Responses of leaf traits to climatic gradients: adaptive variation versus compositional shifts

Dynamic global vegetation models (DGVMs) typically rely on plant functional types (PFTs), which are assigned distinct environmental tolerances and replace one another progressively along environmental gradients. Fixed values of traits are assigned to each PFT; modelled trait variation along gradients is thus driven by PFT replacement. But empirical studies have revealed "universal" scaling relationships (quantitative trait variations with climate that are similar within and between species, PFTs and communities); and continuous, adaptive trait variation has been proposed to replace PFTs as the basis for next-generation DGVMs. Here we analyse quantitative leaf-trait variation on long temperature and moisture gradients in China with a view to understanding the relative importance of PFT replacement vs. continuous adaptive variation within PFTs. Leaf area (LA), specific leaf area (SLA), leaf dry matter content (LDMC) and nitrogen content of dry matter were measured on all species at 80 sites ranging from temperate to tropical climates and from dense forests to deserts. Chlorophyll fluorescence traits and carbon, phosphorus and potassium contents were measured at 47 sites. Generalized linear models were used to relate log-transformed trait values to growing-season temperature and moisture indices, with or without PFT identity as a predictor, and to test for differences in trait responses among PFTs. Continuous trait variation was found to be ubiquitous. Responses to moisture availability were generally similar within and between PFTs, but biophysical traits (LA, SLA and LDMC) of forbs and grasses responded differently from woody plants. SLA and LDMC responses to temperature were dominated by the prevalence of evergreen PFTs with thick, dense leaves at the warm end of the gradient. Nutrient (N, P and K) responses to climate gradients were generally similar within all PFTs. Area-based nutrients generally declined with moisture; Narea and Karea declined with temperature, but Parea increased with temperature. Although the adaptive nature of many of these trait-climate relationships is understood qualitatively, a key challenge for modelling is to predict them quantitatively. Models must take into account that community-level responses to climatic gradients can be influenced by shifts in PFT composition, such as the replacement of deciduous by evergreen trees, which may run either parallel or counter to trait variation within PFTs. The importance of PFT shifts varies among traits, being important for biophysical traits but less so for physiological and chemical traits. Finally, models should take account of the diversity of trait values that is found in all sites and PFTs, representing the "pool" of variation that is locally available for the natural adaptation of ecosystem function to environmental change.

The grass-free lawn: floral performance and management implications

Grass lawns are a ubiquitous feature of urban green-space throughout much of the temperate world. Species poor and intensively managed, lawns are ecologically impoverished, however environmentally aware lawn owners are reluctant to implement alternatives due to aesthetic concerns. Developing an alternative lawn format which is both biodiversity friendly and aesthetically pleasing is an imperative for urban greening. We suggest that such an alternative can be provided by replacing the grass lawn by a forb-based mix. To advance this, we tested the floral performance of three groups of clonal perennial forbs (native, non-native and mixed), each maintained using standard lawn management mowing regimes. Our findings show that both the frequency of mowing and the height at which mowing is applied influence floral performance and lawn aesthetics. Species origin was found to influence floral productivity, floral visibility and floral variety within grass-free lawns, with native species providing the greatest floral performance. The behaviour and management of grass lawns was not found to be a suitable analogue for the management of grass-free lawns and grass-free lawns are sufficiently different from grass lawns to require an entirely original management approach. We suggest that the grass-free lawn can provide an aesthetically and environmentally relevant replacement for the ubiquitous and ecologically-poor grass lawn.

Short-term changes caused by fire and mowing in Brazilian Campos grasslands with different long-term fire histories

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Above- and below-ground biomass and carbon dynamics in Brazilian Cerrado wet grasslands

Questions: Grasslands are usually neglected as potential carbon stocks, partially due to the lack of studies on biomass and carbon dynamics in tropical grasslands. What is the importance of Brazilian tropical wet grasslands as carbon sinks? Does fire frequency and season affect biomass and carbon allocation in Brazilian wet grasslands? Location: Wet grasslands, tropical savanna, Jalapão, Tocantins, northern Brazil. Methods: We determined biomass above- and below-ground, estimated carbon stocks in biennially burned plots (B2) and plots excluded from fire for 4 yr (B4). Moreover, we determined biomass in both rainy and dry seasons. Samples were 0.25 m × 0.25 m × 0.2 m (eight samples per treatment, applying a nested design, total of 48 samples). The biomass was classified in above-ground graminoids, forbs and dead matter, and below-ground roots and other below-ground organs. We used ANOVA to compare variables between treatments and seasons. Results: More than 40% of the total biomass and carbon stocks were located below-ground, mostly in roots. A high proportion of dead biomass (B4) was found in the above-ground material, probably due to low decomposition rates and consequent accumulation over the years. Although these grasslands do not experience water stress, we found significant evidence of resource re-allocation from below-ground organs to the above-ground biomass in the rainy season. Conclusions: We found more dead biomass in the rainy season, probably due to low decomposition rates, which can increase fire risk in these grasslands during the following dry season. These tropical wet grasslands stored high amounts of carbon (621 to 716 g C.m-2), mostly in the roots. Thus, policymakers should consider tropical grasslands as potential carbon stocks, since they are one of the most threatened and unprotected ecosystems in Brazil. © 2012 International Association for Vegetation Science.

Does disturbance affect bud bank size and belowground structures diversity in Brazilian subtropical grasslands?

Brazilian Campos grasslands are ecosystems under high frequency of disturbance by grazing and fires. Absence of such disturbances may lead to shrub encroachment and loss of plant diversity. Vegetation regeneration after disturbance in these grasslands occurs mostly by resprouting from belowground structures. We analyzed the importance of bud bank and belowground bud bearing organs in Campos grasslands. We hypothesize that the longer the intervals between disturbances are, the smaller the size of the bud bank is. Additionally, diversity and frequency of belowground organs should also decrease in areas without disturbance for many years. We sampled 20 soil cores from areas under different types of disturbance: grazed, exclusion from disturbance for two, six, 15 and 30 years. Belowground biomass was sorted for different growth forms and types of bud bearing organs. We found a decrease in bud bank size with longer disturbance intervals. Forbs showed the most drastic decrease in bud bank size in the absence of disturbance, which indicates that they are very sensitive to changes in disturbance regimes. Xylopodia (woody gemmiferous belowground organs with hypocotyl-root origin) were typical for areas under influence of recurrent fires. The diversity of belowground bud bearing structures decreased in the absence of disturbance. Longer intervals between disturbance events, resulting in decrease of bud bank size and heterogeneity of belowground organs may lead to the decline and even disappearance of species that relay on resprouting from the bud bank upon disturbance. (C) 2014 Elsevier GmbH. All rights reserved.

Does fire induce flowering in Brazilian subtropical grasslands?

QuestionsWe aimed to analyse the effect of fire on flowering in subtropical grasslands, by addressing the following questions: will fire history affect flowering? If yes, do fire feedbacks influence flowering or is it just the removal of above-ground biomass? Are there differences in burned and mowed plots?LocationSubtropical grasslands in Southern Brazil (30 degrees 03S, 51 degrees 07W).MethodsWe established plots in areas with different fire histories: 30d (30 plots: five replicates), 1yr (14 replicates), 3yr (30 plots: five replicates) since the last fire, in experimentally burned and mowed plots (14 replicates each). We counted the number of flowering species, as well as the number of flowering stalks.ResultsGraminoid species flowered in highest numbers 1yr after fire, whilst forbs had more species flowering just after fire, indicating different reproductive strategies in post-fire environments. Mowing was not as efficient as fire in stimulating flowering. Finally, the different functional groups showed different flowering responses to time since last fire and to the different types of management.ConclusionsOur results show fire stimulated flowering. Although mowing can be a good alternative for maintaining plant diversity, our study showed that this practice is not as efficient as fire in stimulating flowering. However, fire season should be noted as a limiting factor to the recovery of C-3 grasses in these subtropical grasslands, and annual burns may be harmful to C-4 grasses, since they delay their flowering to the next post-fire growing season.

Avaliação da produtividade em áreas de campo sujo com diferentes históricos de fogo

Fire is a common event in different ecosystems and can both be caused by humans or have natural sources.. In many of these ecosystems, natural fires are an important factor that determines the vegetation. The reduction of tree cover by fire for example, resulted in the evolution of several species-rich ecosystems, dominated by C4 grasses. However, the fire caused by human actions may have greater intensity and lead to negative responses of vegetation, since man changed the fire regime in many parts of the world, such as in the Cerrado. The passage of fire can benefit herbaceous and woody seedlings that cannot compete with the dominant grass layer. It removes the dead biomass and litter (major components of the fuel load), opening up spaces within the grass matrix that allow the establishment of other species. After some time without fire, an increase in shrub cover and decrease herbaceous layer can be observed. One of the major consequences of the absence of fire in savanna and grassland ecosystems is the accumulation of flammable dead biomass (mainly composed of graminoids), which will probably be the fuel load of the next burning thus, fires will be more intense and hotter. Moreover, very frequent burns lead to a reduction in the frequency and density of grasses. Therefore, this study aimed to assess the quantity and quality of biomass in areas with different fire history (fire exclusion for 2 and 7 years) in areas of campo sujo in central Cerrado. Plots (1x1m) were established in both areas and all aboveground biomass of each plot was cut at ground level and put in paper bags in the field. In the laboratory, the material was sorted into live and dead biomass. In addition, live biomass was separated into different functional groups (graminoids, forbs, Vellozia spp, palm and shrubs). The material was oven dried for two days at 80°C and subsequently weighed. In both areas, we found a dominance of graminoid and dead biomass. The area...

Avaliação de diferentes técnicas de manejo para o controle de gramíneas invasoras em unidade de conservação de cerrado

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Efeito da época da queima na dinâmica de campo sujo de Cerrado

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Efeito da época da queima na dinâmica de campo sujo de Cerrado

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

The importance of plant life form on spatial associations along a subtropical coastal dune gradient

Questions Does the spatial association between isolated adult trees and understorey plants change along a gradient of sand dunes? Does this association depend on the life form of the understorey plant? Location Coastal sand dunes, southeast Brazil. Methods We recorded the occurrence of understorey plant species in 100 paired 0.25 m2 plots under adult trees and in adjacent treeless sites along an environmental gradient from beach to inland. Occurrence probabilities were modelled as a function of the fixed variables of the presence of a neighbour, distance from the seashore and life form, and a random variable, the block (i.e. the pair of plots). Generalized linear mixed models (GLMM) were fitted in a backward step-wise procedure using Akaike's information criterion (AIC) for model selection. Results The occurrence of understorey plants was affected by the presence of an adult tree neighbour, but the effect varied with the life form of the understorey species. Positive spatial association was found between isolated adult neighbour and young trees, whereas a negative association was found for shrubs. Moreover, a neutral association was found for lianas, whereas for herbs the effect of the presence of an adult neighbour ranged from neutral to negative, depended on the subgroup considered. The strength of the negative association with forbs increased with distance from the seashore. However, for the other life forms, the associational pattern with adult trees did not change along the gradient. Conclusions For most of the understorey life forms there is no evidence that the spatial association between isolated adult trees and understorey plants changes with the distance from the seashore, as predicted by the stress gradient hypothesis, a common hypothesis in the literature about facilitation in plant communities. Furthermore, the positive spatial association between isolated adult trees and young trees identified along the entire gradient studied indicates a positive feedback that explains the transition from open vegetation to forest in subtropical coastal dune environments.

Recruitment limitation constrains local species richness and productivity in dry grassland

Species coexistence and local-scale species richness are limited by the availability of seeds and microsites for germination and establishment. We conducted a seed addition experiment in seminatural grassland at three sites in southern Switzerland and repeated the experiment in two successive years to evaluate various circumstances under which seed limitation and establishment success affect community functioning. A collection of 144000 seeds of 22 meadow species including grasses and forbs of local provenance was gathered, and seeds were individually sown in a density that resembled natural seed rain. The three communities were seed limited. Three years after sowing, single species varied in emergence (0–50%), survival (0–69%), and establishment rates (0–27%). One annual and 13 perennial species reached reproductive stage. Low establishment at one site and reduced growth at another site indicated stronger microsite limitation compared to the third site. Recruitment was influenced by differences in abiotic environmental conditions between sites (water availability, soil minerals) and by within-site differences in biotic interaction (competition). At the least water-limited site, sowing resulted in an increase in phytomass due to establishment of short-lived perennials in the second and third years after sowing. This increase persisted over the following two years due to establishment of longer-lived perennials. After sowing in a wetter year with higher phytomass, however, productivity did not increase, because higher intensity of competition in an early phase of establishment resulted in less vigorous plants later on. Due to the generally favorable weather conditions during this study, sowing year had a small effect on numbers of established individuals over all species. Recruitment limitation can thus constrain local-scale species richness and productivity, either by a lack of seeds or by reduced seedling growth, likely due to competition from the established vegetation.

The role of landuse and natural determinants for grassland vegetation composition in the Swiss Alps

The Alps provide a high habitat diversity for plant species, structured by broad- and fine-scale abiotic site conditions. In man-made grasslands, vegetation composition is additionally affected by the type of landuse. We recorded vegetation composition in 216 parcels of grassland in 12 municipalities representing an area of 170 x 70 km in the south-eastern part of the Swiss Alps. Each parcel was characterized by a combination of altitudinal level (valley, intermediate, alp). traditional landuse (mown. grazed), current management (mown, grazed, abandoned). and Fertilization (unfertilized, fertilized). For each parcel we also assessed the abiotic factors aspect, slope, pH value, and geographic coordinates, and for each municipality annual precipitation and its cultural tradition. We analysed vegetation composition using (i) variation partitioning in RDA. (ii) cover of graminoids. non-legume forbs, and legumes, and (iii) dominance and frequency of species. Species composition was determined by, in decreasing order of variation explained. landuse, broad-scale abiotic factors, fine-scale abiotic factors. and cultural tradition. Current socio-economically motivated landuse changes, such as grazing of unfertilized former meadows or their abandonment, strongly affect vegetation composition. In our study, the frequency of characteristic meadow species was significantly smaller in grazed and even smaller in abandoned parcels than in still mown ones, suggesting less severe consequences of grazing for vegetation composition than of abandonment. Therefore. low-intensity grazing and mowing every few years should be considered valuable conservation alternatives to abandonment. Furthermore. because each landuse type was characterized by different species. a high variety of landuse types should be promoted to preserve plant species diversity in Alpine grasslands. (C) 2007 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

Impacts of drought stress on water relations and carbon assimilation in grassland species at different altitudes

Grassland is an important ecosystem type which is not only used agriculturally, but also covers sites which cannot be used for other purposes, e.g. in very steep locations or above timberlines. Prolonged summer droughts in the mid-term future, as are predicted for Central Europe, are expected to have a major impact on such ecosystems. To address this topic, rainfall exclusion via shelters was performed on three grassland sites at different altitudes (393, 982 and 1978 m above sea level) in Switzerland. Diurnal drought treatment effects were studied at each study site on a completely sunny day towards the end of an 8–10 week shelter period. Ecophysiological parameters including gas exchange (An, gs and intrinsic WUE) and chlorophyll a fluorescence (Fv/Fm, ΦPSII and NPQ) were considered for several species. The lowland and the Alpine field site were more strongly affected by soil drought than the pre-Alpine site. At all sites, grasses showed different patterns of reductions in stomatal conductance under soil drought compared to legumes and forbs. In addition, grasses were significantly more affected by reductions in assimilation rates at all sites. Time courses of reductions in assimilation rates relative to controls differed between species at the Alpine site, as some species showed reduced assimilation rates at this site in the early morning. Thus, similar rainfall exclusion treatments can trigger different reactions in various species at different sites, which might not become obvious during mere midday measurements. Overall, results suggest strong impacts of prolonged summer drought on grassland net photosynthesis especially at the Alpine site and, within sites, for grasses

Plant regeneration directs changes in grassland composition after extreme drought: a 13-year study in southern Switzerland

1. The cover of plant species was recorded annually from 1988 to 2000 in nine spatially replicated plots in a species-rich, semi-natural meadow at Negrentino (southern Alps). This period showed large climatic variation and included the centennial maximum and minimum frequency of days with ≥ 10 mm of rain. 2. Changes in species composition were compared between three 4-year intervals characterized by increasingly dry weather (1988–91), a preceding extreme drought (1992–95), and increasingly wet weather (1997–2000). Redundancy analysis and anova with repeated spatial replicates were used to find trends in vegetation data across time. 3. Recruitment capacity, the potential for fast clonal growth and seasonal expansion rate were determined for abundant taxa and tested in general linear models (GLM) as predictors for rates of change in relative cover of species across the climatically defined 4-year intervals. 4. Relative cover of the major growth forms present, graminoids and forbs, changed more in the period following extreme drought than at other times. Recruitment capacity was the only predictor of species’ rates of change. 5. Following perturbation, re-colonization was the primary driver of vegetation dynamics. The dominant grasses, which lacked high recruitment from seed, therefore decreased in relative abundance. This effect persisted until the end of the study and may represent a lasting response to an extreme climatic event.