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  • 1
    Publication Date: 2015-09-17
    Description: Aims This study explores the patterns of niche differentiation in a group of seven closely related columbines (genus Aquilegia , Ranunculaceae) from the Iberian Peninsula. Populations of these columbines are subject to complex patterns of divergent selection across environments, which partly explain the taxonomic structure of the group. This suggests the hypothesis that niche divergence must have occurred along the process of diversification of the group. Methods We used MaxEnt to build environmental niche models of seven subspecies belonging to the three species of Aquilegia present in the Iberian Peninsula. From these models, we compared the environmental niches through two different approaches: ENMtools and multivariate methods. Important Findings MaxEnt distributions conformed closely to the actual distribution of the study taxa. ENMtools methods failed to uncover any clear patterns of niche differentiation or conservatism in Iberian columbines. Multivariate analyses indicate the existence of differentiation along altitudinal gradients and along a gradient of climatic conditions determined by the summer precipitation and temperatures. However, climatic conditions related to winter temperature and precipitation, as well as soil properties, were equally likely to show conservatism or divergence. The complex patterns of niche evolution we found suggest that Iberian Columbines have not been significantly constrained by forces of niche conservatism, so they could respond adaptively to the fast and profound climate changes in the Iberian Peninsula through the glacial cycles of the Pleistocene.
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  • 2
    Publication Date: 2015-09-17
    Description: Aims The pattern and driving factors of forest fires are of interest for fire occurrence prediction and forest fire management. The aims of the study were: (i) to describe the history of human-caused fires by season and size of burned area over time; (ii) to identify the spatial patterns of human-caused fires and test for the existence of ‘hotspots’ to determine their exact locations in the Daxing’an Mountains; (iii) to determine the driving factors that determine the spatial distribution and the possibility of human-caused fire occurrence. Methods In this study, K -function and Kernel density estimation were used to analyze the spatial pattern of human-caused fires. The analysis was conducted in S-plus and ArcGIS environments, respectively. The analysis of driving factors was performed in SPSS 19.0 based on a logistic regression model. The variables used to identify factors that influence fire occurrence included vegetation types, meteorological conditions, socioeconomic factors, topography and infrastructure factors, which were extracted and collected through the spatial analysis mode of ArcGIS and from official statistics, respectively. Important Findings The annual number of human-caused fires and the area burnt have declined since 1987 due to the implementation of a forest fire protection act. There were significant spatial heterogeneity and seasonal variations in the distribution of human-caused fires in the Daxing’an Mountains. The heterogeneity was caused by elevation, distance to the nearest railway, forest type and temperature. A logistic regression model was developed to predict the likelihood of human-caused fire occurrence in the Daxing’an Mountains; its global accuracy attained 64.8%. The model was thus comparable to other relevant studies.
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  • 3
    Publication Date: 2015-11-24
    Description: Aims Oxidative stress is one of the most important mechanisms in a plant’s reaction to the effects of different stressors; however, its role in plants’ resistance is still poorly understood. The objective of this study is to evaluate an influence of oxidative stress induced by stress factors of different origin—ozone, ultraviolet (UV)-B radiation, drought, cadmium (Cd) and copper (Cu), to growth of spring barley and to check the hypothesis, that intensification of oxidative stress is the main factor of growth depression induced by strong treatments of different stressors; meanwhile, mitigation of oxidative stress determines eustress-induced growth stimulation. Methods A pot experiment was carried out in phytotron chambers with a controlled environment. Spring barley ( Hordeum vulgare L.) plants were exposed to different doses of investigated environmental stress factors (O 3 , UV-B radiation, drought, Cd and Cu), and their effects on shoots growth, accumulation of superoxide (O 2 .– ), intensification of lipid peroxidation and antioxidative protection (superoxide dismutase, glutathione reductase and catalase activities and concentration of carotenoids) were measured. Analysis of variance (ANOVA) with classical eta-squared ( 2 ) values was used to evaluate and to compare the contribution of non-specific oxidative stress and stressor-specific mechanisms on plants growth. Important Findings Low doses of most stressors stimulated antioxidative protection and growth of barley shoots, reduced the concentration of O 2 .– and/or intensity of lipid peroxidation. Whereas an impairment of growth and intensification of oxidative stress as well as a reduction in concentration of carotenoids and further increase in activity of antioxidative enzymes were noticed when the intensity of the stressors was increased. In the cases of ozone and UV-B stress, the effects of oxidative stress on plant growth was mitigated by strong antioxidative protection—highly increased catalase (CAT) and superoxide dismutase (SOD) activities, respectively. In the cases of drought and Cu, relatively strong oxidative stress was the major cause of plant growth depression. Additionally, mitigation of oxidative stress due to increased SOD activity was likely to be one of the main causes of growth stimulation induced by low doses of UV-B, Cd and Cu stress. Possible reasons for O 3 -induced growth stimulation were increased CAT activity and concentration of carotenoids. Generalizing the effects of different stressors, the contribution of non-specific oxidative stress on plant growth was stronger compared with stressor-specific action mechanisms: oxidative stress determined 42% of the changes in plants’ dry biomass, whereas the contribution of stressor-specific mechanisms accounted for 35% of variability in barley growth.
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  • 4
    Publication Date: 2015-11-24
    Description: Aims The biochemical defense of lichens against herbivores and its relationship to lichen frequency are poorly understood. Therefore, we tested whether chemical compounds in lichens act as feeding defense or rather as stimulus for snail herbivory among lichens and whether experimental feeding by snails is related to lichen frequency in the field. Methods In a no-choice feeding experiment, we fed 24 lichen species to snails of two taxa from the Clausilidae and Enidae families and compared untreated lichens and lichens with compounds removed by acetone rinsing. Then, we related experimental lichen consumption with the frequency of lichen species among 158 forest plots in the field (Schwäbische Alb, Germany), where we had also sampled snail and lichen species. Important findings In five lichen species, snails preferred treated samples over untreated controls, indicating chemical feeding defense, and vice versa in two species, indicating chemical feeding stimulus. Interestingly, compared with less frequent lichen species, snails consumed more of untreated and less of treated samples of more frequent lichen species. Removing one outlier species resulted in the loss of a significant positive relationship when untreated samples were analyzed separately. However, the interaction between treatment and lichen frequency remained significant when excluding single species or including snail genus instead of taxa, indicating that our results were robust and that lumping the species to two taxa was justified. Our results imply lichen-feeding snails to prefer frequent lichens and avoid less frequent ones because of secondary compound recognition. This supports the idea that consumers adapt to the most abundant food source.
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  • 5
    Publication Date: 2015-11-24
    Description: Aims Hydrogeomorphic processes operating at watershed, process zone and site scales influence the distribution of riparian vegetation. However, most studies examining the relationships between hydrogeomorphic processes and riparian vegetation are conducted at site scales. We quantified the relative importance of watershed, process zone and site geomorphic characteristics for predicting riparian plant community types and plant species abundances in four small mountain watersheds in central Nevada, USA. Methods We mapped riparian vegetation types and identified process zones (based on dominant geomorphic process and valley fill material) within the watersheds. We sampled sites in each combination of vegetation type and process zone ( n = 184 sites) and collected data on watershed scale factors, valley and stream geomorphic characteristics and on plant cover of each geomorphic surface. Plant community types were defined by cluster and indicator species analyses of plant cover data, and related to geomorphic variables using ordination analysis (nonmetric multidimensional scaling). Linear mixed effects models were used to predict abundances of indicator species. Important Findings Variables describing position in the watershed (elevation, contributing area) that are related to gradients of temperature, moisture and stream discharge were of primary importance in predicting plant community types. Variables describing local geomorphic setting (valley width, stream gradient, channel sediments, geomorphic surface height) were of secondary importance, but accurately described the geomorphic setting of indicator species. The process zone classification did not include position in the watershed or channel characteristics and only predicted plant community types with unique geomorphic settings. In small mountain watersheds, predicting riparian vegetation distribution requires explicit consideration of scale and geomorphic context within and among watersheds in addition to site variables.
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  • 6
    Publication Date: 2015-11-24
    Description: Aims Plant–soil interaction (PSI) has been implicated as a causative mechanism promoting plant invasions, and some mechanisms underlying PSI effects remain unclear. Here, we attempted to address how altered soil microbes and nutrients influence PSI effects. Methods Soil was cultured by an invasive forb Solidago canadensis for two years. We conducted an experiment, in which S. canadensis and Chinese natives were grown either alone or together in control and cultured soils, and determined the growth of S. canadensis and five natives and the competitive ability of S. canadensis . We analyzed the microbial community composition and nutrients of two types of soils. Important Findings Compared to the control soil, the soil cultured by S. canadensis decreased the subsequent growth of S. canadensis and five Chinese natives, as well as the competitive ability of S. canadensis against Chinese natives. Soil microbial community composition was significantly altered due to soil culturing. Total fatty acids, bacteria, Gram-negative bacteria and Gram-positive bacteria had no responses to soil culturing; fungi, aerobic bacteria and fungi/bacteria ratio significantly decreased with soil culturing; anaerobes and Gram-negative/positive bacteria ratio greatly increased with soil culturing. Soil nitrogen (N) dramatically decreased with soil culturing, whereas soil phosphorus (P) was unchanged. These results suggest that negative PSI effects may be linked to decreases in soil fungi, aerobic bacteria and soil N and increases in soil anaerobic bacteria and the ratio of Gram-negative/positive bacteria. Our findings provide an initial indication that S. canadensis– soil interaction alone could exhibit limited contributions to its success in the early stage of invasion.
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  • 7
    Publication Date: 2015-11-24
    Description: Aims Nitrogen (N) fertilization and lime addition may affect soil microbial and nematode communities and ecosystem functions through changing environmental conditions, such as soil pH and soil organic carbon. The objectives of this experiment were to examine the impact of N input and liming on soil microbial and nematode communities and to identify the key environmental determinant of community composition in a century-old fertilization and crop rotation experiment. Methods The field experiment consisting of a 3-year crop rotation regime was established in 1911 in southeastern USA. Four treatments, (i) no-input control, (ii) NPK with winter legume, (iii) PK with legume and lime and (iv) NPK with legume and lime, were included in this study. Soil samples collected at the 0–5cm depth were used to determine the bacterial growth rate by the 3 H-thymidine incorporation technique. Incorporation of 13 C into neutral lipids, glycolipids and phospholipid fatty acids (PLFAs) was measured after incubation of soil with 13 C-labeled acetate for 24h. Free-living nematodes in fresh soil were extracted using a density sucrose centrifugal flotation method and identified to trophic group level. Important Findings Liming resulted in a 10-fold increase in bacterial growth rates compared with the no-input control, whereas N fertilization had no significant effect. Multivariate analysis of PLFA profiles showed that soil microbial community composition was different among the four treatments; the difference was primarily driven by soil pH. PLFAs indicative of Gram-negative bacteria covaried with soil pH, but not those of fungi and actinobacteria. Liming enhanced 13 C incorporation into neutral lipids, glycolipids and phospholipids by 2–15 times. In addition, 13 C incorporation into 16:0, 16:19, 18:19, 18:17 and 18:26 were greater than other PLFAs, suggesting that Gram-negative bacteria and fungi were more active and sensitive to simple C input. Bacterivorous nematodes were the dominant trophic group in the soil, but no significant differences in nematode communities were found among the treatments. Our results suggest that soil pH had a greater impact than N fertilization on soil microbial community composition and activity in a crop rotation system including legumes.
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  • 8
    Publication Date: 2015-11-24
    Description: Aims In cool-season grasses, systemic and vertically transmitted Epichloë infections often provide a suite of benefits including increased growth, reproduction and competitive abilities. However, these effects of Epichloë endophytes on their hosts often depend upon host and endophyte genotype and environmental factors. Methods Achnatherum robustum (sleepygrass) harbors at least two Epichloë species within natural populations in the Southwest USA. We tested the effects of endophyte infection and species, host population and plant genotype (by experimentally removing the endophyte), and soil moisture (a key limiting factor) on growth and drought stress response of infected A. robustum plants from two populations (Weed and Cloudcroft) in the Sacremento Mountains of New Mexico, USA). Important Findings Although the two populations harbor distinct Epichloë species each with very different chemoprofiles, neither endophyte status (infected vs. uninfected) nor endophyte species affected most growth parameters at 8 or 25 weeks of the experiment, except for leaf length. In high water treatment, infected plants from the Weed population had longer leaf length compared with uninfected plants. In contrast, the population of origin affected all growth parameters, including plant height, leaf number, length and width, tiller number and shoot and root biomass, as well as wilting time. Grasses from the Cloudcroft population generally showed greater growth than grasses from the Weed population. Endophyte infection did affect wilting time, with infection in the Weed population generally reducing time to wilting under low and high water, whereas infection in the Cloudcroft population reduced time to wilting only under high water conditions. Our results suggest that plant population and their associated plant genotypes may play a much larger role in endophyte–host grass interactions in varying environments than previously thought. Asexual Epichloë species may be compatible with only specific host genotypes within populations such that the phenotypic effects due to population may be greater than phenotypic changes influenced by variation in the endophyte.
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  • 9
    Publication Date: 2016-01-29
    Description: Aims Theory suggests that species perform best at intermediate densities, where density-dependent facilitation and antagonism are balanced, but empirical evidence is scarce, particularly in plants. In a self-incompatible perennial herb ( Saussurea nigrescens ), whose recruitment heavily relies on seed output, we test whether both intraspecific facilitation and antagonism significantly affect seed production, resulting in highest seed yield at an intermediate capitulum density. Methods Plots with different S. nigrescens densities were sampled in an Eastern Tibetan meadow during the growing season of 2012 to investigate the relationships between capitulum density and pollinator visitation rate, seed set ratio, parasite ratio, seed damage ratio, and capitulum size. Both simple linear and quadratic models were employed to determine the shape of relationships. Important Findings In line with general theory, hump-shaped relationships of capitulum density versus seed set ratio and number of florets per capitulum indicate intraspecific facilitation in sparse populations, which can be attributed to positive density-dependent pollinator visitation and the amelioration of detrimental physical factors. However, the proportion of seeds damaged by pre-dispersal predators increased monotonically with capitulum density, which may have—in combination with increased intraspecific competition for light and soil nutrients—resulted in density-dependent antagonism. Both positive and negative density-dependent agents acted simultaneously throughout the density range investigated and led to the highest seed yield at intermediate density levels in the Tibetan lotus. More efforts concurrently exploring the two effects are needed to facilitate understanding species abundance and community structure.
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  • 10
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    Oxford University Press
    Publication Date: 2016-07-20
    Description: Aims Vegetation sampling employing observers is prone to both inter-observer and intra-observer error. Three types of errors are common: (i) overlooking error (i.e. not observing species actually present), (ii) misidentification error (i.e. not correctly identifying species) and (iii) estimation error (i.e. not accurately estimating abundance). I conducted a literature review of 59 articles that provided quantitative estimates or statistical inferences regarding observer error in vegetation studies. Important Findings Almost all studies (92%) that tested for a statistically significant effect of observer error found at least one significant comparison. In surveys of species composition, mean pseudoturnover (the percentage of species overlooked by one observer but not another) was 10–30%. Species misidentification rates were on the order of 5–10%. The mean coefficient of variation (CV) among observers in surveys of vegetation cover was often several hundred % for species with low cover, although CVs of 25–50% were more representative of species with mean covers of 〉50%. A variety of metrics and indices (including commonly used diversity indices) and multivariate data analysis techniques (including ordinations and classifications) were found to be sensitive to observer error. Sources of error commonly include both characteristics of the vegetation (e.g. small size of populations, rarity, morphology, phenology) and attributes of the observers (e.g. mental fatigue, personal biases, differences in experience, physical stress). The use of multiple observers, additional training including active feedback approaches, and continual evaluation and calibration among observers are recommended as strategies to reduce observer error in vegetation surveys.
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  • 11
    Publication Date: 2016-07-20
    Description: Aims Understanding relationships between vegetation and environments is of importance for ecosystem restoration and management. However, information on how environments influence the floristic patterns of shrublands is lack, especially in the subtropical China. In this study, we explored how environments regulate species composition of shrublands at landscape scale in mid-subtropical China. Methods We investigated species composition and measured the climate and soil environments for 207 shrubland plots in mid-subtropical China (24°39'–30°08'N, 108°47'–114°15'E). We applied a hierarchical cluster analysis and indicator species analysis based on the Bray–Curtis similarity index to identify the main shrubland types and employed principal coordinate analysis (PCoA) to explore the relationship between floristic composition and environment. Important Findings We identified four shrubland types occurring in different environmental conditions. Montane shrubland, dominated by species suitable for cool climates (e.g. Rhododendron simsii ), were distributed in steep areas at comparatively high altitudes; foothill shrubland, dominated by mesophilous species (e.g. Loropetalum chinense ), were distributed in low mountains and hills; pioneer shrubland, dominated by fast grow and short-life cycles species (e.g. Rhus chinensis ), were distributed at low altitudes with dense population; and finally, limestone shrubland, dominated by calcicole plants (e.g. Coriaria nepalensis ), were distributed in the extensive karst areas. Communities occurring in high pH soils were completely separated from those in low pH soils according to the hierarchical cluster analysis. PCoA ordination associated the four types with distinct edaphic and climatic gradients. Soil pH explained 63.3% of variation in PCoA, followed by soil depth and soil bulk density.
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  • 12
    Publication Date: 2016-07-20
    Description: Aims The loss of canopy trees associated with forest decline can greatly influence the species composition and structure of a forest and have major impacts on the ecosystem. We studied the changes in forest composition and structure 1 and 5 years following nearly total canopy mortality on several hundreds of hectares of xeric oak forests in south-central United States. Because the forests were within an ecotonal vegetation type composed of a mosaic of forest, savanna and grassland, we sought to learn whether forest decline areas would recover to forest or change to more open savanna and grassland conditions in the landscape pattern of vegetation. Because low intensity fire shaped the vegetation type, we sought to learn whether fire would keep the decline areas open. Methods The study was conducted in a xeric oak forest in east-central Oklahoma, USA. Randomly located vegetation and regeneration surveys were conducted in decline and non-decline stands 1 and 5 years following nearly total canopy mortality. Diameter at breast height (DBH), regeneration and sprout origin were recorded for all woody species. Important Findings The major canopy species post oak ( Quercus stellata Wangenh.), blackjack oak ( Quercus marilandica Muenchh.) and black hickory ( Carya texana Buckl.) suffered 85–92% mortality; however, minor canopy components experienced limited mortality. Mortality affected all size classes of canopy trees except those below 5cm breast height diameter. There was abundant regeneration of all species and fire seemed to maintain a high level of sprouting. Decline appeared to decrease the relative importance of stump sprouting and increase other types including root sprouts. Decline areas had abundant true seedlings, with stem origin from a root with the same diameter as the stem, which is very unusual for xeric oak forests. Regeneration height in decline areas was twice that of non-decline forests. Our findings suggest that forest decline may lead to: (i) reduced oak dominance and species change in the canopy, (ii) change in reproduction type to increase success of true seedlings and maintain genetic diversity of oaks.
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  • 13
    Publication Date: 2016-07-20
    Description: Aims To assess the role of moisture in phenological timing in the mediterranean coastal flora of Baja California, and specifically to assess the role of coastal fog and ocean-derived moisture in plant phenology. Moisture seems to be the primary driver of flowering times and durations at the arid end of the mediterranean-climate region, where rainfall is often sporadic (temperature and day length can be expected to play a much lesser role as they are not growth limiting). We aimed to understand: What factors drive climatic variation between sites? Are there general flowering patterns allowing us to identify phenological categories? Do flowering patterns vary in relation to site-specific weather? and most importantly, does maritime influence on weather affect flowering dynamics in coastal mediterranean ecosystems? Methods The southernmost extent of the California Floristic Province (in Baja California, Mexico) is a biological diversity hotspot of high endemism and conservation value, with two steep moisture gradients: rainfall (N–S) and coastal fogs (W–E), providing an ideal study system. We installed five weather stations across the moisture gradients, recording data hourly. We monitored flowering phenology in the square kilometer surrounding each weather station from 2010 to 2013. About 86 plant taxa were monitored across the five sites, every 6–8 weeks. Averaged climatic data is presented with general trends in flowering, and specific flowering syndromes were observed. Data for flowering intensity across the sites was analyzed using a principal components analysis. Important Findings Data analysis demonstrates a general seasonal pattern in flowering times, but distinct differences in local weather and phenology between the five study sites. Three flowering syndromes are revealed in the flora: (i) water responders or spring bloomers, (ii) day-length responders or fall-blooming taxa and (iii) aseasonal bloomers with no seasonal affinity. The two moisture gradients are the strongest drivers of flowering times. Inland sites showed higher phenological variation than coastal sites where seasonality is dampened by ocean-derived moisture, which extends and buffers perennial plant phenology and is a probable driver of local endemism. Phenological controls vary globally with climate and geography; moisture is the primary driver of phenology in mediterranean climates and fog is an important climatic variable in coastal Mexico.
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  • 14
    Publication Date: 2016-07-20
    Description: Aims The aim of the study was to discover what set of variables best explains the transition from warm to mesic forest vegetation. Based on various variables grouped into sets (geomorphological, ecological, structural, soil characteristics and chorological), six models were built and tested by generalized additive mixed models (GAMMs). We assumed that each set of variables has different explanatory power. Our aim was to compare the six different models (sets of variables), to test which model best explains the species turnover in forest communities along the transition between warm and mesic temperate forests and to try to find reasons for the different explanatory power of the models. Methods The research took place in the southern part of the Balkan Peninsula. Field sampling was done according to standard methods. The gradient from warm to mesic forests was defined as the turnover of species and evaluated by projection of samples on the first unconstrained DCA axis. Geomorphological, ecological, structural and soil characteristics, together with chorological sets of variables, were regressed on the turnover of species composition. Based on the five sets of variables, six models were constructed and tested by generalized additive mixed models. Important Findings Ecological conditions best explain the change of forest communities along the gradient; evolution and the development of vegetation reflected in chorotypes are also of high importance; geomorphology and structure seem not to change so dramatically and soil shows the least significant differences of all. Ecological variables are the most important set of variables in the transition between warm and mesic temperate forests but eco-evolutionary dynamics after the Pleistocene should also be taken into consideration.
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  • 15
    Publication Date: 2016-07-20
    Description: Aims Increasing evidence worldwide indicates that high mountain regions are not immune to invasion by alien plants. Here, we aimed to address whether three major woody invaders of low-mountain areas are constrained to lower altitudes due to climatic restrictions or just by low propagule pressure. We hypothesize that the increasing climatic harshness towards higher altitudes restricts seedling establishment and survival of these woody invaders and thus their potential for altitudinal expansion. Methods The study was carried out in the Sierras Grandes Mountains, Córdoba, central Argentina (32º50'S, 64º90'W). We addressed the hypothesis with an experimental approach, dissociating the effect of the environmental gradient from that of propagule pressure, by discarding the influence of seed sources. We translocated seeds of Gleditsia triacanthos L. , Ligustrum lucidum W.T. Aiton. and Pyracantha angustifolia (Franch.) C.K.Schneid. along the altitudinal gradient (from 900 to 2700 m asl). Seven sites were selected along the altitudinal gradient, spaced every 200–400 m. Three plots (4 x 4 m) were selected within each altitudinal site and excluded from livestock; 100 seeds of each species were sown within each plot (6300 seeds in total). Seedling emergence and survival was monitored during two growing seasons and soil temperature and moisture was recorded as well. The field experiment was complemented with lab assays. Important Findings Climate along this altitudinal gradient might be least restrictive at mid elevations, as a result of the intersection between opposite soil temperature and moisture gradients. Sown seeds germinated along the whole altitudinal gradient and seedlings successfully established and over-wintered well above their current altitudinal distribution (up to 2200 m for Ligustrum and 2400 m for Gleditsia and Pyracantha ). Additional lab assays confirmed field patterns and gave some insights into contrasting regeneration strategies between these invaders that might help to overcome stochastic environmental constraints in the germination stage. Overall, seedlings of three major woody invaders of low-mountain woodlands can establish and survive at higher elevations than their current distribution. In contrast to natives, they seem to be broad climate tolerators, rather than specialized stress tolerators, capable of germinating and growing across a wide elevational range. While long-term experiments might be necessary to fully address the potentials for altitudinal expansion, out results on early lifecycle stage suggest that the invaders studied here would have mostly a dispersal barrier rather than a climate barrier to establish in the upper belt of the Sierras Grandes.
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  • 16
    Publication Date: 2016-07-20
    Description: Aims The diversity–productivity relationship is one of the most critical questions in ecology and can be altered by environmental factors. Hydrological fluctuation affects growth of wetland plants, and such effects vary with plant species. Therefore, we hypothesized that hydrological fluctuation changes effects of species richness on productivity of wetland plant communities. Methods We constructed wetland plant communities consisting of three or six wetland plant species and subjected them to hydrological fluctuation (i.e. gradually changing water level) of two frequencies and two ranges, with unchanged water level as the control. We measured height, root and shoot dry mass of each plant at harvest. Important Findings Hydrological fluctuation significantly decreased biomass of wetland plant communities, which was due to impacts of fluctuation range, but not those of fluctuation frequency. Community biomass was significantly higher when species richness was higher, and such an effect did not depend on hydrological fluctuation. Therefore, hydrological fluctuation can decrease the productivity of wetland plant communities but may not alter the diversity–productivity relationship.
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  • 17
    Publication Date: 2016-07-20
    Description: Aims We aimed at determining differences in the leaf spectral signatures of plant species groups growing in habitats along the hydrological gradient of an intermittent wetland and to define leaf traits that explain their variability. We want to contribute to the understanding of the causes for plant spectrum variability at leaf and community levels. Methods We measured leaf reflectance spectra (300–887nm) of representative plant species from different habitats and analyzed spectral differences among species groups. To explain leaf spectra variability within a group, we performed detailed analyses of leaf morphological and biochemical traits in selected species. Important Findings The reflectance spectra of the different species groups differed most in the green, yellow and red spectral ranges. The reflectance spectra of submerged leaves of hydrophytes with simple structures were explained by their biochemical traits (carotenoids), while for more complex aerial leaves, morphological traits were more important. In submerged and natant leaves of amphiphytes, total mesophyll and spongy tissue thickness were the most important traits, and these explained 44% and 47%, respectively, of the spectrum variability of each plant group. In general, the redundancy analysis biplots show that samples of different plant species colonizing the same habitat form separate clusters and are related to the explanatory variables in different ways. The redundancy analysis biplots of helophytes and wet meadow species show clustering of graminoids and dicots into two distinct groups. Leaf encrustation (prickle hair properties and epidermis thickness) is important for graminoids, while leaf thickness and specific leaf area have more important roles in dicots. Our results show that knowledge of the species composition and leaf traits is necessary to interpret the reflectance spectra of such plant communities.
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  • 18
    Publication Date: 2016-07-20
    Description: Aims The physiological responses during dormancy removal and multi-phasic germination were investigated in seeds of Paeonia corsica (Paeoniaceae). Methods Seeds of P. corsica were incubated in the light at a range of temperatures (10–25 and 25/10°C), without any pre-treatment, after W (3 months at 25°C), C (3 months at 5°C) and W + C (3 months at 25°C followed by 3 months at 5°C) stratification, and a GA 3 treatment (250 mg·l –1 in the germination substrate). Embryo growth, time from testa to endosperm rupture and radicle emergence were assessed as separate phases. Epicotyl–plumule emergence was evaluated incubating the germinated seeds at 15°C for 2 weeks, at 5 and 25°C for 2 months on agar water before transplanting to the soil substrate at 10, 15 and 20°C and at 15°C for 2 months on the surface agar water with GA 3 . Important Findings Embryo growth, testa rupture, endosperm rupture (radicle emergence) and growth of the epicotyl were identified as four sequential steps in seeds of P. corsica . Gibberellic acid alone and warm stratification followed by 15°C promoted embryo growth and subsequent seed germination. Cold stratification induced secondary dormancy, even when applied after warm stratification. After radicle emergence, epicotyl–plumule emergence was delayed for ca. 3 months. Mean time of epicotyl–plumule emergence was positively affected by cold stratification (2 months at 5°C) and GA 3 . P. corsica seeds exhibited differential temperature sensitivity for the four sequential steps in the removal of dormancy and germination processes that resulted in the precise and optimal timing of seedling emergence.
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  • 19
    Publication Date: 2016-07-20
    Description: Aims Woody plant-browser systems represent an understudied facet of herbivory. We subjected four genotypes of trembling aspen to artificial browsing, similar to that of a large mammalian herbivore, and applied deer saliva to clipped and unclipped trees to assess: (i) the effects of artificial browsing on aspen growth and phytochemistry of leaves and stems, (ii) genotypic variation in responses and (iii) potential alterations of responses by mammalian saliva. Methods Potted aspen trees were grown outdoors on the University of Wisconsin-Madison campus. The experiment consisted of a fully-crossed, 2 x 2 x 4 randomized complete block design, with two levels of artificial browsing (unclipped and clipped), two levels of saliva application (no saliva and saliva) and four aspen genotypes. To simulate ungulate browsing damage, we removed the upper 50% of the stem of half of the trees by pinching the stem with needle-nosed pliers and then separating it by tearing. For half of the damaged trees, we immediately swabbed the wound with deer saliva. Trees in the unclipped plus saliva treatment were swabbed with saliva at the 50% height mark. To assess the effects of clipping and saliva application, we harvested all trees after 2 months and measured various growth and chemical properties. Growth measurements included height, vertical growth, mass of leaves, stems and roots, leaf number and area and bud set. Chemical parameters included defensive, nutritional and structural components of both foliage and stems. Important Findings Clipping affected most of the growth parameters measured, decreasing tree height, leaf, stem, root and total tree mass and leaf area. Clipped trees had greater vertical growth, more leaves and higher specific leaf area (SLA) than unclipped trees. Deer saliva had little to no effect on plant growth response to the clipping treatment. Terminal budset was delayed by clipping and varied among genotypes but not in response to saliva application. Clipping also affected most of the phytochemical variables measured, reducing defensive compounds (phenolic glycosides and condensed tannins (CTs)) and nutrients (N), but increasing structural components (cellulose and lignin) in both leaves and stems. Saliva had very little effect on tree chemistry, causing only a slight decrease in the amount of CTs in leaves. In general, leaves contained more defensive compounds and nutrients, but much less cellulose, compared with stems. Genotypes differed for all physical and chemical indices, and in tolerance to damage as measured by vertical growth. In addition, for most of the physical and chemical variables measured, genotype interacted with the clipping treatment, suggesting that in natural stands some genotypes will resist or tolerate browsing better than others, affecting forest genetic composition and ultimately forest dynamics.
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  • 20
    Publication Date: 2016-07-20
    Description: Aims In the Core Cape Subregion (CCR), a Mediterranean-climate ecosystem with infertile soils, the legume species Podalyria calyptrata and P. burchellii are in a separate clade to P. leipoldtii and P. myrtillifolia . The closely related species are allopatric, and with the west-east climate gradient and variation in soil nutrient availability in the CCR, it was hypothesized that the two closely related allopatric species would differ in their ecological niche and root:shoot ratio, specific root length (SRL) and organic acid exudation responses to phosphorus (P) supply. Methods With increasing P supply in the glasshouse, we measured plant biomass, leaf nitrogen ([N]), [P], root morphology and release of organic acids. We determined species soil and leaf [N] and [P] and climate in field sites. Important Findings At low P supply, P. calyptrata roots exuded more organic acids than P. burchellii which instead produced roots with a greater SRL, and P. myrtillifolia allocated more biomass to roots than P. leipoldtii . In the field, leaf [P] and climate suggested that P. leipoldtii occupied the most oligotrophic niche followed by P. burchellii and then P. calyptrata and P. myrtillifolia . Closely related allopatric species differed in their mechanisms for P-acquisition and ecological niche, indicating that the environment overrides phylogeny in determining P-acquisition traits for these species, and suggesting that climate regulates nutrient availability, driving distribution and speciation.
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  • 21
    Publication Date: 2013-09-21
    Description: It has been argued that scale is the central problem in ecology ( Levin, 1992 ). Studies on carbon cycles and global climate change, the current major themes in modern ecology, require the interfacing of phenomena that occur on different scales of space, time, and ecological organization. For several decades, tremendous efforts have been made to reveal the general patterns of, and the mechanisms for the global carbon cycles. However, many uncertainties remain, particularly on local to regional scales. To reduce these uncertainties, regional collaborations across the board of nations are required.
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  • 22
    Publication Date: 2013-09-21
    Description: Aims Boreal forest is the largest and contains the most soil carbon among global terrestrial biomes. Soil respiration during the prolonged winter period may play an important role in the carbon cycles in boreal forests. This study aims to explore the characteristics of winter soil respiration in the boreal forest and to show how it is regulated by environmental factors, such as soil temperature, soil moisture and snowpack. Methods Soil respiration in an old-growth larch forest ( Larix gmelinii Ruppr.) in Northeast China was intensively measured during the winter soil-freezing process in 2011 using an automated soil CO 2 flux system. The effects of soil temperature, soil moisture and thin snowpack on soil respiration and its temperature sensitivity were investigated. Important Findings Total soil respiration and heterotrophic respiration both showed a declining trend during the observation period, and no significant difference was found between soil respiration and heterotrophic respiration until the snowpack exceeded 20cm. Soil respiration was exponentially correlated with soil temperature and its temperature sensitivity (Q 10 value) for the entire measurement duration was 10.5. Snow depth and soil moisture both showed positive effects on the temperature sensitivity of soil respiration. Based on the change in the Q 10 value, we proposed a ‘freeze–thaw critical point’ hypothesis, which states that the Q 10 value above freeze–thaw critical point is much higher than that below it (16.0 vs . 3.5), and this was probably regulated by the abrupt change in soil water availability during the soil-freezing process. Our findings suggest interactive effects of multiple environmental factors on winter soil respiration and recommend adopting the freeze–thaw critical point to model soil respiration in a changing winter climate.
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  • 23
    Publication Date: 2013-09-21
    Description: Aims Clear-cutting is a common forest management practice, especially in subtropical China. However, the potential ecological consequences of clear-cutting remain unclear. In particular, the effect of clear-cutting on soil processes, such as the carbon cycle, has not been quantified in subtropical forests. Here, we investigated the response of soil respiration (Rs) to clear-cutting during a 12-month period in a subtropical forest in eastern China. Methods We randomly selected four clear-cut (CC) plots and four corresponding undisturbed forest (UF) plots. Measurements of Rs were made at monthly time points and were combined with continuous climatic measurements in both CC and UF. Daily Rs was estimated by interpolating data with an exponential model dependent on soil temperature. Daily Rs was cumulated to annual Rs estimates. Important Findings In the first year after clear-cutting, annual estimates of Rs in CC (508±23g C m –2 yr –1 ) showed no significant difference to UF plots (480±12g C m –2 yr –1 ). During the summer, soil temperatures were usually higher, whereas the soil volumetric water content was lower in CC than in UF plots. The long-term effects of clear-cutting on Rs are not significant, although there might be effects during the first several months after clear-cutting. Compared with previous work, this pattern was more pronounced in our subtropical forest than in the temperate and boreal forests that have been studied by others. With aboveground residuals off-site after clear-cutting, our results indicate that the stimulation of increasing root debris, as well as environmental changes, will not lead to a significant increase in Rs. In addition, long-term Rs will not show a significant decrease from the termination of root respiration, and this observation might be because of the influence of fast-growing vegetation after clear-cutting in situ .
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  • 24
    Publication Date: 2013-09-21
    Description: Aims Understanding carbon (C) and nitrogen (N) dynamics and their dependence on the stand density of an even-aged, mature forest provides knowledge that is important for forest management. This study investigated the differences in ecosystem total C and N storage and flux between a low-density stand (LD) and a high-density stand (HD) and examined the effects of stand density on aboveground net primary productivity (ANPP), total belowground C allocation (TBCA) and net ecosystem production (NEP) in a naturally regenerated, 65- to 75-year-old Pinus densiflora S. et Z. forest. Methods LD (450 trees ha –1 ) and HD (842 trees ha –1 ) were established in an even-aged, mature P. densiflora forest in September 2006. The forest had been naturally regenerated following harvesting, and the stand density was naturally maintained without any artificial management such as thinning. The diameter at breast height (DBH ≥ 5.0cm) of all live stems within the stands was measured yearly from 2007 to 2011. To compare C and N storage and fluxes in LD and HD, C and N pools in aboveground and belowground biomass, the forest floor, coarse woody debris (CWD) and soil; soil CO 2 efflux ( R S ); autotrophic respiration ( R A ); litter production; and soil N availability were measured. Further, ANPP, TBCA and NEP were estimated from plot-based measurement data. Important Findings Ecosystem C (Mg C ha –1 ) and N (Mg N ha –1 ) storage was, respectively, 173.0±7.3 (mean ± SE) and 4.69±0.30 for LD and 162±11.8 and 4.08±0.18 for HD. There were no significant differences in C and N storage in the ecosystem components, except for soils, between the two stands. In contrast, there were significant differences in aboveground ANPP and TBCA between the two stands ( P 〈 0.05). Litterfall, biomass increment and R S were major C flux components with values of, respectively, 3.89, 3.74 and 9.07 Mg C ha –1 year –1 in LD and 3.15, 2.94 and 7.06 Mg C ha –1 year –1 in HD. Biometric-based NEP (Mg C ha –1 year –1 ) was 4.18 in LD and 5.50 in HD. Although the even-aged, mature P. densiflora forest had similar C and N allocation patterns, it showed different C and N dynamics depending on stand density. The results of the current study will be useful for elucidating the effects of stand density on C and N storage and fluxes, which are important issues in managing natural mature forest ecosystems.
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  • 25
    Publication Date: 2013-09-21
    Description: Aims Tropical forest plays a key role in global C cycle; however, there are few studies on the C budget in the tropical rainforests in Asia. This study aims to (i) reveal the seasonal patterns of total soil respiration ( R T ), litter respiration ( R L ) and soil respiration without surface organic litter ( R NL ) in the primary and secondary Asian tropical mountain rainforests and (ii) quantify the effects of soil temperature, soil moisture and substrate availability on soil respiration. Methods The seasonal dynamics of soil CO 2 efflux was measured by an automatic chamber system (Li-8100), within the primary and secondary tropical mountain rainforests located at the Jianfengling National Reserve in Hainan Island, China. The litter removal treatment was used to assess the contribution of litter to belowground CO 2 production. Important Findings The annual R T was higher in the primary forest (16.73±0.87 Mg C ha –1 ) than in the secondary forest (15.10±0.26 Mg C ha –1 ). The rates of R T , R NL and R L were all significantly higher in the hot and wet season (May–October) than those in the cool and dry season (November–April). Soil temperature at 5cm depth could explain 55–61% of the seasonal variation in R T , and the temperature sensitivity index ( Q 10 ) ranked by R L ( Q 10 = 3.39) 〉 R T (2.17) 〉 R NL (1.76) in the primary forest and by R L (4.31) 〉 R T (1.86) 〉 R NL (1.58) in the secondary forest. The contribution of R L to R T was 22–23%, while litter input and R T had 1 month time lag. In addition, the seasonal variation of R T was mainly determined by soil temperature and substrate availability. Our findings suggested that global warming and increased substrate availability are likely to cause considerable losses of soil C in the tropical forests.
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  • 26
    Publication Date: 2013-09-21
    Description: Aims Understanding of the ecophysiological dynamics of forest canopy photosynthesis and its spatial and temporal scaling is crucial for revealing ecological response to climate change. Combined observations and analyses of plant ecophysiology and optical remote sensing would enable us to achieve these studies. In order to examine the utility of spectral vegetation indices (VIs) for assessing ecosystem-level photosynthesis, we investigated the relationships between canopy-scale photosynthetic productivity and canopy spectral reflectance over seasons for 5 years in a cool, temperate deciduous broadleaf forest at ‘Takayama’ super site in central Japan. Methods Daily photosynthetic capacity was assessed by in situ canopy leaf area index (LAI), (LAI x V cmax [single-leaf photosynthetic capacity]), and the daily maximum rate of gross primary production (GPP max ) was estimated by an ecosystem carbon cycle model. We examined five VIs: normalized difference vegetation index (NDVI), enhanced vegetation index (EVI), green–red vegetation index (GRVI), chlorophyll index (CI) and canopy chlorophyll index (CCI), which were obtained by the in situ measurements of canopy spectral reflectance. Important Findings Our in situ observation of leaf and canopy characteristics, which were analyzed by an ecosystem carbon cycling model, revealed that their phenological changes are responsible for seasonal and interannual variations in canopy photosynthesis. Significant correlations were found between the five VIs and canopy photosynthetic capacity over the seasons and years; four of the VIs showed hysteresis-type relationships and only CCI showed rather linear relationship. Among the VIs examined, we applied EVI–GPP max relationship to EVI data obtained by Moderate Resolution Imaging Spectroradiometer to estimate the temporal and spatial variation in GPP max over central Japan. Our findings would improve the accuracy of satellite-based estimate of forest photosynthetic productivity in fine spatial and temporal resolutions, which are necessary for detecting any response of terrestrial ecosystem to meteorological fluctuations.
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  • 27
    Publication Date: 2013-09-21
    Description: Aims This study aimed to develop radial growth models and to predict the potential spatial distribution of Pinus densiflora (Japanese red pine) and Quercus spp. (Oaks) in South Korea, considering topographic and climatic factors. Methods We used a dataset of diameter at breast height and radial growth estimates of individual trees, topographic and climatic factors in systematic sample plots distributed over the whole of South Korea. On the basis that radial growth is attributed primarily to tree age, we developed a radial growth model employing tree age as an explanatory variable. We estimated standard growth (SG), defined as radial growth of the tree at age 30, to eliminate the influence of tree age on radial growth. In addition, SG estimates including the Topographic Wetness Index, temperature and precipitation were calculated by the Generalized Additive Model. Important Findings As a result of variogram analysis of SG, we found spatial autocorrelation between SG, topographic and climatic factors. Incremental temperature had negative impacts on radial growth of P. densiflora and positive impacts on that of Quercus spp. Precipitation was associated with positive effects on both tree species. Based on the model, we found that radial growth of P. densiflora would be more vulnerable than that of Quercus spp. to climatic factors. Through simulation with the radial growth model, it was predicted that P. densiflora stands would be gradually replaced with Quercus spp. stands in eastern coastal and southern regions of South Korea in the future. The models developed in this study will be helpful for understanding the impact of climatic factors on tree growth and for predicting changes in distribution of P. densiflora and Quercus spp. due to climate change in South Korea.
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  • 28
    Publication Date: 2013-09-21
    Description: Aims Vast grasslands on the Tibetan Plateau are almost all under livestock grazing. It is unclear, however, what is the role that the grazing will play in carbon cycle of the grassland under future climate warming. We found in our previous study that experimental warming can shift the optimum temperature of saturated photosynthetic rate into higher temperature in alpine plants. In this study, we proposed and tested the hypothesis that livestock grazing would alter the warming effect on photosynthetic and respiration through changing physical environments of grassland plants. Methods Experimental warming was carried by using an infrared heating system to increase the air temperature by 1.2 and 1.7°C during the day and night, respectively. The warming and ambient temperature treatments were crossed over to the two grazing treatments, grazing and un-grazed treatments, respectively. To assess the effects of grazing and warming, we examined photosynthesis, dark respiration, maximum rates of the photosynthetic electron transport ( J max ), RuBP carboxylation ( V cmax ) and temperature sensitivity of respiration Q 10 in Gentiana straminea , an alpine species widely distributed on the Tibetan grassland. Leaf morphological and chemical properties were also examined to understand the physiological responses. Important findings 1) Light-saturated photosynthetic rate ( A max ) of G. straminea showed similar temperature optimum at around 16°C in plants from all experimental conditions. Experimental warming increased A max at all measuring temperatures from 10 to 25°C, but the positive effect of the warming occurred only in plants grown under the un-grazed conditions. Under the same measuring temperature, A max was significantly higher in plants from the grazed than the un-grazed condition. 2) There was significant crossing effect of warming and grazing on the temperature sensitivity ( Q 10 ) of leaf dark respiration. Under the un-grazed condition, plants from the warming treatment showed lower respiration rate but similar Q 10 in comparison with plants from the ambient temperature treatment. However, under the grazed condition Q 10 was significantly lower in plants from the warming than the ambient treatment. 3) The results indicate that livestock grazing can alter the warming effects on leaf photosynthesis and temperature sensitivity of leaf dark respiration through changing physical environment of the grassland plants. The study suggests for the first time that grazing effects should be taken into account in predicting global warming effects on photosynthesis and respiration of plants in those grasslands with livestock grazing.
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  • 29
    Publication Date: 2013-09-21
    Description: Aims (i) To explore variations in nutrient resorption of woody plants and their relationship with nutrient limitation and (ii) to identify the factors that control these variations in forests of eastern China. Methods We measured nitrogen (N) and phosphorus (P) concentrations in both green and senesced leaves of 172 woody species at 10 forest sites across eastern China. We compared the nutrient resorption proficiency (NuRP) and efficiency (NuRE) of N and P in plant leaves for different functional groups; we further investigated the latitudinal and altitudinal variations in NuRP and NuRE and the impacts of climate, soil and plant types on leaf nutrient resorptions. Important Findings On average, the leaf N resorption proficiency (NRP) and P resorption proficiency (PRP) of woody plants in eastern China were 11.1mg g – 1 and 0.65 mg g – 1 , respectively; and the corresponding N resorption efficiency (NRE) and P resorption efficiency (PRE) were 49.1% and 51.0%, respectively. Angiosperms have higher NRP (are less proficient) values and lower NRE and PRE values than gymnosperms, but there are no significant differences in NRP, PRP and PRE values between species with different leaf habits (evergreen vs. deciduous angiosperms). Trees have higher NRE and PRE than shrubs. Significant geographical patterns of plant nutrient resorption exist in forests of eastern China. In general, NRP and PRE decrease and PRP and NRE increase with increasing latitude/altitude for all woody species and for the different plant groups. Plant functional groups show more controls than environmental factors (climate and soil) on the N resorption traits (NRP and NRE), while site-related variables present more controls than plant types on PRP and PRE. NRP increases and PRP and NRE decrease significantly with increasing temperature and precipitation for the overall plants and for most groups, except that significant PRE–climate relationship holds for only evergreen angiosperms. Leaf nutrient resorption did not show consistent responses in relation to soil total N and P stoichiometry, probably because the resorption process is regulated by the relative costs of drawing nutrients from soil versus from senescing leaves. These results support our hypothesis that plants growing in P-limited habitats (low latitudes/altitudes or areas with high precipitation/temperature) should have lower PRP and higher PRE, compared with their counterparts in relatively N-limited places (high latitudes/altitudes or areas with low precipitation/temperature). Our findings can improve the understanding of variations in N and P resorption and their responses to global change, and thus facilitate to incorporate these nutrient resorption processes into future biogeochemical models.
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  • 30
    Publication Date: 2013-09-21
    Description: Anthropogenic nitrogen (N) emissions to atmosphere have increased dramatically in China since 1980s, and this increase has aroused great concerns on its ecological impacts on terrestrial ecosystems. Previous studies have showed that terrestrial ecosystems in China are acting as a large carbon (C) sink, but its potential in the future remains largely uncertain. So far little work on the impacts of the N deposition on C sequestration in China’s terrestrial ecosystems has been assessed at a national scale. Aiming to assess and predict how ecological processes especially the C cycling respond to the increasing N deposition in China’s forests, recently researchers from Peking University and their partners have established a manipulation experimental network on the ecological effects of the N deposition: Nutrient Enrichment Experiments in China’s Forests Project (NEECF). The NEECF comprises 10 experiments at 7 sites located from north to south China, covering major zonal forest vegetation in eastern China from boreal forest in Greater Khingan Mountains to tropical forests in Hainan Island. This paper introduces the framework of the NEECF project and its potential policy implications.
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  • 31
    Publication Date: 2013-09-21
    Description: Aims Root and heterotrophic respiration may respond differently to environmental variability, but little evidence is available from large-scale observations. Here we aimed to examine variations of root and heterotrophic respiration across broad geographic, climatic, soil and biotic gradients. Methods We conducted a synthesis of 59 field measurements on root and heterotrophic respiration across China’s forests. Important Findings Root and heterotrophic respiration varied differently with forest types, of which evergreen broadleaf forest was significantly different from those in other forest types on heterotrophic respiration but without statistically significant differences on root respiration. The results also indicated that root and heterotrophic respiration exhibited similar trends along gradients of precipitation, soil organic carbon and satellite-indicated vegetation growth. However, they exhibited different relationships with temperature: root respiration exhibited bimodal patterns along the temperature gradient, while heterotrophic respiration increased monotonically with temperature. Moreover, they showed different relationships with MOD17 GPP, with increasing trend observed for root respiration whereas insignificant change for heterotrophic respiration. In addition, root and heterotrophic respiration exhibited different changes along the age sequence, with insignificant change for root respiration and decreasing trend for heterotrophic respiration. Overall, these results suggest that root and heterotrophic respiration may respond differently to environmental variability. Our findings could advance our understanding on the different environmental controls of root and heterotrophic respiration and also improve our ability to predict soil CO 2 flux under a changing environment.
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  • 32
    Publication Date: 2015-03-28
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  • 33
    Publication Date: 2015-03-28
    Description: Aims Exploring flowering patterns and detecting processes are essential when probing into the nature of reproductive traits during the life history and the interactions among different evolutionary clades. Such patterns are believed to be influenced by many factors, but quantifying these impacts at the community-level remains poorly understood. Methods We investigated the flowering patterns based on long-term herbarium records in a given area from subtropical forest regions in southern China. We obtained 5258 herbarium voucher specimens collected from the Dinghushan National Nature Reserve (DNNR) belonging to 166 families, 943 genera and 2059 species and examined the month when each species was flowering during the period 1920–2007. Important Findings The results showed that plants flowered sequentially almost throughout the whole year, showing the characteristics of subtropical evergreen broad-leaved forests. Flowering spectrums of the entire flora and the four life forms exhibited a clear unimodality that is probably typical of subtropical forest communities. Flowering patterns of the DNNR were positively correlated with mean rainfall, mean air temperature and mean sunshine duration. Median flowering dates of the 38 large species-rich families ranged from early April to late August; 25 families exhibited significant unimodal distribution patterns, whereas the remaining families were unclear or bimodal. Median flowering dates of the 10 most species-rich genera ranged from middle May to later July. While the results are consistent with climatic factors playing a general role in flowering patterns, median flowering dates varied significantly among species-rich families and genera, suggesting that phylogenies could provide specific constraints in subtropical forests.
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  • 34
    Publication Date: 2015-03-28
    Description: Aims Most pollinator fig wasps are host plant specific, with each species only breeding in the figs of one fig tree species, but increasing numbers of species are known to be pollinated by more than one fig wasp, and in rare instances host switching can result in Ficus species sharing pollinators. In this study, we examined factors facilitating observed host switching at Xishuangbanna in Southwestern (SW) China, where Ficus squamosa is at the northern edge of its range and lacks the fig wasps that pollinate it elsewhere, and its figs are colonized by a Ceratosolen pollinator that routinely breeds in figs of F. heterostyla . Methods We recorded the habitat preferences of F. squamosa and F. heterostyla at Xishuangbanna, and compared characteristics such as fig size, location and colour at receptive phase. Furthermore, the vegetative and reproductive phenologies in the populations of F. squamosa and F. heterostyla were recorded weekly at Xishuangbanna Tropical Botanical Garden for 1 year. Important Findings Ficus squamosa is a shrub found near fast-flowing rivers, F. heterostyla is a small tree of disturbed forest edges. Although preferring different habitats, they can be found growing close together. Both species have figs located at or near ground level, but they differ in size when pollinated. Fig production in F. squamosa was concentrated in the colder months. F. heterostyla produced more figs in summer but had some throughout the year. The absence of its normal pollinators, in combination with similarly located figs and partially complementary fruiting patterns appear to have facilitated colonization of F. squamosa by the routine pollinator of F. heterostyla . The figs probably also share similar attractant volatiles. This host switching suggests one mechanism whereby fig trees can acquire new pollinators and emphasizes the likely significance of edges of ranges in the genesis of novel fig tree–fig wasp relationships.
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  • 35
    Publication Date: 2015-03-28
    Description: Aims Within inflorescences of sexually reproducing hermaphrodites, the production of ovules, fruits and seeds commonly declines from basal (early-opening) to distal (late-opening) flowers, while pollen production remains constant or only changes slightly, with the result that distal (late-opening) flowers become functionally male. However, few empirical studies have specifically examined whether or not changes in allocation to pollen production actually lead to changes in the number of seeds sired, a more direct measure of male fitness. In pseudogamous apomicts, fitness depends on the number of seeds produced; thus, a contrasting pattern of variation in the pollen-to-ovule (P/O) ratio within inflorescences might be expected. Methods We investigated floral sex allocation and reproductive success within racemes of Hosta ventricosa , a pseudogamous apomictic hermaphrodite possessing flowers that open acropetally. We quantified variations in pollen number, ovule number, the P/O ratio and fruit and seed production, from 2007 to 2011, among flowers within racemes of four populations of H. ventricosa in southwest China. Ecological causes for fruit and seed production were evaluated by observing patterns of pollen deposition, flower removal and supplemental pollination. Important Findings Pollen number, ovule number and the P/O ratio declined from basal-to-distal positions in all sampled populations (years). Fruit and seed production decreased distally in most populations (years). Low fruit and seed set of distal flowers was not due to pollen limitation because pollen deposition never declined distally and the low fruit and seed set of distal flowers remained even after supplementary pollination was provided. The flower-removal experiment indicated that inter-fruit competition for resources among flowers was common. The low P/O ratio of distal flowers in H. ventricosa might be favored because they were unable to obtain fitness by donating pollen and siring seeds on other plants. Our study may help to understand the adaptive significance of sex allocation among flowers within inflorescences of sexually reproducing hermaphrodites.
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  • 36
    Publication Date: 2016-03-25
    Description: Aims Both high and low densities of macrophyte vegetation can impair its ecosystem service function. Harvesting is often applied to macrophyte vegetation to maintain an appropriate density. Vegetation harvesting has occasionally gone awry and caused catastrophes, such as vegetation disappearance and cyanobacterial dominance in waterways and lakes. Because water depth influences macrophyte density at all life stages, the simultaneous influences of harvesting and water depth should be carefully examined. Thus, this study aims to quantify the effects of differently harvesting Elodea nuttallii on its growth and reproduction at different water depths in field experiments. Methods Four harvest intensities (harvesting E. nuttallii plant heights equal to 25%, 50%, 75% and 100% of the water depth) were applied to E. nuttallii growing at four different water depths (60, 90, 120 and 150cm). Plant length and root length were measured. The node number, root number of each plant and number of floating plants were counted before harvesting. The harvested plant were dried to a constant weight for dry weight determination. Important Findings The rate of increase in the length and shoot number of E. nuttallii varied from –0.012 to 0.440 day –1 and from –0.020 to 0.639 day –1 , respectively. Water depth 〉150cm would limit E. nuttallii growth. Elodea nuttallii responded to increasing water depths and low-intensity harvesting by increasing internodal length and decreasing shoot number. The larger internodal length of E. nuttallii observed in relatively deeper water was also induced by the physical strain generated by its buoyancy as its specific gravity was less than water’s. The physical mechanism of removing the plant canopy by harvesting decreased E. nuttallii buoyancy and prevented floating. Harvesting increased plant production in shallow waters 〈90cm deep. Moreover, it is also necessary to perform three medium-intensity harvests at a water depth of 120cm and one low-intensity harvest or no harvesting at a water depth of 150cm to achieve longer lifetimes and less biomass near the water surface when the plants reach or approach the water surface.
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  • 37
    Publication Date: 2016-03-25
    Description: Aims Distyly has been regarded as an adaptation to improve compatible pollination between two floral morphs with reciprocal herkogamy. The hypothesis that the different positions of anthers and stigmas within flowers as well as their reciprocal position between morphs, reduce the probability of self pollination raised by Darwin has been rarely tested. In this study, we measured stigmatic pollen loads in response to reduced reciprocal herkogamy in two Primula species. Methods To see whether reciprocal herkogamy can increase compatible and/or reduce incompatible pollen deposition, thus promoting compatible pollination, we shortened the distance between anthers and stigmas within the flowers by changing the position of the corolla tube, to which the anthers were fused, i.e. reduced herkogamy in natural populations of Primula secundiflora and P. poissonii and quantified stigmatic pollen loads in the field over 2 years. Important Findings In both species, stigmatic pollen loads were significantly higher in the long-styled (L-morph) than in the short-styled morph (S-morph) in both control and manipulated flowers, but percentage of compatible pollen in S-morph were higher. Flowers manipulated to halve the anther–stigma distance showed a similar pattern for 2 years: total pollen grain counts on stigmas did not differ significantly but compatible pollen grains in L- and S-morphs were significantly decreased in both species. The percentage of compatible pollen loads was decreased by 68.7% in P. secundiflora and 65.3% in P. poissonii in L-morphs, while it decreased by 30.6% and 2.9% in S-morphs, respectively. Our manipulation of the relative position of anthers and stigmas in the two distylous species indicated that a lower degree of herkogamy reduced compatible but incompatible pollen transfer was likely to increase. The higher proportion of compatible pollen in the S-morph than in the L-morph in the two Primula species could be attributed to the accessibility of two-level sexual organs, floral orientations and pollinator behaviors. This is a first attempt to manipulate intraflower herkogamy for understanding adaptation of heterostyly, shedding insights into how the reciprocal herkogamy promotes compatible pollination.
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  • 38
    Publication Date: 2016-03-25
    Description: Aims Deserts are one of the ecosystems most sensitive to global climate change. However, there are few studies examining how changing abiotic and biotic factors under climate change will affect plant species diversity in the temperate deserts of Asia. This study aimed to: (i) characterize species distributions and diversity patterns in an Asian temperate desert; and (ii) to quantify the effects of spatial and environment variables on plant species diversity. Methods We surveyed 61 sites to examine the relationship between plant species diversity and several spatial/environmental variables in the Gurbantunggut Desert. Spatial and environmental variables were used to predict plant species diversity in separate multiple regression and ordination models. Variation in species responses to spatial and environmental conditions was partitioned by combining these variables in a redundancy analysis (RDA) and by creating multivariate regression trees (MRT). Important Findings We found 92 plant species across the 61 sites. Elevation and geographic location were the dominant environmental factors underlying variation in site species richness. A RDA indicated that 93% of the variance in the species–environment relationships was explained by altitude, latitude, longitude, precipitation and slope position. Precipitation and topographic heterogeneity, through their effects on water availability, were more important than soil chemistry in determining the distribution of species. MRT analyses categorized communities into four groups based on latitude, soil pH and elevation, explaining 42.3% of the standardized species variance. Soil pH strongly influenced community composition within homogeneous geographic areas. Our findings suggest that precipitation and topographic heterogeneity, rather than edaphic heterogeneity, are more closely correlated to the number of species and their distributions in the temperate desert.
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  • 39
    Publication Date: 2016-03-25
    Description: Aims The introduction of potentially invasive species through ornamental cultivation or for rehabilitation purposes is a serious environmental problem. They cause damage to biodiversity through loss, increased mortality or ‘ in situ ’ selection phenomena in natural flora. Spartium junceum is a Mediterranean shrub that is not native in most areas of the Iberian Peninsula, although it is extensively grown for the rehabilitation of roadsides. We have investigated the effect on the native vegetation of an old S. junceum (Fabaceae) plantation in a conservation area in the centre of the Iberian Peninsula: the Cuenca Alta del Manzanares nature reserve in Madrid. Methods We compared S. junceum stands with the native nanophanerophytic Cistus ladanifer community at different ecosystem properties: soil properties, temporal soil seed bank contents, standing vegetation and net primary production of annual grasslands growing in these shrublands. Important Findings The results highlighted S. junceum ’s ability to become established in the new environment (marginal areas of the nature reserve) and ensure its successful growth. This is more apparent in northern and eastern exposures where this formation contacts with the core of the best conserved native vegetation in the nature reserve. Soils under Spartium showed a higher nitrogen content, indicating its capacity—in common with other legumes—to fix N, and conferring an advantage over Cistus , which is N-limited. Other soil nutrients such as phosphorus, magnesium and calcium and water availability are also higher in Spartium soils than in Cistus . Phosphorus is usually a constraint for N-fixers, but in our study, it is the most significant soil variable in both shrub formations and is important to the success of Spartium . Water availability is a key factor for Mediterranean vegetation, and particularly in autumn when soils are recharged. The Spartium formation is able to retain water as its growth produces a closer canopy than Cistus , thereby preventing water evaporation and contributing to the success of this species. Perennials are more frequent in the standing vegetation than in the seed bank, whereas therophytes are similar. The standing vegetation has therophytes and chamaephytes as the predominant growth forms in Spartium sites, and hemicryptophytes and phanerophytes in Cistus . Therophytes are dominant in Spartium and Cistus seed banks, although the first formation has more species. Spartium has a higher number of hemicryptophytes and Cistus is higher in phanerophytes. Northern and eastern aspects show significant differences in richness—with a predominance of annual weed species in Spartium —and in above-ground net primary production, probably as a consequence of the nutrients present in the soils. Ruderal annual species under Spartium ( Bromus tectorum , Chenopodium album ) have higher germination rates in the greenhouse than in the standing vegetation, suggesting they are at potential risk if environmental conditions change.
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  • 40
    Publication Date: 2016-03-25
    Description: Aims Positive interactions are defined as non-trophic interactions where at least one of the interacting species is benefited in terms of fitness and the other remains unaffected. Nevertheless, the bidirectional feedbacks between species may be positive, neutral or negative. Thus, if facilitated species induce negative effects on their ‘nurses’, the assumed definition of positive interactions could be reconsidered. Methods We assessed if ecological interactions between cushions of Azorella madreporica and their facilitated species are positive. Specifically, we tested if cover of facilitated species has any costs for cushion plants from an ecophysiological perspective, and if these costs increase with the amount of cover of facilitated species. In addition, through pathway analysis and correlations, we assessed if cover and richness of facilitated species have a direct and/or indirect effect on the fitness of cushion plants. Important Findings We found that facilitated plant species induced a significant cost for their nurses (cushion plants), and this cost increases with cover of the facilitated species. Additionally, the facilitated species exert a strong direct negative effect on the cushion’s fitness and a moderate indirect negative cost evident through the nutrient status and physiological performance of cushion plants. We thus contribute evidence that positive interactions between high mountain cushion plants of central Chile and their ‘facilitated’ species may be an artifact more than a fact, especially when bidirectional effects are considered; contrasting with the majority of studies that document only one side of the interaction.
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  • 41
    Publication Date: 2016-03-25
    Description: Aims Plants of similar life forms and closely related species have been observed to create similar types of plant–soil feedbacks (PSFs). However, investigations of the consistency of PSFs within species have not yielded clear results. For example, it has been reported that species create different types of PSFs in their native and introduced ranges. The aim of this project is to examine if four species create similar PSF types from soils collected from widely distributed areas within their introduced range. The soil for this project was collected from three areas in western North America. With this design, we aim to determine species- and site-specific ability to create PSFs and if the type of PSF created is consistent in all soil from all three collection areas. The species examined are Agropyron cristatum , Centaurea solstitialis, Poa pratensis and Taeniatherum caput-medusae. Methods We used three-field collected soils (from northern Nevada, western Montana and eastern Montana) in a two-phase greenhouse experiment to quantify the type of PSFs created by four invasive species. The first phase was a conditioning phase wherein each invasive species created species-specific changes to the soil. The second phase of the experiment was the response phase wherein both the conditioning species and a native phytometer were grown in the conditioned soil and in unconditioned (control) soil. The final aboveground biomass was used to evaluate the effect of conditioning and to determine the type of PSF created by each invasive species. Important Findings Our results suggest that three of our four study species did show consistency in relation to PSF. Two species A. cristatum and T. caput-medusae consistently created PSF types that benefit conspecifics more than heterospecifics (and thus are ‘invasive’ PSF types) and P. pratensis consistently exhibited no, or ‘neutral’, feedbacks. The fourth species ( C. solstitialis ) was inconsistent: in one soil, no feedback was created; in other soil, an invasive PSF was created and in the last soil, a feedback that relatively benefited the native phytometer was created. Thus, PSFs appear to uniformly contribute to the success of two species ( A. cristatum and T. caput-medusae ) but not C. solstitialis nor P. pratensis.
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  • 42
    Publication Date: 2015-03-28
    Description: Aims Small plant populations may be more likely to suffer more severe pollen limitation due to the lower number of potential mates or suitable pollinators. For invasive species, this phenomenon may be more common when an invading population colonizes a new habitat. Here, we investigated whether pollen limitation occurs in invasive populations of Solanum rostratum during its invasion from North America to China and evaluated the patterns between pollen limitation and population size. Methods Pollen addition experiments were performed on six invasive populations of S. rostratum . By comparing fruit set and seed production with open pollination treatment, we calculated the index of pollen limitation and regressed it to population size and density. Important Findings Among the six sampled invasive populations of S. rostratum , the fruit set and seed production per fruit were 0.346±0.014 and 52.38±9.29, respectively, with open pollination treatment and 0.572±0.022 and 56.28±10.79, respectively, with pollen addition treatment. Compared with open pollination, pollen addition significantly increased fruit set and seed production by 65.3 and 7.4%, respectively. The standardized index of pollen limitation ranged from 0.022 to 0.125, with an average of 0.065, suggesting that invasive populations of S. rostratum do suffer from pollen limitation. The index of pollen limitation was negatively correlated with population size, which is consistent with the pattern that smaller populations suffer from more severe pollen limitation.
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  • 43
    Publication Date: 2015-03-28
    Description: Aims Floral longevity, the duration that a flower remains open and functional, varies greatly among species. Variation in floral longevity has been considered to be optimal strategy for resource allocation under different ecological conditions, mainly determined by the rates of pollination and cost of flower maintenance. However, it is unclear whether an intrinsic factor, floral sexual investment, constrains evolution of floral longevity. The theoretical model also predicts that dichogamy favors long-lived flowers, but empirical studies to test this prediction remain unexplored. Methods To examine the effect of floral sexual investment on floral longevity, we measured flower size together with pollen and ovule production in 37 sympatric flowering plants in a natural community. The duration of the female and male phase in 21 protandrous species and floral longevity of the other 16 adichogamous species were documented in the field. Important Findings Floral longevity varied from 1 day to 15 days, while pollen number per flower varied from 643 to 710880 and ovule number per flower from 1 to 426 in the 37 species. Flower size was correlated with pollen production as well as ovule production. Floral longevity was positively related to pollen production but not to ovule production. Consistent with the prediction that dichogamy favors long-lived flowers, we found the floral longevity of protandrous species was significantly longer than that of adichogamous species. In the protandrous species, pollen production per flower was observed to be positively related to male duration, while ovule production was not related to female duration. Our analyses of variation in floral longevity and sexual investment among different species suggest that the floral sexual investment could be an intrinsic factor contributing to the selected floral longevity, particularly the male phase, and that high pollen production could potentially increase pollen removal, i.e. male productive success.
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  • 44
    Publication Date: 2015-03-28
    Description: Aims The hypothesis of predator satiation has been proposed to explain mast fruiting in various flowering plants. It considers that the simultaneous production of large numbers of seeds by a plant population reduces the risk of seed predation for each individual. Orchids produce huge numbers of seeds per fruit and rarely experience seed predation. It remains unclear which factors may affect fluctuating fruit production in orchids, which generally suffer a widespread pollen limitation. To explore the temporal pattern of fruiting and potential factors related to fluctuation in fruit production, we investigated reproductive success of a long-spurred orchid ( Habenaria glaucifolia ) in an alpine meadow with thousands of individuals over 8 years. Methods To estimate reproductive success, pollinator observation was conducted by day and at night, and pollinia removal and receipt were recorded in the field population for 8 years. To examine whether fruit set and seed set are pollen limited, we conducted supplementary pollination experiments and compared fruit set, seed set and pollinia movement of open-pollinated flowers from 2011 to 2013. We measured lengths of spurs and pollinator proboscises, and nectar volume and concentration, to identify potential pollinators. Important Findings Hawkmoths were seen to be effective pollinators for H. glaucifolia in 3 years, whereas in the remaining 5 years no pollinators were observed, and consequently pollinia were rarely transferred. Numerous pollinia movements were observed in 2012, 2013 and 2014 (pollinia removal: 48, 59 and 85%; pollinia receipt 51, 70 and 80%), and correspondingly fruit set was significantly higher in 2012 and 2013 (59 and 46%) than in 2011 (25%). It was fruit set, rather than seed set, that was pollen limited in this orchid in the 3 years, in that supplementary pollination increased fruit set but did not increase seed set per fruit compared to natural. Three species of hawkmoths had proboscis lengths that matched the spur length of H. glaucifolia . Fruit set in this long spurred orchid depends on the activity of long-tongued hawkmoths, resulting in significant temporal variation in fruit production. Mast fruiting in this alpine orchid could be attributed to a ‘sit and wait’ strategy, awaiting an abundance of effective pollinators.
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  • 45
    Publication Date: 2015-03-28
    Description: Aims Sex allocation in plants is often plastic, enabling individuals to adjust to variable environments. However, the predicted male-biased sex allocation in response to low resource conditions has rarely been experimentally tested in hermaphroditic plants. In particular, it is unknown whether distal flowers in linear inflorescences show a larger shift to male allocation relative to basal flowers when resources are reduced. In this study, we measure position-dependent plasticity of floral sex allocation within racemes of Aconitum gymnandrum in response to reduced resource availability. Methods Using a defoliation treatment in the field applied to potted plants from a nested half-sibling design, we examined the effects of the treatment, flower position, family and their interactions. Important Findings Allocation to male function increased with more distal flower position, while female allocation either did not change with position or declined at the most distal flowers. Defoliation significantly reduced the mass of both the androecium and gynoecium, but not anther number or carpel number. Gynoecial mass declined more strongly with defoliation than did androecial mass, resulting in a significant increase in the androecium/gynoecium ratio as predicted by sex allocation theory. Plastic responses of androecium mass and gynoecium mass were affected by flower position, with less mass lost in basal flowers, but similar plastic magnitude in both sexual traits across flower position lead to consistent variation in the androecium/gynoecium ratio along the inflorescence. A significant treatment*paternal family interaction for the androecium/gynoecium ratio is evidence for additive genetic variation for plastic floral sex allocation, which means that further evolution of allocation can occur.
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  • 46
    Publication Date: 2015-03-28
    Description: Aims When sympatric flowering plant species in a natural community share pollinators, study of plant–plant interactions via interspecific pollen transfer (IPT) is essential for understanding species coexistence. However, little is known about the extent of IPT between interactive species and its causes. Methods To explore how sympatric flowering plants sharing pollinators minimize deleterious effects of IPT, we investigated the pollination ecology of two endemic species, Salvia przewalskii and Delphinium yuanum , in an alpine meadow in the Hengduan Mountains, southwest China. We quantified conspecific and interspecific visits by shared bumblebee pollinators, amounts of pollen placed on different body sites of the pollinators and stigmatic pollen loads on open-pollinated flowers. To examine whether IPT affects female fitness, we measured pollen germination and seed production in the two species in an artificial pollination experiment. Important Findings One bumblebee species, Bombus trifasciatus , was found to be the sole effective pollinator for the two coflowering species. Pollination experiments indicated that deposition of heterospecific pollen could significantly decrease seed set in both species. Experiments showed that S. przewalskii pollen could germinate well on stigmas of D. yuanum , inhibiting conspecific pollen germination in D. yuanum . However, seed set was not lower under open pollination than under cross-pollination within species, suggesting that no female fitness loss was caused by IPT. In foraging bouts with pollinator switches, switches from S. przewalskii to D. yuanum were relatively more frequent (8.27%) than the converse (1.72%). However, IPT from S. przewalskii to D. yuanum accounted for only 1.82% of total stigmatic pollen loads while the reverse IPT to S. przewalskii was 8.70%, indicating that more switches of bumblebees to D. yuanum did not result in higher IPT. By contrast, selection for reduced IPT to S. przewalskii would limit pollinator switches from D. yuanum . We found that a bumblebee generally carried pollen grains from both species but the two species differed in the position of pollen placement on the bumblebee’s body; S. przewalskii ’ s pollen was concentrated on the dorsal thorax while D. yuanum ’ s pollen was concentrated ventrally on the head. This differential pollen placement along with pollinator fidelity largely reduced IPT between the two species with a shared pollinator.
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  • 47
    Publication Date: 2015-03-28
    Description: Aims Studying the ecological significance of highly specialized morphological traits evolved by alpine plants could help us to understand the adaptation and evolution of these plants under severe alpine environment. We explored the adaptive significance of woolly and overlapping leaves for reproduction in Eriophyton wallichii , a perennial herb native to the subnival belt of the Himalaya-Hengduan Mountains. Methods We examined whether the trichomes could influence the leaf wettability, temperature and leaf reflectance spectra in the lab. And we investigated the thermal benefits of the woolly and overlapping leaves for flowers and fruits in the field. Pollen viability and seed germination were also examined in the lab to assess whether these leaves enhance reproductive fitness. Important Findings Our results showed that dense trichomes impart good water repellency, absorption of solar radiation and accumulation of leaf heat. The woolly and overlapping leaves increased the interior temperature of flowers and fruits to an optimal level on sunny hours, but prevented them from overheating when transient intense solar radiation occurs. This kept optimal temperatures in plants’ reproductive organs, thus promoting the development of pollen and seed in alpine environment.
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  • 48
    Publication Date: 2015-03-28
    Description: Aims It is generally accepted that visual displays and floral scent play important roles in communication between flowering plants and their pollinators. However, the relative role of visual and olfactory cues in pollinator attraction is largely unknown. In this study, we determined the roles of both types of cue in attracting pollinators to Cornus capitata , a medium sized tree with each capitulum surrounded by four large, white, petaloid bracts. Methods Pollinator observations and pollination experiments were conducted in a natural population; the inflorescences’ visual and olfactory signals were characterized by spectral and chemical analyses; the responses of pollinators to visual and olfactory cues were tested using dual-choice behavioural bioassays; the relative roles of visual and olfactory cues in pollinator attraction were tested by comparing the responses of pollinators to inflorescences subjected to three experimental treatments (intact, all bracts removed, and capitulum removed) within the natural population. Important Findings For fruit set, C. capitata is entirely dependent on pollinators, with a bee, Anthophora sp., being the main pollinator. Bracts present high colour distance and green contrast against the leaves. Twelve volatile compounds in the floral scent were detected, most of which have previously been reported to be attractive to a broad spectrum of bee species. Behavioural bioassays showed that both, visual cues alone and olfactory cues alone, are attractive to pollinating bees. However, visual cues alone attracted significantly more approaches than olfactory cues alone, while olfactory cues alone elicited a significantly higher landing percentage than visual cues alone. The finding suggests that, in the C. capitata – Anthophora sp. interaction, visual cues are mainly used for location from long distances, while olfactory cues mainly aid landing from short distances. Our results indicate that different modalities of floral cues should be considered together to understand fully the communication between flowering plant and pollinators.
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  • 49
    Publication Date: 2015-03-28
    Description: Aims Flowering time has been suggested to be an important adaptive trait during the dispersal of invasive species, and identifying the molecular mechanisms underlying flowering time may provide insight into the local adaptation during the process of invasion. Here, we conducted a preliminary exploration on the genetic basis of the differentiation of flowering time in Ambrosia artemisiifolia . Methods Using relative real-time fluorescent quantitative polymerase chain reaction, we investigated the expression levels of eight flowering-related genes, including AP1 , FT , SOC1 , CRY2 , FKF1 , GI , CO2 and SPY , in leaves and flowers at different time points in individuals from northern Beijing and southern Wuhan populations that exhibit significant differences in flowering times to identify any rhythmic changes in gene expression and their association with differential flowering times. Important Findings The differentiation of flowering time in the A. artemisiifolia populations was closely associated with five genes involved in flowering pathways. The floral pathway integrators FT and SOC1 and floral meristem identity gene AP1 exhibited increased expression during flowering. The photoreceptor CRY2 in the light-dependent pathway and the SPY gene in the gibberellin pathway displayed specific expression patterns over time. In earlier-flowering Beijing plants, CRY2 expression was lower and SPY expression was higher than in Wuhan plants. The expression patterns of these five genes suggest a molecular basis for the differentiation of flowering time in A. artemisiifolia .
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  • 50
    Publication Date: 2015-03-28
    Description: Aims Hybridization usually leads to gene introgression between related species in hybrid zones, associated with complex patterns of morphological variation. Nevertheless, previous studies have tended to ignore the effects of geographic variation in hybridization rates on species taxonomy. This study aims to investigate the variation of morphological traits between two sympatric and taxonomically confused oak species, Quercus liaotungensis and Q. mongolica , and reveal the effects of hybridization rates on morphological traits and the taxonomic boundary. Methods We used seven microsatellite loci to evaluate species status and measured 15 morphological traits in 26 trees in the recent hybrid zone between Q. liaotungensis and Q. mongolica , and we characterized the differences between the two oak species and their hybrids for the investigated traits. Important Findings Molecular analyses indicated that 74% of 78 sampled maternal trees were hybrids between Q. liaotungensis and Q. mongolica although the observed morphological variation suggested that they had remained distinct species. Across all of the differentiated leaf and reproductive traits, the hybrids expressed patterns similar to Q. liaotungensis , which may suggest dominant expression of parental characters. These results are consistent with our expectation that hybrids will be difficult to distinguish from parental species in a recent hybrid zone.
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  • 51
    Publication Date: 2015-03-28
    Description: Aims The calyx, the outermost whorl of a flower (usually green), has been considered to function to protect flowers. In some species, however, calyces are colorful and retained during seed development. Limonium species have been exploited as cut flower crops because the calyces persist for several months after the corolla has closed. To explore the adaptive significance of the persistent calyx in a desert plant Limonium leptolobum , we ask whether persistence of calyces can enhance pollinator attraction by enlarging floral displays, increasing reproductive success in this self-incompatible species. Methods The yellow flower of L. leptolobum lasted 1–2 days but its white, membranous calyx extended fully after the corolla closed, and persisted for over 2 months in the field, making hundreds of ‘showy flowers’ on one individual. To examine the ecological function of calyces, we test the pollinator attraction hypothesis. In an experimental population, we compared the difference in visit frequency and visitor behavior between intact inflorescences and inflorescences with their calyces removed on the same individual plants. Important findings In four experimental plots four types of floral visitors were observed including bees, butterflies, syrphid flies and day-flying moths. No significant preference was observed between calyx-free and intact inflorescences for both first arrivals and total visit frequency of all types of floral visitors, indicating that the persistence of calyces did not make plants more attractive to potential pollinators. The pollinator attraction hypothesis for the showy calyces was not supported by the current data. Whether the calyx in this desert plant helps seed development or has other functions needs further study.
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  • 52
    Publication Date: 2015-03-28
    Description: Aims Floral nectar plays a vital role in plant reproductive success by attracting pollinators. Nectar traits of a flower can depend directly on plant characteristics other than environmental factors and exhibit extensive flower- and plant-level variations. Studies on nectar traits frequently focused on intraplant variation for dichogamous plants, but few have paid attention to both intra- and interplant nectar variations in relation to plant characteristics. Revealing within- and among-plant variation and its relative magnitude is important for our understanding of how pollinator-mediated selection can act on nectar traits and evolution of nectar traits. Methods Through investigating protandrous Aconitum gymnandrum populations at the Alpine Meadows and Wetland Ecosystems Research Station of Lanzhou University, we examined the relationships between nectar production per flower and plant characteristics (e.g. flower position within inflorescences, floral sexual phases, flowering time, inflorescence size and floral attractive traits). Important Findings A. gymnandrum exhibited a declining gradient in the nectar volume along inflorescences, with more nectar in basal flowers than distal ones. Protandrous flowers of A. gymnandrum did not show gender-biased nectar production while the nectar volume varied with different stages of floral sexual phases. The significant correlation between the first flowering date of individuals and the mean nectar volume per flower was positive in 2013, but became negative in 2014, suggesting complex effects of biotic and abiotic factors. The mean nectar volume per flower was not related to inflorescence size (the number of total flowers per plant). Furthermore, nectar production was weakly associated with floral attractive traits (the petal width and the galea height), even if the effect of flowering time of individuals was removed, suggesting that the honesty of floral traits as signals of nectar reward for pollinators is not stable in this species.
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  • 53
    Publication Date: 2015-03-28
    Description: Aims Exploring the characteristics and function of a flower color polymorphism contributes to our understanding of floral evolution in angiosperms. However, little information is available on stigma color polymorphisms in flowering plants despite their important functional role in plant reproduction. Methods We studied a stigma color polymorphism at the individual level in Acer pictum subsp. mono (aka A. mono ) by investigating stigma-color morph proportion and comparing stigma performance and components of female fitness (pollen adhesion, pollen germination, fruit set, seed set and fruit weight) between the two morphs (red and greenish-yellow stigma) within natural populations at Dongling Mountain in the north of China. In the flowering period, we conducted hand pollination in natural populations and then made microscopic observations using the aniline blue fluorescence method. Important Findings Individuals in the studied populations did not change their stigma color between years, and flowers produced by a single tree were uniform in their stigma color. This strongly suggests that stigma color variation is genetically controlled. The percentage of the red stigma flowers with germinated pollen grains was significantly higher than that of the greenish-yellow stigma flowers when hand pollination was conducted in the early flowering period, but not so in the late flowering period. There was no significant difference in the percentage of flowers with pollen adhesion to the stigma between the two morphs. Fruit set of the red morph was significantly higher than that of the greenish-yellow morph. Our findings suggest that the red morph may be more advantageous for pollen germination and fruit set than the greenish-yellow morph, which may provide a functional explanation for the high incidence of the red stigma morph in the studied populations. Alternative explanations for the stigma color polymorphism are discussed to stimulate further work.
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  • 54
    Publication Date: 2015-03-28
    Description: Aims In heterantherous plants, ‘division of labor’ among structurally different stamens, i.e. pollinating and feeding functions, has been thought to reduce the evolutionary conflict of relying on pollen both as the carrier of male gametes and as the food for pollinators. The key to successful division of labor among different sets of stamens is the size match between stamens and legitimate pollinators, which results in the precise deposition of pollen onto specific locations on pollinator’s body and facilitates cross pollination. However, the potential impact of small illegitimate insects that are ubiquitous during the pollination process on the plant reproduction in heterantherous species has been largely neglected in previous studies and never been demonstrated experimentally. Methods Here, we investigated the functions of three different types of stamens in Commelina communis . The pollinator visitation, pollen removal and deposition were compared among flowers with different types of anthers emasculated at two natural populations. Moreover, the mating systems of C. communis in wild populations were estimated using microsatellite markers. Important Findings Our data showed that the main floral visitors for C. communis at the two studied populations were small illegitimate bees rather than legitimate pollinators, accounting for 77.5 and 92.2% of total flower visits, respectively. Flower manipulations in C. communis demonstrated that the two types of brightly yellow stamens separately functioned as ‘deceptive attraction’ and ‘feeding’ functions. Although the brown inconspicuous stamens of C. communis with the largest amount of fertile pollen had the potential function in offering pollen for cross pollination, the high ratio of illegitimate visitation by small bees significantly affected the dispersal and deposition of pollen from the pollinating anthers, and subsequently decreased the levels of outcrossing ( t m = 0.23–0.32) in wild populations. Our work further confirmed that the size match between pollinators and the floral morphology is the prerequisite to successfully fulfill the functional differentiation among different sets of stamens in heterantherous plants. Local high ratio of illegitimate visitation by size unmatched insects could significantly weaken the potential functions of heteranthery, affecting the dispersal and deposition of functional pollen in heterantherous plants and further the whole mating systems.
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  • 55
    Publication Date: 2016-09-21
    Description: Aims Facilitation by nurse plants is a common interaction in harsh environments and this positive plant–plant interaction may promote vegetation recovery in ecosystems affected by human activities. Determining the relevance of this process, however, requires assessing how nurse plants influence the establishment of other species, as well as the proportion of species in the regional species pool that would benefit from the presence of nurse plants in human-disturbed areas. Further, since vegetation recovery is a time-dependent process, the community-level consequences of facilitation are likely to vary among landscapes with different disturbance history. Thus, an integrative perspective of the relevance of nurse plants for vegetation recovery could be obtained by measuring their effects across different human-disturbed landscapes of the target region. This study focuses on these issues and uses a regional-scale approach to assess the community-level effects of a widespread nurse plant of American deserts, the creosotebush ( Larrea tridentata ). Methods This study was conducted in the southernmost portion of Chihuahuan Desert because most floodplain valleys of this region have been affected by human activities during the past centuries. For this study, we selected 10 floodplain valleys differing in their age (i.e. the time elapsed after human activities were ceased). At each landscape, we measured the cover of creosotebushes and the proportion of plant species positively associated with them, as well as the density of seeds in the soil beneath creosotebush canopies. All these data were regressed against the age of the landscapes. Further, to assess whether positive association patterns were due to facilitation or other processes, we conducted field experiments and measured the ecophysiological performance of plant species established beneath and outside creosotebush canopies. Important Findings Most plant species from the target region were positively associated to creosotebushes, and our field experiments and ecophysiological measures indicated that these distribution patterns can be attributed to facilitative interactions. In most landscapes, the density of seeds was higher beneath creosotebushes than in the surrounding habitats, suggesting that these shrubs may also act as seed traps. The community-level effects of creosotebushes increased with landscape age and creosotebush cover, indicating that magnitude of these effects depends on the disturbance history of each site. These results highlight the relevance of performing large-scale assessments for identifying the consequences of facilitation on vegetation recovery across space and time. We then propose that this kind of large-scale approach should be taken into account in the development of conservation programs aimed at the recovery and preservation of plant biodiversity in harsh environments.
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  • 56
    Publication Date: 2016-09-21
    Description: Aims Forest fragmentation and the associated augmentation of forest edge zones are increasing worldwide. Forest edges are characterized by altered plant species richness and community composition. As the tree layer and its species composition has been shown to influence herb layer composition, changes in tree species composition or richness may weaken or strengthen edge effects in forest ecosystems. We studied effects of the edge–center transition, tree species composition and their potential interaction on the understory vegetation in the Hainich National Park, Germany’s largest connected deciduous forest, allowing to cover large edge–center transects. Methods We established 12 transects in an area of 75 km 2 of continuous forest, 6 beech-dominated and 6 in multispecies forest stands. Each transect reached from the forest edge up to 500 m into the forest interior. Vegetation relevés were conducted in regular, logarithmic distances along each transect. Important Findings Herb species richness was influenced by an interaction of edge effects and tree diversity level. With increasing distance from the forest edge, herb species richness remained constant in multispecies forest stands but rapidly decreased in beech-dominated forest stands. Further, herb richness was higher in the interior of multispecies forest stands. Percent forest specialists increased and percent generalists decreased with distance from the edge and this contrasting pattern was much more pronounced in beech-dominated transects. By using structural equation modeling, we identified litter depth mediated by tree species composition as the most important driver of herb layer plant species richness.
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  • 57
    Publication Date: 2016-09-21
    Description: Aims Grazing and water availability are the primary drivers of vegetation dynamics in grazing-dominated regions of Mongolia with a semi-arid climate and frequent droughts. Nomadic animal husbandry still plays a large part in the economy of Mongolia, but more variable precipitation regime and increase in livestock number have severely affected grassland ecosystems through overgrazing, leading to pasture degradation. This study aimed to examine the effects of grazing exclusion, interannual variation of plant-available precipitation (PAP) and their interaction on the aboveground biomass (AGB) of each dominant species, the AGB of annual species and the total AGB in a Mongolian dry steppe, using long-term field data. Methods To detect the effect of grazing on vegetation dynamics, vegetation surveys were conducted in a non-grazed exclosure zone and a fully grazed area outside the exclosure. We assessed the effects of grazing, PAP and their interaction on AGB parameters using a generalized linear model. A detrended correspondence analysis (DCA) was used to visualize the effects of grazing and PAP on the AGB of each species. Important Findings Grazing, PAP and their interaction had significant effects on AGB. The effect of grazing on AGB was larger with higher precipitation and higher amounts of AGB (i.e. forage) while AGB was strongly limited in drought years, which resulted in a smaller grazing effect. The current year PAP had the highest impact ( r = 0.88, P 〈 0.01) on AGB. The dominance of annual species was characterized by the amount of PAP in the current and preceding years: annuals dominated in wet years that followed consecutive dry years. The DCA Axis 1 reflected the variation of AGB with interannual variation of PAP while the DCA Axis 2 differentiated the grazing effect. The DCA scatter diagram based on species score illustrated that Artemisia adamsii (an unpalatable herb) was clearly linked to grazing disturbance whereas palatable perennials such as Agropyron cristatum , Stipa krylovii and Cleistogenes squarrosa were related to grazing abandonment and wetter conditions. In brief, number of livestock, hence the grazing impacts on vegetation dynamic in this region could have driven by forage availability, which is mainly controlled by current-year PAP.
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  • 58
    Publication Date: 2016-09-21
    Description: Aims Foliar herbivory and water stress may affect floral traits attractive to pollinators. Plant genotypes may differ in their responses to the interplay between these factors, and evolution of phenotypic plasticity could be expected, particularly in heterogeneous environments. We aimed at evaluating the effects of simulated herbivory and experimental drought on floral traits attractive to pollinators in genetic families of the annual tarweed Madia sativa , which inhabits heterogeneous environments in terms of water availability, herbivore abundance and pollinator abundance. Methods In a greenhouse experiment with 15 inbred lines from a M . sativa population located in central Chile (Mediterranean-type climate), we measured the effects of apical bud damage and reduced water availability on: number of ray florets per flower head, length of ray florets, flower head diameter, number of open flower heads per plant, flowering plant height and flowering time. Important Findings Apical damage and water shortage reduced phenotypic expression of floral traits attractive to pollinators via additive and non-additive effects. Plants in low water showed decreased height and had fewer and shorter ray florets, and fewer and smaller flower heads. Damaged plants showed delayed flowering, were less tall, and showed shorter ray florets and smaller flower heads. The number of ray florets was reduced by damage only in the low water treatment. Plant height, flowering time and number of flower heads showed among-family variation. These traits also showed genetic variation for plasticity to water availability. Ray floret length, flower head size and time to flowering showed genetic variation for plastic responses to apical damage. Plasticity in flowering time may allow M. sativa to adjust to the increased aridity foreseen for its habitat. Because genetic variation for plastic responses was detected, conditions are given for evolutionary responses to selective forces acting on plastic traits. We suggest that the evolution of adaptive floral plasticity in M. sativa in this ecological scenario (heterogeneous environments) would result from selective forces that include not only pollinators but also resource availability and herbivore damage.
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  • 59
    Publication Date: 2016-09-21
    Description: Aims The two coexisting Leymus chinensis ecotypes exhibit remarkable divergences in adaptive strategies under drought and salinity in semi-humid meadows and semi-arid steppes. In order to detect the major genetic and environmental factors dominating the intraspecific phenotype variations and ecotype formation, the questions regarding the two distinct phenotypic forms (ecotypes) in L. chinensis were addressed: (i) did environments drive the L. chinensis ecotype formation? (ii) was there a molecular basis for the morphological divergence between the two ecotypes? (iii) which driving force dominated the intraspecies divergence, divergent natural selection, genetic drift or stabilizing selection? Methods We applied a series experiments on demographical, morphological and physiological traits of two Leymus chinensis ecotypes with gray green (GG) and yellow green (YG) leaf color in nine wild sites along a longitudinal gradient from 114° to 124°E in northeast China. The environmental data including mean annual precipitation, mean annual temperature, elevation and soil properties were collected. We compared the differences of morphological, physiological and genetic differentiations between the two ecotypes. Important Findings The GG type exhibited stronger fitness than YG type from the population densities, morphological traits (e.g. shoot height, leaf area, leaf and seed weights et al.), leaf mass per area (LMA) and physiological traits [relative water content (RWC), proline, soluble sugar contents]. Most of above phenotypes (e.g. total shoot densities, spike length et al.) were significantly correlated with mean annual precipitation, mean annual temperature and soil water content (SWC), rarely a correlated with soil pH and soil nutrient. Transplanted populations showed convergence trend by their leaf chlorophyll contents and osmotic adjustments (proline and soluble sugar contents) in the greenhouse, but still exhibited their divergences between two ecotypes in the outdoor transplantation, suggesting that whether L. chinensis ecotype differentiated could be largely affected by the environmental conditions. Furthermore, by the comparison result of quantitative genetic variation ( Q ST ) values from phenotypes with theoretical neutral genetic differentiation ( F ST ), differentiation in phenotypic traits greatly surpassed neutral predictions, implying that directional natural selection played a crucial role in L. chinensis ecotype differentiation. In addition, microsatellite analysis from Neighbor-joining tree and Bayesian assignment generated into two groups according to ecotypes, indicating molecular genetic differentiation also propelled the two ecotypes divergence. We conclude that L. chinensis population variations were driven by combing divergent natural selection (precipitation, temperature and SWCs) along the large-scale gradient and significantly intrinsic genetic differentiation.
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  • 60
    Publication Date: 2015-05-21
    Description: 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.
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  • 61
    Publication Date: 2015-05-21
    Description: 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 CO 2 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 CO 2 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.
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  • 62
    Publication Date: 2015-05-21
    Description: 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.
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  • 63
    Publication Date: 2015-05-21
    Description: 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.
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  • 64
    Publication Date: 2015-05-21
    Description: 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 .
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  • 65
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    Oxford University Press
    Publication Date: 2015-05-21
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  • 66
    Publication Date: 2015-05-21
    Description: 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.
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  • 67
    Publication Date: 2015-05-21
    Description: 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.
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  • 68
    Publication Date: 2015-05-21
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  • 69
    Publication Date: 2015-05-21
    Description: 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 x 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.
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  • 70
    Publication Date: 2015-05-21
    Description: Aims With a close association with plant water availability, foliar 13 C had been investigated extensively in alpine regions; however, foliar 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 13 C and 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 13 C and 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 13 C exhibited a significant positive linear relationship with altitude; in contrast, foliar 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 13 C, while differences in foliar 15 N were likely the result of soil N availability. (iii) Principal component analysis revealed a clear association between 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 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.
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  • 71
    Publication Date: 2015-01-23
    Description: Aims Measures of plot-to-plot phylogenetic dissimilarity and beta diversity are providing a powerful tool for understanding the complex ecological and evolutionary mechanisms that drive community assembly. Methods Here, we review the properties of some previously published dissimilarity measures that are based on minimum or average phylogenetic dissimilarity between species in different plots. Important Findings We first show that some of these measures violate the basic condition that for two identical plots the measures take the value zero. They also violate the condition that the dissimilarity between two identical plots should always be lower than that between two different plots. Such erratic behavior renders these measures unsuitable for measuring plot-to-plot phylogenetic dissimilarity. We next propose a new measure that satisfies these conditions, thus providing a more reasonable way for measuring phylogenetic dissimilarity.
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  • 72
    Publication Date: 2015-01-23
    Description: Aims To clarify whether variation in leaf traits with climate differs with scale, i.e. across species and within a species, and to detect whether plant functional group affects species-specific response. Methods Leaf dry matter content (LDMC), specific leaf area (SLA), mass- and area-based leaf N ( N mass , N area ) and leaf P concentrations ( P mass , P area ) and leaf chlorophyll concentration (SPAD) were measured for 92 woody plant species in two botanical gardens in China. The two gardens share plant species in common but differ in climate. Leaf trait variation between the two gardens was examined via mean comparison at three scales: all species together, species grouped into plant functional groups and within a species. A meta-analysis was performed to summarize the species-specific responses. Important Findings At the scale of all species together, LDMC, SLA, P mass and N mass were significantly lower in the dry-cold habitat than in the wet-warm one, whereas N area and SPAD showed an inverse pattern, indicating a significant environmental effect. The meta-analysis showed that the above-mentioned patterns persisted for SLA, N area and SPAD but not for the other variables at the species-specific scale, indicating that intraspecific variation affects the overall pattern of LDMC, P mass and N mass and P area . In terms of species-specific response, positive, negative or nonsignificant patterns were observed among the 92 species. Contrary to our prediction, species-specific responses within a functional group were not statistically more similar than those among functional groups. Our results indicated that leaf trait variation captured climatic difference yet species-specific responses were quite diverse irrespective of plant functional group, providing new insights for interpreting trait variability with climate.
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  • 73
    Publication Date: 2015-01-23
    Description: Aims Plant functional traits determine how plants respond to environmental factors and influence ecosystem processes. Among them, root traits and analyses of relations between above and below-ground traits in natural communities are scarce. Methods we characterized a set of above- and below-ground traits of three dominant shrub species in a semiarid shrub-steppe that had contrasting leaf phenological habits (deciduous, semideciduous and evergreen). We analysed if there was coordination among above- and below-ground resource economics patterns: i.e. patterns of biomass allocation, construction costs and lifespan. Important Findings Above- and below-ground traits and their resource economics relations pointed to species-specific functional strategies to cope with drought and poor soils and to a species ranking of fast to slow whole-plant strategies in terms of resource uptake, biomass construction costs and turnover. The deciduous shrub, Proustia cuneifolia , had relatively deep and even distribution of roots, and high proportion of short-lived tissues of low C construction costs: it had high fine to coarse root and high leaf-to-stem biomass ratios, high specific leaf area (SLA), and stems of low wood density. This strategy allows Proustia to maximize and coordinate above- and below-ground resources uptake as long as the most limiting factor (water) is available, but at the cost of having relative high plant biomass turnover. The evergreen Porlieria chilensis, instead, displayed a more conservative and slow strategy in terms of resource economics. It had ~80% of the roots in the 40cm topsoil profile, low proportion of fine compared with coarse roots and low leaf-to-stem ratios, low SLA and stems of high wood density, i.e. it invested in C costly tissues that, overall, persist longer but probably at the cost of having lower plant resource uptake rates. Traits in the semideciduous Adesmia bedwellii were in between these two functional extremes. Our results revealed high functional diversity and above- and below-ground complementarity in resource economics among these three codominant species in the Chilean coastal desert.
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  • 74
    Publication Date: 2015-01-23
    Description: Aims Soil CO 2 emission from steppes is affected by soil properties and vegetation in different successional stages. Primary and secondary succession of plants frequently occurred at the meadow steppe in Songnen Plain, Northeast China, which indicates the large uncertainty associated with CO 2 emission in this environment. This study aims to investigate the temporal variations of soil respiration (Rs) and the effect of plant succession on cumulative soil CO 2 emission during the growing season. Methods Using a LI-6400 soil CO 2 flux system, Rs of five vegetation types which represented different stages of plant succession in meadow steppes of Songnen Plain, China, was investigated during the growing seasons of 2011 and 2012. Important Findings Soil temperature (Ts) was the dominant controlling factor of Rs, which could explain ~64% of the change in CO 2 fluxes. The Q 10 values of Rs were ranged from 2.0 to 6.7, showing a decreasing trend with the plant successional stages. The cumulative CO 2 emission increased with the degree of vegetation succession and it averaged to 316±6g C m –2 (ranges: 74.8±6.7 to 516.5±11.4g C m –2 ) during the growing season. The magnitude of soil CO 2 emission during the growing season was positively correlated with aboveground plant biomass, soil organic carbon content and mean soil water content, while negatively linked to mean Ts, pH, electrical conductivity and exchangeable sodium percentages. The results implied that soil CO 2 emission increased with the development of plant communities toward more advanced stages. Our findings provided valuable information for understanding the variations of CO 2 emission in the process of vegetation succession.
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  • 75
    Publication Date: 2015-01-23
    Description: Aims Understanding the relationships among disturbance, invasion and species change is essential for effective management of many systems. We investigated relationships among fire history, invasion by a native tree species, Allocasuarina huegeliana , and diversity change to understand the potential drivers of plant community alteration in a complex and biodiverse system. Methods We used plant species surveys from 1983 and 2011 to quantify species loss/gain and thence compositional changes. Additionally, we surveyed population densities of the invasive species and collated long-term fire history data for each site. General linear models and non-parametric models were used to assess the strength of relationships between the three variables of interest. Important Findings Within the last 30 years, ~11% of the plant species richness was lost from the reserve. At an individual site level, we found only a 4% average decrease in overall plant species richness, but large species losses and gains that imply considerable compositional shifts. Though such shifts might be expected over 30 years, many of the gained species were common, potentially opportunistic species, while those lost were often locally rare woody perennials. In addition, gained species tended to be expanding their recorded range westward suggesting that they may be responding to the regional drying climate. The relationship between invasion density and species loss was strong over all spatial scales. We identified a potential state change to dominance by the native invasive particularly as high densities prevented species gain at the site scale. In these extreme cases of high invasive density and high biodiversity loss, we argue that there may be a need to directly address the expanding native population.
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  • 76
    Publication Date: 2015-01-23
    Description: Aims Soil heterogeneity is common in natural habitats. It may trigger foraging responses (placing more ramets and/or roots in nutrient-rich patches than in nutrient-poor patches) and further affect the growth of plants. However, the impact of soil heterogeneity on competitive interactions has been little tested. Methods We conducted a greenhouse experiment to investigate the effects of soil heterogeneity on intraspecific competition with a stoloniferous herb Hydrocotyle vulgaris . We grew one (without competition) or nine ramets (with competition) of H . vulgaris under a homogeneous environment and two heterogeneous environments differing in patch size (large or small patches). In the heterogeneous treatment, the soil consisted of the same number of nutrient-rich and nutrient-poor patches arranged in a chessboard manner, and in the homogeneous treatment, the soil was an even mixture of the same amount of the nutrient-rich and the nutrient-poor soil. Important Findings Irrespective of intraspecific competition, H. vulgaris showed foraging responses to soil heterogeneity in the large patch treatment, e.g. it produced significantly more biomass, ramets, aboveground mass and root mass in the nutrient-rich patches than in the nutrient-poor patches. In the small patch treatment, foraging responses were observed when intraspecific competition was present, but responses were not observed when there was no competition. However, we find a significant effect of soil heterogeneity on neither overall growth nor competitive intensity of H. vulgaris . Our results suggest that foraging responses to soil heterogeneity may not necessarily be adaptive and intraspecific competition may not be influenced by soil heterogeneity.
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  • 77
    Publication Date: 2015-01-23
    Description: Aims The Inclusive Niche Hypothesis has not been validated for plants using ecophysiological performance. The few experiments have measured growth and competition but not the physiological response of plants. A metallophyte plant that hyperaccumulates aluminium (Al), Plantago almogravensis , showed a defined spatial distribution by occurring mostly on vegetation gaps associated with metalliferous areas (geochemical islands). This case was used to determine, in situ , whether the Inclusive Niche Hypothesis was suitable to explain the extent of the species realized niche. Methods The vegetation associated with P. almogravensis geochemical islands in the SW coast of Portugal was mapped. The biotic (neighbouring plants) and abiotic (edaphic) components of the niche were correlated with parameters of the plant’s ecological and physiological performances (plant density and cover; leaf C and N concentration and isotopic composition; growth). The results were obtained using image analysis, abundance and morphological measures, isotopic signatures and chemical composition. Important Findings The species showed better physiological performance where its ecological performance was lower due to trade-offs with environmental constraints. The species’ realized niche was mostly limited by shrub competition and soil Al-toxicity. These limits contribute to explain the rarity status of the species: the species has a poor capacity to compete but, due to an enhanced Al-tolerance and Al-hyperaccumulator trait, has the ability to find refuge in geochemical islands that are too harsh for most other species. This work, based on ecophysiological field studies, provides support for the Inclusive Niche Hypothesis relating to plant species.
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  • 78
    Publication Date: 2015-01-23
    Description: Aims Grassland is the most widely distributed vegetation type on the Xizang Plateau. Accurate remote sensing estimation of the grassland aboveground biomass (AGB) in this region is influenced by the types of vegetation indexes (VIs) used, the grain size (resolution) of the remote sensing data and the targeted ecosystem features. This study attempts to answer the following questions: (i) Which VI can most accurately reflect the grassland AGB distribution on the Xizang Plateau? (ii) How does the grain size of remote sensing imagery affect AGB reflection? (iii) What is the spatial distribution pattern of the grassland AGB on the plateau and its relationship with the climate? Methods We investigated 90 sample sites and measured site-specific AGBs using the harvest method for three grassland types (alpine meadow, alpine steppe and desert steppe). For each sample site, four VIs, namely, Normalized Difference VI (NDVI), Enhanced VI, Normalized Difference Water Index (NDWI) and Modified Soil-Adjusted VI (MSAVI) were extracted from the Moderate Resolution Imaging Spectroradiometer (MODIS) products with grain sizes of 250 m and 1 km. Linear regression models were employed to identify the best estimator of the AGB for the entire grassland and the three individual grassland types. Paired Wilcoxon tests were applied to assess the grain size effect on the AGB estimation. General linear models were used to quantify the relationships between the spatial distribution of the grassland AGB and climatic factors. Important Findings The results showed that the best estimator for the entire grassland AGB on the Xizang Plateau was MSAVI at a 250 m grain size (MSAVI 250 m ). For each individual grassland type, the best estimator was MSAVI at a grain size of 250 m for alpine meadow, NDWI at a grain size of 1 km for alpine steppe and NDVI at a grain size of 1 km for desert steppe. The explanation ability of each VI for the grassland AGB did not significantly differ for the two grain sizes. Based on the best fit model (AGB = –10.80 + 139.13 MSAVI 250 m ), the spatial pattern of the grassland AGB on the plateau was characterized. The AGB varied from 1 to 136g m –2 . Approximately 59% of total spatial variation in the AGB for the entire grassland was explained by the combination of the mean annual precipitation (MAP) and mean annual temperature. The explanatory power of MAP was weaker for each individual grassland type than that for the entire grassland. This study illustrated the high efficiency of the VIs derived from MODIS data in the grassland AGB estimation on the Xizang Plateau due to the vegetation homogeneity within a 1 x 1 km pixel in this region. Furthermore, MAP is a primary driver on the spatial variation of AGB at a regional scale.
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  • 79
    Publication Date: 2015-01-23
    Description: Aims Arid environments are resource limited, with scarcity of water the key limiting factor, but hyper-arid environments are rarely studied. We test for spatial and temporal variation in ecologically important characteristics to deduce plant adaptations to the extreme climate. Methods The endangered Sinai Thyme ( Thymus decussatus ) exists as a set of patches on mountaintops within the St Katherine Protectorate, South Sinai, a hyper-arid environment with rare events of good rains (every 10–15 years). Important Findings From spatial and temporal patterns of plant mortality, size, condition and flowering among 10 patches on the Mt Sinai massif, we deduce that the incidence and amount of flowering responds relatively quickly (1–2 years) to rainfall fluctuations, but plant growth respond only very slowly. Small individuals are most at risk of death during drought, and a high proportion of plants were dead at the end of 8 years of very low or no rainfall. No recruitment of seedlings was observed even in years of good rainfall. Droughts are expected to become increasingly frequent due to climate change; this may have important consequences for Sinai Thyme and also its associated herbivores, such as the Critically Endangered Sinai Baton Blue ( Pseudophilotes sinaicus ) whose larval stage feeds exclusively upon the flowers of this plant.
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  • 80
    Publication Date: 2015-01-23
    Description: Aims The majority of angiosperms are pollinated by animals, and this interaction is of enormous importance in both agricultural and natural systems. Pollinator behavior is influenced by plants’ floral traits, and these traits may be modified by interactions with other community members. In recent years, knowledge of ecological linkages between above- and belowground organisms has grown tremendously. Soil communities are extremely diverse, and when their interactions with plants influence floral characteristics, they have the potential to alter pollinator attraction and visitation, but plant–pollinator interactions have been neglected in studies of the direct and indirect effects of soil organism–root interactions. Here, we review these belowground interactions, focusing on the effects of nitrogen-fixing bacteria, arbuscular mycorrhizal fungi and root-feeding herbivores, and their effects on floral traits and pollinators. Further, we identify gaps in our knowledge of these indirect effects and recommend promising directions and topics that should be addressed by future research. Important Findings Belowground organisms can influence a wide variety of floral traits that are important mediators of pollinator attraction, including the number and size of flowers and nectar and pollen production. Other traits that are known to influence pollinators in some plant species, such as floral volatiles, color and nectar composition, have rarely or never been examined in the context of belowground plant interactions. Despite clear effects on flowers, relatively few studies have measured pollinator responses to belowground interactions. When these indirect effects have been studied, both arbuscular mycorrhizal fungi and root herbivores were found to shift pollinator visitation patterns. Depending on the interaction, these changes may either increase or decrease pollinator attraction. Finally, we discuss future directions for ecological studies that will more fully integrate belowground ecology with pollination biology. We advocate a multilevel approach to these questions to not only document indirect effect pathways between soil interactions and pollination but also identify the mechanisms driving changes in pollinator impacts and the resultant effects on plant fitness. A more thorough understanding of these indirect interactions will advance ecological theory and may inform management strategies in agriculture and conservation biology.
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  • 81
    Publication Date: 2012-10-10
    Description: Aim Due to the important role of lianas in the functioning of forest ecosystem, knowledge of the factors that affect them are important in the management of forests. Currently, there are conflicting reports on the response of liana communities to disturbance, calling for more research in the area. The present study was carried out to investigate the response of liana diversity and structure to human disturbance within two major forests in the Penang National Park, Malaysia. The study also looked at the implication of the findings for conservation. Methods A total of 15 40 x 40-m 2 (or 40-m x 40-m) plots each were randomly located across a range of habitats in a primary forest and disturbed secondary forest. Trees with diameter at breast height ≥10 cm were examined for lianas with diameter ≥2 cm. Both lianas and trees were enumerated and compared between the two forests. Diversity and structural variables of lianas were compared between the two forests using the t -test analysis. Tree abundance was also compared between the two forests with t -test, while linear regression analysis was run to determine the effects of tree abundance on liana abundance. Important Findings A total of 46 liana species belonging to 27 genera and 15 families were identified in the study. Human disturbance significantly reduced liana species richness and species diversity in the secondary forest. Liana abundance remained the same in both forests whereas liana basal area was significantly higher in the primary forest. Twiners and hook climbers were significantly more abundant in the primary and secondary forest, respectively. Large diameter lianas were more abundant in the primary forest compared with the secondary forest. The diameter distribution of most families in the primary forest followed the inverted J-shaped curve whereas only a few of the families in the secondary forest did so. Tree abundance was significantly higher in the primary forest. The abundance of lianas significantly depended on tree abundance in all the forests. The study has provided evidence of negative effects of human disturbance on liana diversity and structure that does not auger well for biodiversity in the forest. In view of the critical role of lianas in maintaining biodiversity in the forest ecosystem, lianas in the national park should be protected from further exploitation.
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  • 82
    Publication Date: 2012-10-10
    Description: Aims The present study aims (i) to examine if recently reported interspecific shoot-level biomass allocational trade-offs, i.e. isometric trade-offs between leaf mass (LM) and stem mass (SM) and between leaf size and leaf number, hold intraspecifically and (ii) to explore whether those scaling relationships are independent of shoot type (i.e. long vs. short shoots). Methods In order to address our questions, we used Fagus sylvatica saplings growing under a broad light range that were sampled in the Western Carpathians Mountains (Slovakia). Important Findings We found that: (i) intraspecific shoot-level biomass allocational trade-offs differ from those reported interspecifically and that (ii) long and short shoots differ in biomass allocation scaling coefficients. Allometric relationships with slopes statistically smaller than 1.0 or higher than –1.0, were found between SM and LM and between mean leafing intensity and individual leaf mass, respectively, in long shoots. In contrast, isometric scaling was found in short shoots. This suggests that leaf mass in short shoots is unaffected by shoot stem mass, in contrast to long shoots. Short shoots also had a larger fraction of biomass allocated to leaves. Beech shoots, as has been observed in other shoot dimorphic species, are specialized, with short shoots specializing in carbon gain and long shoots in space acquisition. A greater shift in LM than in SM among species during speciation shifting from allometric intraspecific relationships to an isometric interspecific scaling relationship between those traits could explain the discrepancies between the outputs of the present intraspecific study and others similar studies. This study draws attention to the importance of considering shoot types in future studies dealing with allocation rules in species with dimorphic shoots.
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  • 83
    Publication Date: 2012-04-13
    Description: Aims To determine if an experimentally applied anomalous weather year could have effects on species composition and community structure that would carry over into the following year. Methods We conducted a field experiment applying two levels of temperature (ambient and +4°C) and two levels of precipitation (ambient and doubled) and followed cover of plant species during the treatment year and one post-treatment year. Data analysis included ordination analysis, examination of species frequency distributions and comparison of cover of functional groups and individual species. Important Findings A drought during the summer and fall of the treatment year resulted in significant differences in community structure between the 2 years. C 3 and winter annual species were depressed in the spring of the second year following the dry autumn. Species richness and legume cover increased in the second, wetter, year. Treatments caused no overall differences in community structure but did alter the dominance hierarchy of species among treatments as well as years. Warming decreased relative cover of winter annuals and early spring-flowering species but increased other annuals. Warming and double precipitation together increased cover of C 4 perennial graminoids. In particular, the warming and precipitation treatments both increased the abundance of Andropogon gerardii , not individually altering the dominance hierarchy but together nearly doubling the relative cover of A.gerardii , making it the most abundant species in the combined treatment, while the cover of Bromus arvensis , the former dominant, decreased by 25%. The following year, Andropogon relative cover increased further in the former warmed plots, becoming dominant in both the formerly warmed and warmed plus double precipitation treatments. The year following treatments also saw an increase in relative cover of summer-blooming species in the formerly warmed plots and differences among the former treatments in species richness of functional groups. If the effects of one anomalous year on plant abundance can carry over into the following year, several warm years could have a significant impact on plant community structure.
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  • 84
    Publication Date: 2012-04-13
    Description: Aims In recent years, coastal mangroves have been frequently affected by large disturbances (cyclones, hurricanes, flooding and tsunamis) and post-disturbance vegetation is often dominated by small stature mangrove, mangrove-associate and non-mangrove species potentially affecting ecosystem functioning. Knowledge on the processes of mangrove vegetation development and recovery (succession) following normal and large disturbances will benefit practitioners in designing robust ecosystem management/restoration plans. Here we propose a conceptual model of disturbance-mediated succession in mangroves. Methods Based on field observations and species’ life history traits, we develop conceptual models of mangrove succession under normal disturbance regime and recently experienced increased frequency of large disturbances. We evaluate our conceptual models by conducting a scenario testing experiment. Important Findings We suggest two predominant processes affecting mangrove succession after disturbance: propagule limitation due to damage of seed producing mature trees and dispersal barrier resulting from biological invasion associated with large disturbance. We argue that large disturbances affect mature trees more than the small-stature non-tree (shrubs, herbs and climbers) species creating a larger propagule shortage for mangrove tree species than non-tree species. Secondly, large disturbances facilitate invasion of free-floating aquatics, which may interfere with the flow-facilitated propagule dispersal and seedling establishment of mangrove species. In a scenario testing experiment, we have shown that similar levels of disturbance impact vegetation development and recovery differently depending on the presence or absence of invasive species. We conclude that since biological invasion is one of the major drivers of post-disturbance mangrove succession, the dimension of biological invasion should be included in prediction, management and restoration of mangrove forests.
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  • 85
    Publication Date: 2012-04-13
    Description: Aims In view of the growing interest in modelling the potential spread of invasive species, prediction of plant invasiveness on the basis of native range size holds considerable promise. Our objective was to use a simple model to evaluate whether a wider native range predisposes plant species to become invasive in non-native regions and to easily identify potential invaders on this basis. The Kashmir Himalayan alien flora, of which a large proportion is native to Europe, was used to test this model. Methods The Kashmir Himalayan alien flora comprises 436 species of vascular plants at different stages of invasion. We focussed on plant species at two critical invasion stages ( sensu Colautti and MacIsaac 2004 ), i.e. Stage II (species that are just at the earliest phase of introduction) and Stage V (species that are widespread and dominant in the invaded region and are thus considered invasive). We used the territorial distribution in Europe (number of countries) as a surrogate for the native range size of plants of European origin. Important Findings Using a subset of 88 species, for which information on the native European range was available, we showed that a large proportion (68%) of Stage II species growing in the Kashmir Valley had a relatively restricted European range (present in ≤20 countries); on the other hand, 77% of Stage V species had an extensive native range (present in 〉20 countries). We consequently hypothesized that 14 Kashmir Himalayan Stage II species of European origin that are distributed in 〉20 European countries are at risk of becoming future invaders in Kashmir. On the other hand, those Kashmir Himalayan Stage II species of European origin distributed in ≤20 European countries are less likely to become invasive. Although this analysis is quite simple, the data suggest that a wider native range is a good predictor of plant invasiveness and could be used as a simple and low-cost early warning tool in predicting potential invasive species.
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  • 86
    Publication Date: 2012-04-13
    Description: Aims Vegetation type is important in determining variations in soil carbon (C) efflux under grassland managements. This study was conducted to examine the effects of mowing and nitrogen (N) addition on soil respiration and their dependences upon vegetation types in an oldfield grassland of northern China. Methods Soil respiration, temperature, moisture and aboveground net primary productivity (ANPP) and belowground net primary productivity (BNPP) were examined in response to mowing and N addition among the three patches dominated by different species (named as grass, forb and mixed patches, respectively) in the growing seasons (May–October) from 2006 to 2008. Important Findings Across the 3 years, soil respiration in the grass patch was greater than those in the forb and mixed patches, which could have been ascribed to the higher soil moisture (SM) in the grass patch. Mowing had no impact on soil respiration due to unaltered SM and plant growth. Soil respiration was stimulated by 6.53% under N addition, and the enhancement was statistically significant in 2006 but not in 2007 or 2008 because of the limited water availability in the later 2 years. There were no interactive effects between mowing and N addition on soil respiration. Soil respiration showed positive dependence upon SM, ANPP and BNPP across plots. The results suggest that soil water availability and plant growth could be the primary factors in controlling the temporal and spatial variations in soil respiration and its response to different treatments. Our observations indicate that grassland managements (i.e. mowing for hay once a year) may have little influence on soil respiration of the oldfield grassland in northern China.
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  • 87
    Publication Date: 2012-04-13
    Description: Aims Numerous studies have showed that the balance between negative and positive plant–plant interactions shifted along environmental gradients. But little is known how the positive or negative plant–plant interactions varied with temporal fluctuating habitat conditions and plant ontogenetic phases. Methods In a 2-year experiment, the four perennial grasses ( Kobresia humilis , Stipa aliena , Elymus nutans and Saussurea superba ) were grown under four interaction treatments (no root or shoot interaction, only shoot interaction, only root interaction, root and shoot interaction). Intensity of above- and belowground interactions is proposed to vary with the fluctuation of seasonal climatic conditions and soil available nutrients. Here we report measurements of above- and belowground interactions during entire growing season. Correlation between plant interaction intensity and seasonal soil available N as well as habitat climate conditions was also performed. Important findings Our experiment found that root interactions had negative effect on plant growth for the four species during growing season. However, both negative and positive shoot interactions occurred among the four species. Despite there being shoot facilitative effect for E. nutans and S. superba , the full interaction was negative, suggested that root interaction take more important role on plant growth than that of shoot interaction. The interaction between root and shoot effect varied as a function of species identity and growth phases. The weak correlation of plant interaction intensity to habitat environmental factors suggested that plant ontogenetic characteristics may be primary factors causing temporal variation in plant interaction.
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  • 88
    Publication Date: 2012-04-13
    Description: Aims Ecosystem carbon models often require accurate net ecosystem exchange of CO 2 (NEE) light-response parameters, which can be derived from the Michaelis–Menten equation. These parameters include maximum net ecosystem exchange (NEE max ), apparent quantum use efficiency ( α ) and daytime ecosystem respiration rate ( R e ). However, little is known about the effects of land conversion between steppe and cropland on these parameters, especially in semi-arid regions. To understand how these parameters vary in responses to biotic and abiotic factors under land conversions, seasonal variation of light-response parameters were evaluated for a steppe and a cropland of Inner Mongolia, China, during three consecutive years (2006–08) with different precipitation amounts. Methods NEE was measured over a steppe and a cropland in Duolun, Inner Mongolia, China, using the eddy covariance technique, and NEE light-response parameters (NEE max , α and R e ) were derived using the Michaelis–Menten model. Biophysical regulations of these parameters were evaluated using a stepwise regression analysis. Important Findings The maximum absolute values of NEE max occurred in the meteorological regimes of 15°C ≤ T a 〈 25°C, vapor pressure deficit (VPD) 〈 1 KPa and 0.21 m 3 m – 3 ≤ volumetric soil water content at 10 cm (SWC) 〈 0.28 m 3 m – 3 for both the steppe and the cropland ecosystems. The variations of α and R e showed no regular variation pattern in different T air , VPD and SWC regimes. Under the same regime of T air , VPD and SWC, the cropland had higher absolute values of NEE max than the steppe. Canopy conductance and leaf area index (LAI) were dominant drivers for variations in NEE light-response parameters of the steppe and the cropland. The seasonal variation of NEE light-response parameters followed the variation of LAI for two ecosystems. The peak values of all light-response parameters for the steppe and the cropland occurred from July to August. The values of NEE light-response parameters (NEE max , α and R e ) were lower in the driest year (2007). Seasonally averaged NEE light-response parameters for the cropland surpassed those for the steppe. Land conversion from steppe to cropland enhanced NEE light-response parameters during the plant growing period. These results will have significant implications for improving the models on regional NEE variation under climate change and land-use change scenarios.
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  • 89
    Publication Date: 2012-04-13
    Description: Aims Road effects from maintenance and traffic have the potential to alter plant communities, but the exact relationships between these effects and changes in plant community composition have not often been studied in diverse environments. To determine the direction and level of community composition changes in saline environment due to road effects, we conducted a study along roads of different ages and in nearby non-road (i.e. natural) areas in the Yellow River Delta, China. Additionally, to potentially elucidate the mechanisms underlying the changes in the richness and composition of plant communities along roads, we evaluated physiochemical changes in soil of roadside and non-road areas. Methods Floristic and environmental data were collected along roadside of different ages and nearby non-road areas. To evaluate plant communities at each site, six 2 m x 2 m quadrats were placed at 3-m intervals along roads and six quadrats were arranged randomly in non-road areas. To determine the difference in plant community composition between roadside and non-road areas, we measured species richness and the abundance of each species, examined species turnover and floristic dissimilarity between the two areas and positioned plant species and sites in an abstract multivariate space. Plant community (species richness, percentage of halophytes) and soil physicochemical properties (pH, salinity, moisture content, bulk density, nitrate and ammonium nitrogen concentration) were compared between roadside and non-road areas (young roadside vs. corresponding non-road areas, old roadside vs. corresponding non-road areas) by using t -tests. Classification and ordination techniques were used to examine the relationship between vegetation and related environmental variables in both roadside and non-road areas. Important Findings For both the young and old roadside areas, species richness in roadside areas was significantly higher than in non-road areas and high floristic dissimilarity values indicated that roadside and non-road areas differed greatly in community composition. In both the young and old roadside areas, the plant communities in roadside areas had lower percentages of halophytes than non-road communities. Correspondence analysis and two-way indicator species analysis showed that halophytes dominated in the non-road areas, while a number of typical non-salt-tolerant species dominated in the roadside areas. Compared to non-road areas, activities associated with roads significantly decreased soil moisture, bulk density and salinity and increased soil pH and nitrate content. Forward selection for the environmental variables in canonical correspondence analysis showed that soil salinity was the most important factor related to the variation of species composition between roadside and non-road areas. Our study demonstrates that road effects have a significant impact on the associated vegetation and soil, and these changes are consistent across roads of different ages in our system.
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  • 90
    Publication Date: 2012-04-13
    Description: Aims Australian alpine ecosystems currently experience high precipitation in the snow-free season, but they are predicted to experience drier conditions under climate change. We observed high mortality of the dominant alpine grasses following drought in 2007. Our aims were as follows: to test the involvement of plant-available water (PAW) and other environmental variables in grass mortality in the field; to detect possible species differences in drought response and to link soil moisture to precipitation using soil properties and climate data. Methods The dominant tussock grasses of the Australian alpine zone, Poa hothamensis var. hothamensis N.G. Walsh, P oa hiemata Vickery and P oa phillipsiana Vickery (Poaceae), all exhibited mortality following drought in the Bogong High Plains, Victoria, Australia in 2007. PAW was calculated using soil water potential measurements, and past drought occurrence was modelled using climate data. We then tested the effects of PAW and soil depth on grass survival both at a large spatial scale spanning the elevational range of the alpine zone and at a smaller scale. Poa hothamensis and P. phillipsiana were compared in a common-garden experiment to test drought tolerance. Important Findings Poa hothamensis survival was predicted by dry-season PAW at the small spatial scale; at the large scale, soil depth and elevation were more important predictors of P. hothamensis survival, but dry-season PAW predicted P. hiemata survival. Common-garden experiments supported field observations that P. hothamensis is more drought-sensitive than is P. phillipsiana . We also present a simple polynomial relationship between rainfall and field soil moisture, which predicts that the alpine soils dry below wilting point several times a year. We suggest the timing of long rain-free periods may be more important than their duration.
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  • 91
    Publication Date: 2012-04-13
    Description: Aims Forest growth and recruitment and their relationships to climate are complex. The aims of our study are (i) to examine the patterns in tree radial growth and recruitment along an altitudinal gradient in Mt. Everest region and (ii) to identify the climatic factors that are responsible to the observed patterns in tree growth and recruitment. Methods Four plots, each 30 x 60 m in size, were established from the lower to upper limits of the eastern Himalayan fir forest in Dingjie County of the Mt. Everest Nature Reserve, China. Dendrochronological techniques were applied to obtain information about the radial growth and age of the trees in the plots. Correlation analysis was used to identify the relationships between radial growth and recruitment of trees and climatic variables, i.e. monthly mean temperature and Palmer Drought Severity Index (PDSI). The population age structure was analyzed to investigate the recruitment history. Important Findings The timberline plot was characterized by significant tree recruitments in the recent three decades and sporadic recruitments in earlier periods. The other three plots showed recruitment pulses during 1880–1910 in Plot 3 700 m, during 1870–80 and 1920–30 in Plot 3 520 m and during 1900–40 in Plot 3 410 m. The recruitment of fir trees in the timberline was sensitive to summer (June–September) temperatures, but it was mainly controlled by episodic disturbances in lower altitudes. Fir radial growth in the upper two plots was positively correlated with previous winter and current August temperature. Fir radial growth at the two lower plots was positively correlated with PDSI from previous September to current September.
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  • 92
    Publication Date: 2012-04-13
    Description: Aims Possible shifts in the phenotypic performance along invasive plants’ spreading route are rarely examined due to the discontinuous and incomplete records of exotic species. As the invasion history of common ragweed ( Ambrosia artemisiifolia L.) is well documented in Hungary, its residence time is known for each location. By sampling a sequence of older to more recently established populations, we aimed to determine (i) whether there are phenotypic divergences along the historical spreading route of A.artemisiifolia ; (ii) which traits are under selection during the invasion process and (iii) the extent of maternal effects on the individual's performance. Methods We used a hierarchical sampling design to collect seeds from 64 individuals belonging to eight sites in four residence time categories (seven populations along the historical spreading route of ragweed in Hungary and one recently invaded site in Romania). We selected four large and four small individual plants in each population to control for maternal effects. The offspring were reared in a common garden located in Romania. Five vegetative phenotypic traits were measured at the end of the experiments and used in the subsequent analysis (plant height, basal diameter, number of secondary axes, length of the longest secondary axis and biomass). To summarize the variation of these highly correlated traits, principal component analysis was performed first and then the important components were used in linear mixed effect models. Important Findings The residence time categories were significantly distinguished by the first component, which compresses the variation of all five measured traits. The measures gradually decrease from populations with the longest residence time (introduced more than 65 years ago) towards the most recently established populations (established less than 30 years ago). These differences might reflect the invasion history of the populations: the longer the residence time the higher the chance to develop relevant traits beneficial in invasion process. The size of the mother plant significantly influenced not only the seed mass (inversely) but also the adult performance of its offspring (directly).
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  • 93
    Publication Date: 2012-04-13
    Description: Aims Although stem CO 2 efflux is critical to ecosystem carbon and energy balance and its feedback to future climate change, little information is available on stem CO 2 efflux and its responses to temperature, especially in subtropical China. This study aims to (i) evaluate the temporal and spatial variations of stem CO 2 efflux of three species, including oak ( Quercus acutissima Carr.), masson pine ( Pinus massoniana Lamb.) and loblolly pine ( Pinus taeda Linn.) in subtropical China and (ii) analyze the temperature sensitivity of stem CO 2 efflux in the three species based on 2-year field measurements. Methods We measured stem CO 2 efflux and stem temperature (at 3 cm depth) of the three species using the horizontally oriented soil chamber technique from September 2008 to August 2010. We also conducted a 24-h measurement to examine the diurnal variation of stem CO 2 efflux in three consecutive days in April 2009. Important findings The temporal dynamics of stem CO 2 efflux followed the change of the stem temperature in a 3-cm depth with a bell-shaped curve in the three species. Stem temperature explained 77–85% of the seasonal variations of stem CO 2 efflux over the entire study period in the three species. The temperature sensitivity ( Q 10 ) of stem CO 2 efflux was obviously different among the three species with higher Q 10 value found in oak (2.24) and lower values in the coniferous species (1.76 and 1.63). Our results also showed that the Q 10 values of stem CO 2 efflux in all the three species were lower in the growing season than that in the non-growing season, indicating that the growth and maintenance respiration had different temperature responses. Moreover, we found that the temperature-normalized stem CO 2 efflux ( R 10 ) changed greatly between the growing and non-growing seasons in oak and masson pine, but not in loblolly pine. Additionally, we also found that in the non-growing season, the principal factor responsible for the spatial variation of stem CO 2 efflux among the 15 sampling trees was sapwood volume, whereas in the growing season, stem CO 2 efflux was closely related to annual dry-matter production in the three subtropical species.
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  • 94
    Publication Date: 2012-04-13
    Description: Aims Combining field data analysis and modeling, this study investigates factors influencing the diurnal boundary layer (BL) development in boreal forest. Methods Field data analysis used both air sounding and surface flux measurements collected during the Boreal Ecosystem–Atmosphere Study field campaigns in central Canada. Model study applied a non-local transilient turbulence theory (TTT) to simulate the impact of the heterogeneous boundary conditions together with initial conditions on the BL development at the Candle Lake and Thompson release sites over boreal forests. Boundary conditions were characterized by the integrated surface flux measurements from different forest stands. The lake effect was included in constructing the surface fluxes at Candle Lake release site. Important Findings Analyses of serial upper air sounding data and tower flux data indicate strong linear impacts of surface sensible heat forcing on the diurnal BL development above boreal forests. The regression slopes on the relationship between the BL development and the surface fluxes reflect the influences of initial boundary conditions to the BL developments. Both the modeled and the measured diurnal BLs show that lakes reduce sensible heat flux, leading to a shallower boundary in Candle Lake than in Thompson. Comparison of the model results and field measurements on the BL profiles indicates that the TTT model has the capability to simulate the BL development above boreal forests for sunny, rainfall or cloudy days. This study demonstrates the importance of lake on surface fluxes and BL development. The modeling effort shows the potential to couple the transilient theory with a land surface process model to study land surface and atmosphere interaction in boreal forest.
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  • 95
    Publication Date: 2015-09-17
    Description: Aims Understanding relationships between the distributions of species and their surrounding environment provides a basis for forecasting how species will respond to future environmental changes. In this study, we examined the effects of environmental factors and human developmental features associated with disturbances on probability of occurrence of juveniles of invasive Triadica sebifera and three native plant species, Baccharis halimifolia , Ilex vomitoria and Morella cerifera within a typical coastal transition in coastal Mississippi, USA. Methods We recorded presence of juveniles of focal species and measured environmental factors (soil salinity, canopy openness, soil texture and soil carbon to nitrogen ratio) along an 11.3 km transect located at Grand Bay National Estuarine Research Reserve. Further, we documented anthropogenic features and associated activities as a proxy for human disturbance. Important Findings With the exception of I. vomitoria , all other species occurrences were significantly influenced by an interaction between soil salinity and canopy openness. The occurrence of I. vomitoria sharply decreased with salinity. B. halimifolia occurrence peaked at moderate levels of salinity and low to moderate levels of canopy openness. Occurrences of M. cerifera and the invasive T. sebifera were highest at low levels of salinity and canopy openness. These results indicate that salinity is a strong driver that limits distribution of juvenile native and invasive species in coastal transitions. Logistic regression confirmed the positive effects of anthropogenic disturbances on T. sebifera and I. vomitoria occurrence. It appears that while soil salinity is likely to drive retreat of the seaward boundaries of woody plant species distributions, increased human developments along the coast likely enhance the inland spread of species, in particular the invasive T. sebifera by increasing disturbances and facilitating dispersal. Results from this study can offer insight for the development of T. sebifera management and preventive measures for further spread along coastal areas of the southeastern USA.
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  • 96
    Publication Date: 2015-09-17
    Description: Aims Understanding the effect of long-term fertilization on the sensitivity of grain yield to temperature changes is critical for accurately assessing the impact of global warming on crop production. In this study, we aim to assess the impacts of temperature changes on grain yields of winter wheat ( Triticum aestivum L.) under different fertilization treatments in a long-term manipulative experiment in North China. Methods We measured grain yields of winter wheat under four fertilization treatments at the Yucheng Comprehensive Experimental Station each year from 1993 to 2012. We also measured air temperature at 0200, 0800, 1400 and 2000h each day since 1 January 1980. We then used the first-difference method and simple linear regression models to examine the relationship of crop yield changes to mean air temperature, mean daytime and nighttime air temperature in crop growing seasons. Important Findings We found that increases in mean daily temperature, mean daytime temperature and mean nighttime temperature each had a positive impact on the grain yield of winter wheat. Grain yield increased by 16.7–85.6% for winter wheat in response to a 1°C increase in growing season mean daily temperature. Winter wheat yield was more sensitive to variations of nighttime temperature than to that of daytime temperature. The observed temperature impacts also varied across different fertilization treatments. Balanced fertilization significantly enhanced grain yields for winter wheat under a warming climate. Wheat plots treated with nitrogen and phosphorous balanced fertilization (NPK- and NP-treated plots) were more responsive to temperature changes than those without. This report provides direct evidence of how temperature change impacts grain yields under different fertilization treatments, which is useful for crop management in a changing global climate.
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  • 97
    Publication Date: 2015-09-17
    Description: Aims Woody invasions into grasslands have increased globally due to changing land use, climate and introduced woody species, but spatial processes generating and sustaining these invasions are not well understood. To gain insight into the patterns of spread of tree populations within grasslands, and to propose a full spatial analytical toolbox for studying native and non-native woody species spread when long-term data are not available, we tested if 50 years of grassland invasion in Western Carpathians by Norway spruce ( Picea abies Karst.) proceeded by one of the two traditionally competing hypotheses of species spread: (i) by frontier expansion, or (ii) by advanced groups established ahead of the population frontier. We also tested whether the pattern of invasion changed over time. Methods We analyzed the spatial demographic and genetic patterns of a Norway spruce population invading a Western Carpathian grassland using Ripley’s L ( t ) and genetic kinship coefficients ( F ij ). We mapped and genotyped spruce trees across the invasion front (from the invasion leading edge to fully colonized grassland near the source population) using three demographic classes (adults, juveniles and seedlings) to approximate the temporal aspects of the invasion. We studied how the spatial patterns of invasion by individual demographic classes and their genetic kinship varied among adjacent plots established at different distances from the source population (ranging from 0 to 160 m, in 40-m distance increments). Important Findings Juveniles were positively genetically related to adults on fine scales (〈4 m), suggesting that adults within the grassland acted as a seed source and accelerated early invasion. However, adults did not act as nucleation centers for the formation of advanced juvenile groups. Instead, genetically unrelated juveniles formed groups independently of adults. These groups were small and separate at the leading edge but they increased in size and graded into a continuous zone near the source population. Thus, juvenile recruitment occurred as a frontier expansion near the source population and as advanced groups controlled by environmental variation at the leading edge. Unlike juveniles, seedlings were clustered on all scales across the invasion front and formed groups around adult crowns at the invasion leading edge. The bulk of seedling establishment occurred at intermediate distances from the source population, independently from the adults, suggesting that the invasion front continued to expand as a frontier, gradually coalescing with the advanced groups at the leading edge. Thus, the grassland invasion was driven by a gradual frontier expansion of the original population during the first 50 years, with advanced groups enhancing but not driving the invasion process. Frontier expansion appeared more important as a mechanism of woody species spread early in the invasion process in this study, while advanced groups may play a larger role over longer temporal scales.
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  • 98
    Publication Date: 2015-09-17
    Description: Aims The relationship between biodiversity and ecosystem functioning has intrigued ecologists for several decades, but the effect of loss of a dominant species on community structure and functioning along a nutrient gradient remains poorly understood. The aim of this paper was to test the effect of a dominant species on community structure and function by conducting a species removal experiment along a fertilization gradient. Methods We removed the population of a dominant species ( Elymus nutans ) in a long-term fertilization field in an alpine meadow on the Tibetan Plateau, China. Univariate general linear models were used to evaluate the effects of fertilization and removal on above-ground vegetation characteristics, including photosynthetically active radiation in the understory, species richness, Shannon–Weiner diversity index, Simpson’s dominance index, above-ground biomass (including different functional groups) and seedling richness and density. Important Findings Results revealed that after two plant growing seasons, there was no significant effect of the removal of a dominant species on species richness and diversity of the remaining vegetation, but the biomass of forbs and seedling recruitment were significantly increased indicative of the potential for long-term effects. Moreover, removal had a large effect at high fertilization levels, but little effect when fertilization levels were low. Our studies indicated that community response to loss of a dominant species was mainly dependent on resource availability and the remaining functional group identities. We also found seedling recruitment was usually more sensitive to the influence of competition of dominant species than the established vegetation in the short term.
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  • 99
    Publication Date: 2015-09-17
    Description: Aim Lianas are abundant and diverse throughout the world and constitute an important structural and functional component of tropical forests. This study aims to investigate liana diversity, abundance and their functional traits in Indian tropical dry evergreen forest (TDEF). Methods A total of ten 1-ha plots, one each in 10 Indian TDEF sites were demarcated. Each 1-ha plot was divided into one-hundred 10- x 10-m quadrats to facilitate woody species inventory. All lianas ≥1cm diameter measured at 130cm from the rooting point and all trees ≥10-cm girth at breast height (gbh) were recorded from the study sites to analyze the patterns of liana diversity and abundance and also to compare the contribution of lianas to the total woody species richness, density and basal area. Liana variables across the study sites were compared using one-way analysis of variance. The qualitative functional traits of inventoried lianas and trees were assessed on the field and referring to pertinent field manuals. Important Findings A total of 9237 liana individuals (ranged from 408–1658 individuals ha –1 ) representing 52 species, 45 genera and 28 families were encountered from the 10 study sites. Liana species richness ranged from 11–31 species ha –1 in 10 sites, which averaged 23.4 (±5.7) species ha –1 . The total basal area of lianas in the study sites was 7. 3 m 2 (0.20–1.76 m 2 ha –1 ). There was a significant variation in liana species richness, density and basal area across the studied sites. On the whole, lianas contributed 52%, 49.3% and 4.1% to the total woody species (lianas and trees) richness, density and basal area, respectively. Liana trait analysis revealed the majority (50%) of lianas belonged to brevi-deciduous type. Stem twining was the chief climbing mechanism, exhibited by 21 species (52.6% of total abundance). More than half of the liana species (34 species; 6925 individuals) had microphyllous leaves. Fleshy-fruited lianas mostly bearing berries and drupes constituted the major fruit type in the studied sites. Zoochory was the predominant dispersal mode observed in 63.4% of species. Considering the ecological and functional role of lianas in Indian TDEF, the need for conservation is emphasized.
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  • 100
    Publication Date: 2015-09-17
    Description: Aims In grassland biodiversity experiments, positive biodiversity effects on primary productivity increase over time. Recent research has shown that differential selection in monoculture and mixed-species communities leads to the rapid emergence of monoculture and mixture types, adapted to their own biotic community. We used eight plant species selected for 8 years in such a biodiversity experiment to test if monoculture and mixture types differed in metabolic profiles using infrared spectroscopy. Methods Fourier transform infrared spectroscopy (FTIR) was used to assess metabolic fingerprints of leaf samples of 10 individuals of each species from either monocultures or mixtures. The FTIR spectra were analyzed using multivariate procedures to assess (i) whether individuals within species could be correctly assigned to monoculture or mixture history based on the spectra alone and (ii) which parts of the spectra drive the group assignment, i.e. which metabolic groups were subject to differential selection in monocultures vs. mixtures. Important Findings Plant individuals within each of the eight species could be classified as either from monoculture or mixture selection history based on their FTIR spectra. Different metabolic groups were differentially selected in the different species; some of them may be related to defense of pathogens accumulating more strongly in monocultures than in mixtures. The rapid selection of the monoculture and mixture types within the eight study species could have been due to a sorting-out process based on large initial genetic or epigenetic variation within the species.
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