradient, and for habitat specialists and generalists separately. We then disentangled and quantified the contribution of intraspecific trait variability and interspecific trait differences in shaping SLA patterns.
• We revealed that the increase in community‐level SLA with decreasing light availability was primarily driven by beech forest specialists (and, to a lesser extent, by forest generalists), and this pattern was mainly determined by specialists’ high intraspecific variability. Community‐level SLA was therefore formed by different responses at different organizational levels, i.e. within and among species, and for separate species groups.
• This study provides insights into factors shaping the shade tolerance strategy in beech forest understorey plants; specialists persistence under putative less favourable conditions (i.e. high irradiation) may be fostered by their ability to adjust their light capture strategies intraspecifically.

     

Differential effect of drought regimes on the seedling performance of six floodplain grassland species

• The performance of seedlings is crucial for the survival and persistence of plant populations. Although drought frequently occurs in floodplains and can cause seedling mortality, studies on the effects of drought on seedlings of floodplain grasslands are scarce. We tested the hypotheses that drought reduces aboveground biomass, total biomass, plant height, number of leaves, leaf area and specific leaf area (SLA), and increases root biomass and root‐mass fraction (RMF) and that seedlings from species of wet floodplain grasslands are more affected by drought than species of dry grasslands.
• In a greenhouse study, we exposed seedlings of three confamilial pairs of species (Pimpinella saxifraga, Selinum carvifolia, Veronica teucrium, Veronica maritima, Sanguisorba minor, Sanguisorba officinalis) to increasing drought treatments. Within each plant family, one species is characteristic of wet and one of dry floodplain grasslands, confamilial in order to avoid phylogenetic bias of the results.
• In accordance with our hypotheses, drought conditions reduced aboveground biomass, total biomass, plant height, number of leaves and leaf area. Contrary to our hypotheses, drought conditions increased SLA and decreased root biomass and RMF of seedlings. Beyond the effects of the families, the results were species‐specific (V. maritima being the most sensitive species) and habitat‐specific. Species indicative of wet floodplain grasslands appear to be more sensitive to drought than species indicative of dry grasslands.
• Because of species‐ and habitat‐specific responses to reduced water availability, future drought periods due to climate change may severely affect some species from dry and wet habitats, while others may be unaffected.

     

An intraspecific application of the leaf‐height‐seed ecology strategy scheme to forest herbs along a latitudinal gradient

We applied the leaf‐height‐seed (LHS) ecology strategy scheme (a combination of three ecologically important traits: specific leaf area (SLA), seed mass and plant height) intraspecifically to two widespread European forest herbs along a latitudinal gradient. The aims of this study were to quantify LHS trait variation, disentangle the environmental factors affecting these traits and compare the within‐species LHS trait relationships with latitude to previously established cross‐species comparisons. We measured LHS traits in 41 Anemone nemorosa and 44 Milium effusum populations along a 1900–2300 km latitudinal gradient from N France to N Sweden. We then applied multilevel models to identify the effects of regional (temperature, latitude) and local (soil fertility and acidity, overstorey canopy cover) environmental factors on LHS traits. Both species displayed a significant 4% increase in plant height with every degree northward shift (almost a two‐fold plant height difference between the southernmost and northernmost populations). Neither seed mass nor SLA showed a significant latitudinal cline. Temperature had a large effect on the three LHS traits of Anemone. Latitude, canopy cover and soil nutrients were related to the SLA and plant height of Milium. None of the investigated variables appeared to be related to the seed mass of Milium. The variation in LHS traits indicates that the ecological strategy determined by the position of each population in this three‐factor triangle is not constant along the latitudinal gradient. The significant increase in plant height suggests greater competitive abilities for both species in the northernmost populations. We also found that the studied environmental factors affected the LHS traits of the two species on various scales: spring‐flowering Anemone was affected more by temperature, whereas early‐summer flowering Milium was affected more by local and other latitude‐related factors. Finally, previously reported cross‐species correlations between LHS traits and latitude were generally unsupported by our within‐species approach.     

A Race for Space? How Sphagnum fuscum stabilizes vegetation composition during long‐term climate manipulations

Strong climate warming is predicted at higher latitudes this century, with potentially major consequences for productivity and carbon sequestration. Although northern peatlands contain one‐third of the world's soil organic carbon, little is known about the long‐term responses to experimental climate change of vascular plant communities in these Sphagnum‐dominated ecosystems. We aimed to see how long‐term experimental climate manipulations, relevant to different predicted future climate scenarios, affect total vascular plant abundance and species composition when the community is dominated by mosses. During 8 years, we investigated how the vascular plant community of a Sphagnum fuscum‐dominated subarctic peat bog responded to six experimental climate regimes, including factorial combinations of summer as well as spring warming and a thicker snow cover. Vascular plant species composition in our peat bog was more stable than is typically observed in (sub)arctic experiments: neither changes in total vascular plant abundance, nor in individual species abundances, Shannon's diversity or evenness were found in response to the climate manipulations. For three key species (Empetrum hermaphroditum, Betula nana and S. fuscum) we also measured whether the treatments had a sustained effect on plant length growth responses and how these responses interacted. Contrasting with the stability at the community level, both key shrubs and the peatmoss showed sustained positive growth responses at the plant level to the climate treatments. However, a higher percentage of moss‐encroached E. hermaphroditum shoots and a lack of change in B. nana net shrub height indicated encroachment by S. fuscum, resulting in long‐term stability of the vascular community composition: in a warmer world, vascular species of subarctic peat bogs appear to just keep pace with growing Sphagnum in their race for space. Our findings contribute to general ecological theory by demonstrating that community resistance to environmental changes does not necessarily mean inertia in vegetation response.     

Potential effects of warming and drying on peatland plant community composition

Boreal peatlands may be particularly vulnerable to climate change, because temperature regimes that currently constrain biological activity in these regions are predicted to increase substantially within the next century. Changes in peatland plant community composition in response to climate change may alter nutrient availability, energy budgets, trace gas fluxes, and carbon storage. We investigated plant community response to warming and drying in a field mesocosm experiment in northern Minnesota, USA. Large intact soil monoliths removed from a bog and a fen received three infrared warming treatments crossed with three water‐table treatments (n = 3) for five years. Foliar cover of each species was estimated annually. In the bog, increases in soil temperature and decreases in water‐table elevation increased cover of shrubs by 50% and decreased cover of graminoids by 50%. The response of shrubs to warming was distinctly species‐specific, and ranged from increases (for Andromeda glaucophylla) to decreases (for Kalmia polifolia). In the fens, changes in plant cover were driven primarily by changes in water‐table elevation, and responses were species‐ and lifeform‐specific: increases in water‐table elevation increased cover of graminoids – in particular Carex lasiocarpa and Carex livida– as well as mosses. In contrast, decreases in water‐table elevation increased cover of shrubs, in particular A. glaucophylla and Chamaedaphne calyculata. The differential and sometimes opposite response of species and lifeforms to the treatments suggest that the structure and function of both bog and fen plant communities will change – in different directions or at different magnitudes – in response to warming and/or changes in water‐table elevation that may accompany regional or global climate change.     

Drought mildly reduces plant dominance in a temperate prairie ecosystem across years

1. Shifts in dominance and species reordering can occur in response to global change. However, it is not clear how altered precipitation and disturbance regimes interact to affect species composition and dominance.
2. We explored community‐level diversity and compositional similarity responses, both across and within years, to a manipulated precipitation gradient and annual clipping in a mixed‐grass prairie in Oklahoma, USA. We imposed seven precipitation treatments (five water exclusion levels [?20%, ?40%, ?60%, ?80%, and ?100%], water addition [+50%], and control [0% change in precipitation]) year‐round from 2016 to 2018 using fixed interception shelters. These treatments were crossed with annual clipping to mimic hay harvest.
3. We found that community‐level responses were influenced by precipitation across time. For instance, plant evenness was enhanced by extreme drought treatments, while plant richness was marginally promoted under increased precipitation.
4. Clipping promoted species gain resulting in greater richness within each experimental year. Across years, clipping effects further reduced the precipitation effects on community‐level responses (richness and evenness) at both extreme drought and added precipitation treatments.
5. Synthesis: Our results highlight the importance of studying interactive drivers of change both within versus across time. For instance, clipping attenuated community‐level responses to a gradient in precipitation, suggesting that management could buffer community‐level responses to drought. However, precipitation effects were mild and likely to accentuate over time to produce further community change.

     

Drought responses,phenotypic plasticity and survival of Mediterranean species in two different microclimatic sites

• Climate models predict a further drying of the Mediterranean summer. One way for plant species to persist during such climate changes is through acclimation. Here, we determine the extent to which trait plasticity in response to drought differs between species and between sites, and address the question whether there is a trade‐off between drought survival and phenotypic plasticity.
• Throughout the summer we measured physiological traits (photosynthesis – A_max, stomatal conductance – g_s, transpiration – E, leaf water potential – ψl) and structural traits (specific leaf area – SLA, leaf density – LD, leaf dry matter content – LDMC, leaf relative water content – LRWC) of leaves of eight woody species in two sites with slightly different microclimate (north‐ versus south‐facing slopes) in southern Spain. Plant recovery and survival was estimated after the summer drought period.
• We found high trait variability between species. In most variables, phenotypic plasticity was lower in the drier site. Phenotypic plasticity of SLA and LDMC correlated negatively with drought survival, which suggests a trade‐off between them. On the other hand, high phenotypic plasticity of SLA and LDMC was positively related to traits associated with rapid recovery and growth after the drought period.
• Although phenotypic plasticity is generally seen as favourable during stress conditions, here it seemed beneficial for favourable conditions. We propose that in environments with fluctuating drought periods there can be a trade‐off between drought survival and growth during favourable conditions. When climate become drier, species with high drought survival but low phenotypic plasticity might be selected for.

     

Heterogeneous flows foster heterogeneous assemblages: relationships between functional diversity and hydrological heterogeneity in riparian plant communities

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Grazing effects on intraspecific trait variability vary with changing precipitation patterns in Mongolian rangelands

Functional traits are proxies for plant physiology and performance, which do not only differ between species but also within species. In this work, we hypothesized that (a) with increasing precipitation, the percentage of focal species which significantly respond to changes in grazing intensity increases, while under dry conditions, climate‐induced stress is so high that plant species hardly respond to any changes in grazing intensity and that (b) the magnitude with which species change their trait values in response to grazing, reflected by coefficients of variation (CVs), increases with increasing precipitation. Chosen plant traits were canopy height, plant width, specific leaf area (SLA), chlorophyll fluorescence, performance index, stomatal pore area index (SPI), and individual aboveground biomass of 15 species along a precipitation gradient with different grazing intensities in Mongolian rangelands. We used linear models for each trait to assess whether the percentage of species that respond to grazing changes along the precipitation gradient. To test the second hypothesis, we assessed the magnitude of intraspecific trait variability (ITV) response to grazing, per species, trait, and precipitation level by calculating CVs across the different grazing intensities. ITV was most prominent for SLA and SPI under highest precipitation, confirming our first hypothesis. Accordingly, CVs of canopy height, SPI, and SLA increased with increasing precipitation, partly confirming our second hypothesis. CVs of the species over all traits increased with increasing precipitation only for three species. This study shows that it remains challenging to predict how plant performance will shift under changing environmental conditions based on their traits alone. In this context, the implications for the use of community‐weighted mean trait values are discussed, as not only species abundances change in response to changing environmental conditions, but also values of traits considerably change. Including this aspect in further studies will improve our understanding of processes acting within and among communities.     

Resource partitioning between ungulate populations in arid environments

Herbivores are major drivers of ecosystem structure, diversity, and function. Resilient ecosystems therefore require viable herbivore populations in a sustainable balance with environmental resource availability. This balance is becoming harder to achieve, with increasingly threatened species reliant on small protected areas in increasingly harsh and unpredictable environments. Arid environments in North Africa exemplify this situation, featuring a biologically distinct species assemblage exposed to extreme and volatile conditions, including habitat loss and climate change‐associated threats. Here, we implement an integrated likelihood approach to relate scimitar‐horned oryx (Oryx dammah) and dorcas gazelle (Gazella dorcas) density, via dung distance sampling, to habitat, predator, and geographic correlates in Dghoumes National Park, Tunisia. We show how two threatened sympatric ungulates partition resources on the habitat axis, exhibiting nonuniform responses to the same vegetation gradient. Scimitar‐horned oryx were positively associated with plant species richness, selecting for vegetated ephemeral watercourses (wadis) dominated by herbaceous cover. Conversely, dorcas gazelle were negatively associated with vegetation density (herbaceous height, litter cover, and herbaceous cover), selecting instead for rocky plains with sparse vegetation. We suggest that adequate plant species richness should be a prerequisite for areas proposed for future ungulate reintroductions in arid and semi‐arid environments. This evidence will inform adaptive management of reintroduced ungulates in protected environments, helping managers and planners design sustainable ecosystems and effective conservation programs.     

Plasticity in seedling morphology,biomass allocation and physiology among ten temperate tree species in response to shade is related to shade tolerance and not leaf habit

• Mechanisms of shade tolerance in tree seedlings, and thus growth in shade, may differ by leaf habit and vary with ontogeny following seed germination. To examine early responses of seedlings to shade in relation to morphological, physiological and biomass allocation traits, we compared seedlings of 10 temperate species, varying in their leaf habit (broadleaved versus needle‐leaved) and observed tolerance to shade, when growing in two contrasting light treatments – open (about 20% of full sunlight) and shade (about 5% of full sunlight).
• We analyzed biomass allocation and its response to shade using allometric relationships. We also measured leaf gas exchange rates and leaf N in the two light treatments.
• Compared to the open treatment, shading significantly increased traits typically associated with high relative growth rate (RGR) – leaf area ratio (LAR), specific leaf area (SLA), and allocation of biomass into leaves, and reduced seedling mass and allocation to roots, and net assimilation rate (NAR). Interestingly, RGR was not affected by light treatment, likely because of morphological and physiological adjustments in shaded plants that offset reductions of in situ net assimilation of carbon in shade. Leaf area‐based rates of light‐saturated leaf gas exchange differed among species groups, but not between light treatments, as leaf N concentration increased in concert with increased SLA in shade.
• We found little evidence to support the hypothesis of a increased plasticity of broadleaved species compared to needle‐leaved conifers in response to shade. However, an expectation of higher plasticity in shade‐intolerant species than in shade‐tolerant ones, and in leaf and plant morphology than in biomass allocation was supported across species of contrasting leaf habit.

     

Geographical variation and the role of climate in leaf traits of a relict tree species across its distribution in China

• Intraspecific trait variation and trait–climate relationships are crucial for understanding a species’ response to climate change. However, these phenomena have rarely been studied for tree species. Euptelea pleiospermum is a relict tree species with a wide distribution in China that offers a novel opportunity to examine such relationships.
• Here, we measured 13 leaf traits of E. pleiospermum in 20 sites across its natural distribution in China. We investigated the extent of trait variation at local and regional scales, and developed geographic and climate models to explain trait variation at the regional scale.
• We documented intraspecific trait variation among leaf traits of E. pleiospermum at local and regional scales. Five traits exhibited relatively high trait variation: leaf area, leaf density and three leaf economic traits (leaf dry matter content, specific leaf area [SLA] and leaf phosphorus concentration). Significant trait–geography correlations were mediated by local climate. Most leaf trait variation could be explained (from 24% to 64%) by geographic or climate variables, except leaf width, leaf thickness, leaf dry matter content and leaf length–width ratio. Latitude and temperature were the strongest predictors of trait variation throughout the distribution of E. pleiospermum in China, and temperature explained more leaf trait variation than precipitation. In particular, we showed that leaves had longer petiole lengths, higher SLA and lower densities in northern E. pleiospermum populations. We suggest that northern E. pleiospermum populations are adapting to higher latitudinal environments via high growth rate (higher SLA) and low construction investment strategies (lower leaf densities), benefitting northern migration.
• Overall, we demonstrate that intraspecific trait variation reflects E. pleiospermum response to the local environment. We call for consideration of intraspecific trait variation to examine specific climate response questions. In addition, provenance experiments using widely distributed species are needed to separate trait variation resulting from genetic differentiation and plastic responses to environmental change.

     

Size matters: nest colonization patterns for twig‐nesting ants

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Using plant traits to compare sward structure and composition of grass species across environmental gradients

Abstract. Plant traits which may give an indication of a plant's strategy for nutrient acquisition and regeneration are known for numerous grassland species. This study aimed to establish whether there is any relationship between two plant traits: specific leaf area (SLA) and number of reproductive tillers, and sward structural characteristics which influence herbage intake by grazers (bulk density and digestibility, leaf:stem ratio). Comparison is made for nutrient‐rich (Dactylis glomerata) and nutrient‐poor (Festuca rubra) grass species. We hypothesized that these traits are responsive to environmental gradients and also act on the processes of the ecosystem. Both grasses were compared with two P‐fertilizer rates in two localities (200 and 1300 m a.s.l.) which differed in their temperature:radiation ratios. For the vegetative phase SLA was well correlated with sward characteristics: D. glomerata, which has the higher SLA, has the lower bulk density and higher digestibility. The values of SLA and vegetation bulk density varied according to growing conditions (P‐rate and temperature:radiation ratio), but the ranking of the species remained the same because the phenotypic plasticity that exists for plant traits was also observed for sward structure and composition. That suggested the possibility of grouping natural grassland species for their relevant characteristics for grazers according to SLA values. Over the reproductive phase, the proportion of stems was well correlated to the percentage of reproductive tillers. However, the percentage of reproductive tillers was a very plastic trait for both species, depending on the growing conditions, and resulting in a density‐dependent effect, particularly for F. rubra. The species studied were too plastic and too similar in their regenerative strategy so that there is no unique relationship between percentage of reproductive tillers and stem proportion, regardless of the species and the growing conditions. The number of reproductive tillers is not a suitable plant trait which could be used to rank species for leaf and stem proportions in the sward.     

Conserving species‐rich predator assemblages strengthens natural pest control in a climate warming context

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The role of functional traits in species distributions revealed through a hierarchical model

Quantifying how functional traits relate to environmental gradients provides insight into mechanisms governing species distributions. Here, we bring together the fields of species distribution modelling and functional trait ecology with hierarchical modelling by explicitly incorporating traits into a multi‐species distribution model. We combined traits from the leaf‐height‐seed strategy scheme (specific leaf area (SLA), plant height and seed mass) with a distribution model for 20 eucalypt taxa in Victoria, Australia. The key insight of this approach is how traits modulate species responses to environmental gradients. The strongest link was between SLA and percent rock cover (species with low SLA had positive responses to rockiness, whereas high SLA species responded negatively to rockiness). We found evidence for complex yet potentially important interactions. For instance, the probability of species occurrence increased with rainfall and solar radiation on average yet the response varied depending on species height and SLA. Tall species were predicted to increase with rainfall and solar radiation across the range of SLA values (tall species with low SLA were especially sensitive to rainfall). Short species responded positively to rainfall and solar radiation only if they had low SLA. This framework readily accounts for interactions between combinations of traits and environmental variables unlike multi‐step approaches. Further application of this concept will contribute to a generalized mechanistic understanding of how traits influence species distributions along environmental gradients, with implications for understanding the response of species to global change.