Aims Functional traits are supposed to play an important role in determining the colonization success of new species into established communities. Short-term experimental studies have documented higher resistance of more diverse grasslands against colonization by new species. However, little is known about which traits colonizers should have to successfully invade diverse plant communities in the longer term and how community history may modify the resistance of diverse communities against colonization.
Methods In a grassland biodiversity experiment (Jena Experiment) established with different species richness (SR; 1, 2, 4, 8 and 16) and functional group (FG) number and composition (1 to 4; legumes, grasses, small herbs, tall herbs), we studied colonization of naturally dispersed species in split-plots (i) with different duration of weeding (never weeded, weeded for 3 or 6 years and then un-weeded for 1 year) and (ii) with different duration of colonization (7 years, 4 years and 1 year after cessation of weeding).
Important findings Resistance against colonization by new species declined with increased duration of weeding (on average 13, 17 and 22 colonizer species in 1-, 4- and 7-year-old communities, respectively). Communities established at low diversity accumulated more colonizer species with a longer duration of weeding than more diverse communities. Duration of colonization had only small effects on the number of colonizer species. Colonizers with early successional traits, i.e. annual life cycle, reproduction by seeds, small seeds, long-lived seeds and an earlier start of a longer flowering period, were favoured in species-poor newly established experimental plant communities (short duration of weeding) and early after cessation of weeding (short duration of colonization). A change from early- to mid-successional traits, i.e. taller growth, perennial life cycle, vegetative reproduction, characterized colonization at increased plant diversity and in communities with legumes or without grasses. Legume absence/grass presence and increased duration of weeding led to a shift in colonizer strategies from rapid nutrient uptake and cycling (higher specific leaf area) to nutrient retention and symbiotic N 2 fixation. Our study shows that non-random trait spectra of naturally dispersed colonizers encompass trade-offs between different functions (reproduction, persistence, growth) reflected in a change from early- to mid-successional traits at increasing plant diversity, with a longer duration of weeding and a longer time of colonization.

Aims We analyse here the variations in species composition and richness and the geographic ranges of the tree species occurring in South American subtropical Atlantic and Pampean forests. Our goals were to assess (i) the floristic consistency of usual classifications based on vegetation physiognomy, climate and elevation; (ii) the leading role of temperature-related variables on the variations in species composition and richness; (iii) the predominance of species with tropical–subtropical ranges, possibly as a result of forest expansion over grasslands after the Last Glacial Maximum (LGM); (iv) the restriction of most subtropical endemics to stressful habitats as a possible result of past forest refuges during the LGM.
Methods The region was defined by the Tropic of Capricorn to the north, the Rio de la Plata to the south, the Atlantic shoreline to the east and the catchment areas of the upper Paraná and Uruguay Rivers to the west. Multivariate analyses, multiple regression modelling and variance partition analyses were performed on a database containing 63 994 occurrence records of 1555 tree species in 491 forest sites and 48 environmental variables. All species were also classified according to their known geographic range.
Important findings A main differentiation in species composition and richness was observed between the eastern windward coastlands (rain and cloud forests) and western leeward hinterlands (Araucaria and semi-deciduous forests). Pre-defined forest types on both sides were consistent with variations in tree species composition, which were significantly related to both environmental variables and spatial proximity, with extremes of low temperature playing a chief role. Tree species richness declined substantially towards the south and also from rain to seasonal forests and towards the highland summits and sandy shores. Species richness was significantly correlated with both minimum temperature and actual evapotranspiration. About 91% of the subtropical flora is shared with the much richer tropical flora, probably extracting species that can cope with frost outbreaks. The 145 subtropical endemics were not concentrated in harsher habitats.

Aims Litter decomposition is a critical pathway linking the above- and belowground processes. However, factors underlying the local spatial variations in forest litter decomposition are still not fully addressed. We investigated leaf litter decomposition across contrasting forest stands in central China, with objective to determine the spatial variations and controlling factors in forest floor leaf litter decomposition in relation to changes in forest stands in a temperate forest ecosystem.
Methods Leaf litter decomposition was studied by using litterbag method across several typical forest stand types in Baotianman Nature Reserve, central China, including pure stands of Quercus aliena var. acuteserrata, Q. glandulifera var. brevipetiolata and Q. variabilis, respectively, and mixed pine/oak stands dominated by Pinus armandii and Q. aliena var. acuteserrata, as well as stands of pure Q. aliena var. acuteserrata trees ranging in stand age from ~40 to>160 years. Measurements were made on litter mass remaining and changes in litter chemistry during decomposition over a 2-year period, along with data collections on selective biotic and environmental factors. A reciprocal transplant experiment involving Q. aliena var. acuteserrata and Q. variabilis was concurrently carried out to test the occurrence of 'home-field advantage (HFA)' in local forests when only considering contrasting oak tree species. Correlation analyses and path analyses were performed to identify the dominant drivers and their relative contributions to variations in leaf litter decomposition.
Important findings Significant variations were found in the rate of leaf litter decomposition among stands of different tree species but not among stand age classes. The values of decay constant, k, varied from 0.62 in Q. aliena var. acuteserrata stands to 0.56 in Q. variabilis stands. The reciprocal litter transplant experiment showed that the rate of leaf litter decomposition was on average 5% slower in home-fields than on reciprocal sites. Path analysis identified litter acid-unhydrolyzable residue (AUR) to N ratio, soil microbial biomass carbon (MBC), soil pH and soil organic carbon (SOC) as most prominent factors controlling the rate of leaf litter decomposition, collectively accounting for 57.8% of the variations; AUR/N had the greatest negative effect on k value, followed by weaker positive effects of SOC and MBC. Our findings suggest that tree species plays a primary role in affecting forest floor leaf litter decomposition by determining the litter quality, with site environment being a secondary factor contributing to the local variations in leaf litter decomposition in this temperate forest ecosystem.

Aims Phenotypic plasticity and local adaptation of populations at their distributional limits are crucial to understand species colonization and persistence in novel or marginal environments, as well as species divergence and niche width evolution. We assess the contribution of these processes to shape current elevational limits and determine elevational phenotypic divergence between two subspecies of Aquilegia vulgaris (subsp. vulgaris and nevadensis).
Methods We conducted sowing and transplant experiments considering four elevations throughout the current elevational range of A. vulgaris in southern Iberian Peninsula. Experiments were designed to explore, on the one hand, local adaptation through three components of performance (germination, survival and growth) and, on the other hand, the phenotypic differentiation and/or plasticity associated to local adaptation. Four populations per subspecies (three from the elevational core and one from the elevational boundary) were used as seed sources. Patterns of local adaptation and phenotypic differentiation are examined in the context of the 'centre-periphery' hypothesis.
Important findings Central populations of both subspecies performed better at their local elevations while marginal populations were maladapted, confirming the hypothesis and contributing to explain the current elevational segregation of these subspecies. Density of glandular pubescence and germination timing seem to be related to local adaptation, through phenotypic differentiation between subspecies or elevations. The widespread subsp. vulgaris showed signals of adaptive plasticity in the timing of germination while it was not the case in the endemic subsp. nevadensis .

Aims It has been well recognized that understory vegetation plays an important role in driving forest ecosystem processes and functioning. In subtropical plantation forests, understory removal and fertilization have been widely applied; however, our understanding on how understory removal affects soil respiration and how the process is regulated by fertilization is limited. Here, we conducted an understory removal experiment combined with fertilization to evaluate the effects of the two forest management practices and their interactions on soil respiration in subtropical forest in southern China.
Methods The study was conducted in a split-plot design with fertilization as the whole-plot factor, understory removal as the subplot factor and block as the random factor in subtropical Eucalyptus plantations. In total, there were four treatments: control with unfertilized and intact understory (CK), understory removal but without fertilization (UR), with fertilization but without understory removal (FT) and with fertilization + understory removal (FT + UR). Eucalyptus above- and belowground biomass increment, fine root biomass, soil temperature, soil moisture and soil respiration were measured in the present study. Understory respiration (R U) was quantified in different ways: R u = R CK ? R UR or R u = R FT ? R (FT + UR); fertilization increased soil respiration (R FI) was also quantified in different ways: R FI = R FT ? R CK or R FI = R (FT + UR) ? R UR .
Important findings Over a 2-year experiment, our data indicate that understory removal significantly decreased soil respiration, while fertilization increased soil respiration. Understory removal decreased soil respiration by 28.8% under fertilization, but only 15.2% without fertilization. Fertilization significantly increased soil respiration by 23.6% with the presence of understory vegetation, and only increased by 3.7% when understory was removed, indicating that fertilization increased soil respiration mainly by increasing the contribution of the understory. Our study advances our understanding of the interactive effects of understory management and fertilization on soil respiration in subtropical plantations.

Aims The plasticity of ecosystem responses could buffer and postpone the effects of climates on ecosystem carbon fluxes, but this lagged effect is often ignored. In this study, we used carbon flux data collected from three typical grassland ecosystems in China, including a temperate semiarid steppe in Inner Mongolia (Neimeng site, NM), an alpine shrub-meadow in Qinghai (Haibei site, HB) and an alpine meadow steppe in Tibet (Dangxiong site, DX), to examine the time lagged effects of environmental factors on CO₂ exchange.
Methods Eddy covariance data were collected from three typical Chinese grasslands. In linking carbon fluxes with climatic factors, we used their averages or cumulative values within each 12-month period and we called them 'yearly' statistics in this study. To investigate the lagged effects of the climatic factors on the carbon fluxes, the climatic 'yearly' statistics were kept still and the 'yearly' statistics of the carbon fluxes were shifted backward 1 month at a time.
Important findings Soil moisture and precipitation was the main factor driving the annual variations of carbon fluxes at the alpine HB and DX, respectively, while the NM site was under a synthetic impact of each climatic factor. The time lagged effect analysis showed that temperature had several months, even half a year lag effects on CO₂ exchange at the three studied sites, while moisture's effects were mostly exhibited as an immediate manner, except at NM. In general, the lagged climatic effects were relatively weak for the alpine ecosystem. Our results implied that it might be months or even 1 year before the variations of ecosystem carbon fluxes are adjusted to the current climate, so such lag effects could be resistant to more frequent climate extremes and should be a critical component to be considered in evaluating ecosystem stability. An improved knowledge on the lag effects could advance our understanding on the driving mechanisms of climate change effects on ecosystem carbon fluxes.

Aims With a close association with plant water availability, foliar δ 13 C had been investigated extensively in alpine regions; however, foliar δ 15 N has rarely been concurrently used as an indicator of plant nitrogen availability. Due to the positive correlations between leaf nitrogen content and foliar δ 13 C and δ 15 N found in previous studies, we expected that they should show consistent patterns along an altitudinal gradient.
Methods To test our hypothesis, we measured foliar δ 13 C and δ 15 N in conjunction with multiple key leaf functional traits of Quercus aquifolioides, a dominant species of alpine forest on the eastern slopes of the Sygera Mountains, southeastern Tibetan Plateau from 2500 to 3800 m.
Important findings (i) Contrary to our hypothesis, foliar δ 13 C exhibited a significant positive linear relationship with altitude; in contrast, foliar δ 15 N initially increased and subsequently decreased with altitude, the change in trend occurring around 3300 m. (ii) Our analyses indicated that leaf internal resistance and stomatal conductance, rather than photosynthetic capacity indicated by leaf N concentration, apparently explained the altitudinal variation in foliar δ 13 C, while differences in foliar δ 15 N were likely the result of soil N availability. (iii) Principal component analysis revealed a clear association between δ 13 C and a tradeoff between water loss and carbon gain, indicated by traits related to gas exchange such as leaf thickness, density, stomatal properties. In contrast, the second axis was associated with δ 15 N and nitrogen acquisition strategy in Q. aquifolioides across its altitudinal distribution, represented by traits related to nitrogen concentration and stomata per gram of leaf nitrogen.

Aims Ecological systems, especially soils, have been recently recognized as an important source of atmospheric nitric oxide (NO). However, the study on the contribution of plants to atmospheric NO budget is significantly lagged. The specific objectives of this study are to reveal the phylogenetic variation in NO emission potential existing in various plant species and find out the possible leaf traits affecting NO emission potential.
Methods We measured NO emission potential, leaf N and C content, C:N ratio, specific leaf area, net photosynthetic rate (P n) and estimated photosynthetic N use efficiency (PNUE) of 88 plant species. Further investigation of the relationships between NO emission potential and leaf traits were performed by simple linear regression analysis and pair-wise correlation coefficients analysis.
Important findings Major results are as follows: (1) NO emission from plant species exhibited large variations, ranging from 0 to 41.7 nmol m ?2 h^-1, and the species frequency distributions of NO emission potential could be fitted to a log-normal curve. (2) Among 88 species, NO emission potential was the highest in Podocarpus macrophyllus, but lowest in Zanthoxylum nitidum and Vernicia montana. (3) NO emission potential has strong correlation to leaf N content, P n and PNUE. The variations in NO emission potential among diverse plant species may be closely related to leaf N level and net photosynthetic ability.

Aim European and North American studies have suggested that nitrogen (N) depositions reduce plant diversity and increase primary productivity due to changes in plant traits. To predict the vegetation response to future global change, experimental validations from other regions are widely needed. We assessed the effects of N treatment by urea fertilization on the diversity and biomass of the herbaceous plant traits (HPTs) in a dry tropical environment of India.
Methods Diversity and biomass of different HPTs were determined on the basis of data collected in year 2010, from 135, 1 m × 1 m plots distributed over 15 locations. The plots were treated with urea fertilizer in different doses (Control, 60kgNha^-1 yr^-1 and 120kg N ha^-1 yr^-1) since 1st January 2007. The plots were ordinated and data were subjected to appropriate statistical analyses.
Important findings Correspondence analysis (CA) suggested uniqueness of species composition due to N amendment. Species number and biomass of the trait categories varied due to N fertilization and traits. All studied trait categories (except N-fixers) yielded maximum mean species number at moderate level of N fertilization. Different levels of N fertilization exhibited different species diversity–primary productivity (D-P) relationships. Further, study showed reduction in plant diversity due to increase in biomass at high rates of N addition.Conclusions Tall, erect, non N-fixers, annuals, grasses HPTs were favoured by N enrichment. N dose above 60kg enhanced the biomass of fast growing, erect, annuals, non N-fixers, nitrophilic HPTs. The changes in traits with N addition, especially the increase in annuals and grasses and decrease in typically N-rich N-fixers, have implications for sustainable cattle production.