Differences in Phenotypic Plasticity between Invasive and Native Plants Responding to Three Environmental Factors.

The phenotypic plasticity hypothesis suggests that exotic plants may have greater phenotypic plasticity than native plants. However, whether phenotypic changes vary according to different environmental factors has not been well studied. We conducted a multi-species greenhouse experiment to study the responses of six different phenotypic traits, namely height, leaf number, specific leaf area, total biomass, root mass fraction, and leaf mass fraction, of native and invasive species to nutrients, water, and light. Each treatment was divided into two levels: high and low. In the nutrient addition experiment, only the leaf mass fraction and root mass fraction of the plants supported the phenotypic plasticity hypothesis. Then, none of the six traits supported the phenotypic plasticity hypothesis in the water or light treatment experiments. The results show that, for different environmental factors and phenotypes, the phenotypic plasticity hypothesis of plant invasion is inconsistent. When using the phenotypic plasticity hypothesis to explain plant invasion, variations in environmental factors and phenotypes should be considered.

Contrasting Effects of Nitrogen Addition on Vegetative Phenology in Dry and Wet Years in a Temperate Steppe on the Mongolian Plateau.

Changes in spring and autumn phenology and thus growing season length (GSL) pose great challenges in accurately predicting terrestrial primary productivity. However, how spring and autumn phenology in response to land-use change and nitrogen deposition and underlying mechanisms remain unclear. This study was conducted to explore the GSL and its components [i.e., the beginning of growing season and ending of growing season (EGS)] in response to mowing and nitrogen addition in a temperate steppe on the Mongolia Plateau during 2 years with hydrologically contrasting condition [dry (2014) vs. wet (2015)]. Our results demonstrated that mowing advanced the BGS only by 3.83 days, while nitrogen addition advanced and delayed the BGS and EGS by 2.85 and 3.31 days, respectively, and thus prolonged the GSL by 6.16 days across the two growing seasons from 2014 to 2015. When analyzed by each year, nitrogen addition lengthened the GSL in the dry year (2014), whereas it shortened the GSL in the wet year (2015). Further analyses revealed that the contrasting impacts of nitrogen on the GSL were attributed to monthly precipitation regimes and plant growth rate indicated by the maximum of normalized difference vegetation index (NDVmax). Moreover, changes in the GSL and its two components had divergent impacts on community productivity. The findings highlight the critical role of precipitation regimes in regulating the responses of spring and autumn phenology to nutrient enrichment and suggest that the relationships of ecosystem productivity with spring and autumn phenology largely depend on interannual precipitation fluctuations under future increased nitrogen deposition scenarios.

Water table decline alters arthropod community structure by shifting plant communities and leaf nutrients in a Tibetan peatland.

Water table decline is one of the most serious environmental problems in the peatland in the Qinghai-Tibetan Plateau. However, the effect of water table decline on the structure of aboveground arthropod communities is still not clear. We investigated changes in the abundance of different arthropod groups, and estimated the abundance, height, and biomass of the plant community in a soil water table reduction experiment to reveal the effect of water table decline on the arthropod community structure. The effect of water level decline on herbivorous arthropods varied according to the feeding habits. Specifically, water table decline treatment decreased the abundance of grass-preferring herbivores but increased the abundance of forb-preferring herbivores. However, the density of predators (e.g., spiders) did not change significantly. The variations in arthropod communities were correlated with the increase in forbs and leaf nitrogen content in the water table decline treatments. Our experiment demonstrated that the effect of water table decline on plant communities cascades upwardly to alter the arthropod community. Such trophic interactions should be considered in studies aimed at predicting shifts in the arthropods communities in a changing climate.

Linkage between species traits and plant phenology in an alpine meadow.

Plant phenology differs largely among coexisting species within communities that share similar habitat conditions. However, the factors explaining such phenological diversity of plants have not been fully investigated. We hypothesize that species traits, including leaf mass per area (LMA), seed mass, stem tissue mass density (STD), maximum plant height (Hmax), and relative growth rate in height (RGRH), explain variation in plant phenology, and tested this hypothesis in an alpine meadow. Results showed that both LMA and STD were positively correlated with the onset (i.e., beginning) and offset (i.e., ending) times of the four life history events including two reproductive events (flowering and fruiting) and two vegetative events (leafing and senescing). In contrast, RGRH was negatively correlated with the four life phenological events. Moreover, Hmax was positively correlated with reproductive events but not with vegetative events. However, none of the eight phenological events was associated with seed size. In addition, the combination of LMA and STD accounted for 50% of the variation in plant phenologies. Phylogenetic generalized least squares analysis showed plant phylogeny weakened the relationships between species traits vs. phenologies. Phylogeny significantly regulated the variation in the ending but not the beginning of phenologies. Our results indicate that species traits are robust indicators for plant phenologies and can be used to explain the diversity of plant phenologies among co-occurring herbaceous species in grasslands. The findings highlight the important role of the combination of and trade-offs between functional traits in determing plant phenology diversity in the alpine meadow.

Spatiotemporal Variation of Osmanthus fragrans Phenology in China in Response to Climate Change From 1973 to 1996.

Climate change greatly affects spring and autumn plant phenology around the world consequently, and significantly impacts ecosystem function and the social economy. However, autumn plant phenology, especially autumn flowering phenology, has not been studied so far. In this study, we examined the spatiotemporal pattern of Osmanthus fragrans phenology, including both leaf phenology (the date of bud-bust, BBD; first leaf unfolding, FLD; and 50% of leaf unfolding, 50 LD) and flowering phenology (the date of first flowering, FFD; peak of flowering, PFD; and end of flowering, EFD). Stepwise multiple linear regressions were employed to analyze the relationships between phenophases and climatic factors in the long term phenological data collected by the Chinese Phenological Observation Network from 1973 to 1996. The results showed that spring leaf phenophases and autumn flowering phenophases were strongly affected by latitude. BBD, FLD, and 50LD of O. fragrans were delayed by 3.98, 3.93, and 4.40 days as per degree of latitude increased, while FFD, PFD and EFD in O. fragrans advanced 3.11, 3.26, and 2.99 days, respectively. During the entire study period, BBD was significantly delayed across the region, whereas no significant trends were observed either in FLD or 50LD. Notably, all flowering phenophases of O. fragrans were delayed. Both leaf and flowering phenophases negatively correlated with growing degree-days (GDD) and cold degree-days (CDD), respectively. BBD and FLD were negatively correlated with total annual precipitation. In addition to the effects of climate on autumn flowering phenology, we found that earlier spring leaf phenophases led to delayed autumn flowering phenophases. Our results suggest that future climate change and global warming might delay the phenological sequence of O. fragrans. Our findings also advanced the flowering mechanism study of autumn flowering plants, and facilitated the accurate prediction of future phenology and climate change.

Short-term phosphorus addition augments the effects of nitrogen addition on soil respiration in a typical steppe.

Soil respiration is one of the largest carbon (C) sources in terrestrial ecosystems and is sensitive to soil nutrient variation. Although nitrogen (N) availability affects soil respiration, other nutrients, such as phosphorous (P), which play pivotal roles in plant growth and microbial activity, may also affect soil respiration. In addition, N and P have been widely reported to interactively affect plant growth; however, their interactive effects on soil respiration have rarely been studied. Therefore, we conducted a short-term, two-factor experiment (from 2013 to 2015) to determine whether N and P addition can interactively affect soil respiration in a northern Chinese steppe. Nitrogen addition elevated soil respiration by 9.5%, whereas P addition did not affect soil respiration in the studied steppe across all treatments. However, neither N nor P addition significantly affected soil respiration alone in the experiment. Furthermore, N and P interactively affected soil respiration. Nitrogen addition did not affect soil respiration in the ambient P plots, but significantly elevated soil respiration (by 17.7%) in P addition plots across the three growing seasons. The effects of N addition on soil respiration were primarily correlated with the responses of vegetation cover and litter biomass to N addition in the experiment. Our results demonstrate that P addition augments the effects of N addition on soil respiration. Soil nutrient contents should be incorporated into predictive models for terrestrial C cycle response to N addition.

Direct effects of nitrogen addition on seed germination of eight semi-arid grassland species.

Seed germination plays an important role in mediating plant species composition of grassland communities under nitrogen (N) enrichment. Shifts of plant community structure with N-enhanced deposition in terrestrial ecosystems have occurred globally. Despite numerous studies about the effects of enhanced N deposition on mature plant communities, few studies have focused on seed germination. Using a laboratory experiment, we report the effects of five N concentrations, including 0, 5, 10, 20, and 40 mM N (NH4NO3) on seed germination of eight semi-arid grassland species. Results showed that low N concentrations (5- and 20-mM N) promoted mean final germination proportion of all eight species by 4.4% and 6.4%, but high concentrations (40 mM N) had no effect. The mean germination rate was decreased 2.1% and 5.1% by higher N concentration (20- and 40-mM N) levels, but germination start time showed the opposite trend, delayed by 0.7, 0.9, and 1.8 d for the 10, 20, and 40 mM N treatments. Final germination proportion, mean germination rate, and germination start time were significantly different among species in response to N concentration treatments. The final germination proportion of Allium tenuissimum and Chenopodium glaucum were suppressed by increased N concentration, whereas it increased for Potentilla bifurca, Plantago asiatica, and Setaria viridis. Our findings provide novel insights into N deposition-induced species loss based on seed germination factors in semi-arid grassland communities.

A meta-analysis of 1,119 manipulative experiments on terrestrial carbon-cycling responses to global change.

Direct quantification of terrestrial biosphere responses to global change is crucial for projections of future climate change in Earth system models. Here, we synthesized ecosystem carbon-cycling data from 1,119 experiments performed over the past four decades concerning changes in temperature, precipitation, CO2 and nitrogen across major terrestrial vegetation types of the world. Most experiments manipulated single rather than multiple global change drivers in temperate ecosystems of the USA, Europe and China. The magnitudes of warming and elevated CO2 treatments were consistent with the ranges of future projections, whereas those of precipitation changes and nitrogen inputs often exceeded the projected ranges. Increases in global change drivers consistently accelerated, but decreased precipitation slowed down carbon-cycle processes. Nonlinear (including synergistic and antagonistic) effects among global change drivers were rare. Belowground carbon allocation responded negatively to increased precipitation and nitrogen addition and positively to decreased precipitation and elevated CO2. The sensitivities of carbon variables to multiple global change drivers depended on the background climate and ecosystem condition, suggesting that Earth system models should be evaluated using site-specific conditions for best uses of this large dataset. Together, this synthesis underscores an urgent need to explore the interactions among multiple global change drivers in underrepresented regions such as semi-arid ecosystems, forests in the tropics and subtropics, and Arctic tundra when forecasting future terrestrial carbon-climate feedback.

Elevated CO2 does not stimulate carbon sink in a semi-arid grassland.

Elevated CO2 is widely accepted to enhance terrestrial carbon sink, especially in arid and semi-arid regions. However, great uncertainties exist for the CO2 fertilisation effects, particularly when its interactions with other global change factors are considered. A four-factor (CO2 , temperature, precipitation and nitrogen) experiment revealed that elevated CO2 did not affect either gross ecosystem productivity or ecosystem respiration, and consequently resulted in no changes of net ecosystem productivity in a semi-arid grassland despite whether temperature, precipitation and nitrogen were elevated or not. The observations could be primarily attributable to the offset of ecosystem carbon uptake by enhanced soil carbon release under CO2 enrichment. Our findings indicate that arid and semi-arid ecosystems may not be sensitive to CO2 enrichment as previously expected and highlight the urgent need to incorporate this mechanism into most IPCC carbon-cycle models for convincing projection of terrestrial carbon sink and its feedback to climate change.

Fire rather than nitrogen addition affects understory plant communities in the short term in a coniferous-broadleaf mixed forest.

Increasing fire risk and atmospheric nitrogen (N) deposition have the potential to alter plant community structure and composition, with consequent impacts on biodiversity and ecosystem functioning. This study was conducted to examine short-term responses of understory plant community to burning and N addition in a coniferous-broadleaved mixed forest of the subtropical-temperate transition zone in Central China. The experiment used a pair-nested design, with four treatments (control, burning, N addition, and burning plus N addition) and five replicates. Species richness, cover, and density of woody and herbaceous plants were monitored for 3 years after a low-severity fire in the spring of 2014. Burning, but not N addition, significantly stimulated the cover (+15.2%, absolute change) and density (+62.8%) of woody species as well as herb richness (+1.2 species/m2, absolute change), cover (+25.5%, absolute change), and density (+602.4%) across the seven sampling dates from June 2014 to October 2016. Light availability, soil temperature, and prefire community composition could be primarily responsible for the understory community recovery after the low-severity fire. The observations suggest that light availability and soil temperature are more important than nutrients in structuring understory plant community in the mixed forest of the subtropical-temperate transition zone in Central China. Legacy woody and herb species dominated the understory vegetation over the 3 years after fire, indicating strong resistance and resilience of forest understory plant community and biodiversity to abrupt environmental perturbation.

Effects of nitrogen addition and mowing on rodent damage in an Inner Mongolian steppe.

Rodent damage is a serious threat to sustainable management of grassland. The effects of nitrogen (N) deposition and grassland management on rodent damage have been scarcely studied. Here, we reported the effects of 2 years of N addition and mowing on burrow density and damage area of Citellus dauricus in a semiarid steppe in Inner Mongolia. N addition significantly aggravated, while mowing alleviated rodent damage in the grassland under study. Burrow density and damage area increased 2.8-fold and 4.7-fold, in N addition plots compared to the ambient N addition treatment, respectively. Conversely, mowing decreased burrow density and damage area by 75.9% and 14.5%, respectively, compared to no mowing plots. Observed changes in rodent damage were mainly due to variations in plant community cover, height, and aboveground net primary productivity. Our findings demonstrate that N addition and mowing can affect the rodent density and activity in grassland, suggesting that the effects of a changing atmospheric composition and land use on rodent damage must be considered in order to achieve better grassland management.

Global warming reduces plant reproductive output for temperate multi-inflorescence species on the Tibetan plateau.

• Temperature is projected to increase more during the winter than during the summer in cold regions. The effects of winter warming on reproductive effort have not been examined for temperate plant species. • Here, we report the results of experimentally induced seasonal winter warming (0.4 and 2.4°C increases in growing and nongrowing seasons, respectively, using warmed and ambient open-top chambers in a Tibetan Plateau alpine meadow) for nine indeterminate-growing species producing multiple (single-flowered or multi-flowered) inflorescences and three determinate-growing species producing single inflorescences after a 3-yr period of warming. • Warming reduced significantly flower number and seed production per plant for all nine multi-inflorescence species, but not for the three single-inflorescence species. Warming had an insignificant effect on the fruit to flower number ratio, seed size and seed number per fruit among species. The reduction in seed production was largely attributable to the decline in flower number per plant. The flowering onset time was unaffected for nine of the 12 species. Therefore, the decline in flower production and seed production in response to winter warming probably reflects a physiological response (e.g. metabolic changes associated with flower production). • Collectively, the data indicate that global warming may reduce flower and seed production for temperate herbaceous species and will probably have a differential effect on single- vs multi-inflorescence species.

Shifting phenology and abundance under experimental warming alters trophic relationships and plant reproductive capacity.

Phenological mismatches due to climate change may have important ecological consequences. In a three-year study, phenological shifts due to experimental warming markedly altered trophic relationships between plants and insect herbivores, causing a dramatic decline of reproductive capacity for one of the plant species. In a Tibetan meadow, the gentian (Gentiana formosa) typically flowers after the peak larva density of a noctuid moth (Melanchra pisi) that primarily feeds on a dominant forb (anemone, Anemone trullifolia var. linearis). However, artificial warming of approximately 1.5 degrees C advanced gentian flower phenology and anemone vegetative phenology by a week, but delayed moth larvae emergence by two weeks. The warming increased larval density 10-fold, but decreased anemone density by 30%. The phenological and density shifts under warmed conditions resulted in the insect larvae feeding substantially on the gentian flowers and ovules; there was approximately 100-fold more damage in warmed than in unwarmed chambers. This radically increased trophic connection reduced gentian plant reproduction and likely contributed to its reduced abundance in the warmed chambers.