Anthropogenic phosphorus (P) inputs are rapidly altering terrestrial P cycling through plant - soil - microbial interactions; however, global patterns and underlying mechanisms driving these changes remain poorly understood. By performing a global meta-analysis of 1315 observations from 176 studies across diverse natural terrestrial ecosystems, we found that P addition increased P concentrations in foliage, stems, roots, and litter by 62%, 114%, 100% and 63%, respectively. Soil total P, plant-available P, and microbial P concentrations rose by 43%, 221%, and 70%, while leaf P-resorption efficiency and soil phosphatase activity declined by 23% and 15%, respectively. Stem P and soil phosphatase activity exhibited consistent trends across tropical, temperate, and boreal zones, suggesting climate-specific P acquisition strategies. In addition, foliar P responses diverged among ecosystem and plant functional types. These responses were primarily regulated by background soil total P concentration, precipitation, soil pH, and P addition duration and rate. Our findings provide critical insights into the potential consequences of increasing anthropogenic P inputs in natural terrestrial ecosystems, improving our understanding of nutrient cycling and informing future ecosystem management under ongoing global change.
Leaf phosphorus (P) concentration has traditionally been assumed to increase from the equator to the poles. However, whether there exists a uniform or symmetrical latitudinal pattern across both hemispheres has never been examined due to variation in the geological histories, land-sea distribution, and climate of the hemispheres. We analyzed global latitudinal trends in woody plant leaf P concentrations across hemispheres to evaluate four hypotheses underlying these patterns. We show that leaf P concentration was significantly higher in the Northern Hemisphere than in the Southern Hemisphere, increasing with latitude in the former but decreasing in the latter. Key drivers of leaf P concentration differed between hemispheres: temperature dominated Northern Hemisphere variations, supporting the Temperature-Plant Physiology Hypothesis, while soil available P (indicative of substrate age) primarily influenced Southern Hemisphere trends, supporting the Soil Substrate Age Hypothesis. Temperature and precipitation play opposite roles in forming the leaf P latitudinal patterns in the two hemispheres. Our findings challenge the notion of a traditional latitudinal P gradient and emphasize incorporating hemisphere-specific climatic and edaphic drivers into forest productivity and global P-cycling models. Resolving these asymmetries will improve predictions of vegetation responses to environmental change and enhance Earth system model accuracy.
Abstract Caragana microphylla is a primary species that drives shrub encroachment across the Eurasian steppe. Understanding how its resource allocation strategies shift during ontogeny is crucial for understanding its expansion mechanism. Here, we quantified organ-level biomass allocation and the stoichiometric characteristics of carbon (C), nitrogen (N), and phosphorus (P) along an individual size gradient. We sampled 150 shrubs on the shrub-encroached grassland of Inner Mongolia, and classified them into three size classes. For each individual, we measured the biomass and C, N, and P contents of five organs—leaves, current-year twigs, perennial branches, fine roots, and coarse roots. Biomass allocation patterns were analyzed using allometric equations, and resource strategies were evaluated using principal component analysis. Our results revealed that, as shrub size increased, only the coarse-root mass fraction increased, whereas allocation to aboveground organs and fine roots decreased or stabilized. Allometric analyses revealed that underground organs scaled positively with total biomass (α > 1), whereas aboveground organs approached isometry at larger sizes. These patterns indicate that a trade-off occurs between organs across size classes, representing a shift from photosynthetic and absorptive tissues toward structural and hydraulic consolidation as shrubs grow larger. Compared with inactive organs (perennial branches and coarse roots), active organs (leaves, current-year twigs and fine roots) maintained greater amounts of N and P. Overall, as individuals grow larger, C. microphylla shifts from a “conservative” to an “acquisitive” strategy. This ontogenetic shift provides a new physiological perspective on the life history strategies facilitating shrub encroachment in arid environments.
The growing demand for ruminant livestock products in China has intensified the conflict with grassland grazing pressure. The intensification of cultivated forage production is increasingly recognized as a key strategy to support increasing livestock feed demand and to reduce grazing pressure. This study developed a novel approach to estimate potential yields of three major cultivated forages, including alfalfa, silage maize, and forage oats, under data-limited conditions, using data sourced from 713 papers published between 1985 and 2023, covering 11,822 entries on forage yields across China. By combining K-means clustering with high-yield sample selection and random forest modeling, we mapped potential yields of alfalfa, silage maize, and forage oats across China. Results show that improving yields without expanding sown area could provide an additional 35.6 million tonnes (Mt) of forage nationwide, equivalent to 48.3% of their current production of these forages. This yield increment corresponds to an additional 171.0 PJ of net energy supply, sufficient to meet 10.9% of the energy requirements of beef cattle, dairy cattle, sheep, and goats in 2022. Consequently, 36–41% (46.7–52.3 million hectares) of currently utilized grasslands could be spared for ecological protection if the potential forage yield predicted in this study can be achieved. These findings highlight the role of cultivated forage in securing feed supply, alleviating grassland degradation, and supporting national ecological restoration, while also supporting the “small land for large protection” principle, which advocates using modest areas of cultivated land to safeguard natural grasslands.
Tree growth rates generally increase with individual size, but how nutrient enrichment alters the size-growth relationship remains poorly understood. In this study, we asked whether a decade of nitrogen (N) addition altered the size-growth relationship of trees, and explored the potential mechanisms underlying such changes. We found a nonlinear shift in the size-growth relationship along the N addition gradient. The near size-symmetric growth observed in control plots shifted to a pattern favoring smaller individuals under low-level N addition (20 kg N ha(-1) yr(-1)). This occurred because small trees had higher leaf N resorption efficiency in controls, reflecting stronger N conservation and allowing a stronger growth response to moderate N inputs. With further increases in N addition (> 50 kg N ha(-1) yr(-1)), the relationship reversed toward size symmetry. This reversal was associated with N-induced changes in nutrient stoichiometry, as smaller trees showed greater phosphorus and potassium resorption efficiencies, suggesting they experienced stronger secondary limitation by these nutrients and consequently lost their growth advantage relative to larger trees. Our findings provide novel insights into how N addition rate reshapes the size-growth relationship of trees, highlighting multiple nutrient limitation as a potential driver of stand development. These results suggest that size-dependent growth responses should be incorporated into forest management and projections of long-term biomass accumulation under increasing N deposition.
Forest line is one of the hot spots in ecological research. It is well documented that the dynamics of forest line under climatic warming, yet how soil microbial community composition shifts across forest line ecotone remains unclear. This study investigated the change in microbial communities of rhizosphere and bulk soils at locations above, at, and below the forest line within an alpine forest-shrubland ecotone in the Hengduan Mountains, China. The results showed that the relative abundances of dominant phyla in surface rhizosphere soil changed more significantly along the forest line ecotone, with Proteobacteria and Basidiomycota decreasing but Chloroflexi and Mortierellomycota increasing, compared to bulk soil. Both the network complexity and key nodes of soil microbial communities changed significantly along the forest line ecotone. Specifically, bacterial network complexity between rhizosphere and bulk soils depended on soil depth, whereas fungal network complexity between rhizosphere and bulk soils varied with forest line. Microbial community compositions were co-regulated by both environmental and vegetation factors. Furthermore, microbial community changes were primarily driven by the cover and height of shrub and herb layers, highlighting the importance of shrubs and herbs in shaping soil microbial community composition. Our findings reveal the distribution patterns and underlying shift mechanisms of microbial communities along the forest line ecotone. These insights are crucial for mountain biodiversity conservation and for improving predictions of forest responses to climate change.
Trophic transfer efficiency (TTE) describes the proportion of energy or nutrients transferred from one trophic level to the next. A common assumption holds that ~10% of energy is transferred upward. This rule of thumb has shaped ecological models and discussions of sustainability, but broad empirical tests are lacking. We compiled 2052 TTE estimates from 122 studies across ecosystems. Average energy transfer efficiency (TTEe) was 5.92%, well below 10%, while nutrient transfer efficiency (TTEn) averaged 11.13%. Marine ecosystems had the highest TTEe (8.13%), followed by freshwater (5.53%) and terrestrial (1.52%) systems. TTEe declined with temperature in freshwater and was lower for consumers feeding on autotrophs, endotherms, and higher trophic levels. Our findings challenge long-standing assumptions and highlight the need for better understanding of TTE variation.
Between 2023 and 2024, Amazonian rainforests experienced two consecutive, record-breaking droughts-each more intense than any previously observed-yet their impacts remain largely unquantified. Using newly developed monthly radar satellite observations (1992 to 2025) that track forest moisture and biomass dynamics, we analyzed the long-term responses of intact Amazonian rainforests to past major droughts-particularly the 2023-2024 event-and projected their post-drought recovery. We found a biome-wide sharp decline in radar signal during 2023-2024, marking the lowest level observed since 1992. Spatially, 26.8% of the forests reached their three-decade minima during this period, primarily in eastern Amazonia. This ratio is more than double that recorded during the 2005 drought, when 11.0% of the forests reached such minima. Moreover, projections based on both historical and future CMIP6 precipitation scenarios consistently indicated that, even 7 y after the 2023-2024 droughts, less than 50% of the affected areas are expected to recover to predrought conditions, and these forests are associated with lower soil cation concentrations, higher soil sand content, and lower canopy height-characteristics that lessen the risk of hydraulic failure. Given that severe droughts have occurred approximately every 7 y over the past three decades, Amazonian rainforests may face another drought before fully recovering from the 2023-2024 event. Our results therefore highlight the growing vulnerability of the Amazonian rainforests to intensifying climate extremes driven by El Niño events and ongoing anthropogenic climate change, providing evidence that these forests are approaching the limits of their preindustrial operating space.
Leaf traits influence biotic interactions and ecosystem functions in forests. However, biogeographic drivers of leaf traits remain highly uncertain, limiting their integration into global vegetation models. Using a global dataset of forest plots, we show that community canopy leaf traits align along three dimensions: leaf economy, leaf density, and nitrogen-to-phosphorus (N:P) ratio. Changes in these trait dimensions across forest communities were driven by different variables: leaf economy was primarily shaped by the proportion of deciduous trees, leaf density by soil available P and temperature, and leaf N:P ratio by temperature. These three leaf trait dimensions together explained 60.2% of the variation in ecosystem-scale forest maximum photosynthesis, with the leaf N:P ratio showing the strongest association, followed by the dimensions of leaf economy and leaf density. Forests with moderate leaf N:P ratios and more acquisitive traits had higher photosynthesis than other forests. Our findings highlight the potential of community canopy leaf traits in predicting biogeographic variation in ecosystem-scale forest functioning.
Context: Filling China's massive forage gap requires intensified silage maize (Zea mays L.) production with high and stable yield. Modifying irrigation and sowing time comprise two practical, cost-effective measures for such intensification, yet their spatial effects remain insufficiently quantified. Methods: Using the APSIM Classic 7.10 version for Maize, calibrated with experimental data using a differential evolution algorithm, we assessed biomass yield potential, irrigation benefits, and sowing strategies at 10 km resolution across five regions in China (1980-2017). In addition, partial correlation analysis was employed to identify the climatic drivers of irrigation benefits, while sensitivity analysis was conducted to evaluate the effects of sowing date and climate variability on biomass yield. Results: Under full irrigation and non-nitrogen-limiting conditions, potential yield (Yp) exceeded 24 Mg/ha in Northeast (NE) and Northwest (NW) China, while rainfed potential (Yw) was highest in Southwest (SW; 20 Mg/ ha) and Southeast (SE; 18 Mg/ha). Irrigation expanded high-yield stable (HS) areas from low-yield unstable (LU) zones by 26%, particularly in arid and semi-arid northern zones, with impacts of irrigation modulated by mean annual precipitation (MAP) during growing season. Optimal sowing strategies varied by region and water regime: early sowing by 30 days most improved HS areas in North China Plain (NC; +49%), SE (+28%), and SW (+26%) under sufficient water, while NE and NW favored 20-day and 10-day advances, respectively. Under rainfed conditions, southern regions benefited from early sowing, whereas the north required moderate delays for yield stability. Sensitivity analysis further confirmed that sowing date dominated irrigated yield variability in the north, while precipitation and radiation were key limiting factors in rainfed northern (MAP < 650 mm) and southern (MAP > 1000 mm) zones, respectively. Conclusions: This study provides the first high-resolution national assessment of silage maize, offering a scientific basis for tailoring its deployment to regional climate constraints and water availability, thereby supporting resilient forage intensification under changing environmental conditions.
Global change ecology demands predictive models that reconcile data-driven learning with mechanistic theory to address complex, interconnected ecosystem challenges. Traditional process-based approaches struggle with spatiotemporal parameterization, while purely data-driven machine learning approaches suffer from extrapolation, interpretability, and physical consistency. Knowledge-guided machine learning (KGML) bridges this divide by systematically integrating ecological principles (e.g., physical first principles, stoichiometry, process understanding, disturbance regimes) into how models are designed, trained, and adjusted to generalize across different ecosystems. The emerging KGML paradigm offers tremendous opportunities to advance the research of global change ecology. This review synthesizes KGML's transformative potential, showcasing its capacity to enhance the prediction of carbon-water-nutrient cycles and other ecological processes and lay groundwork for ecological foundation models. Emerging applications in decision support and symbolic regression further illustrate its role in deriving actionable insights and novel theoretical hypotheses. Future directions emphasize adaptive integration of data and knowledge, uncertainty quantification, causal embedding in foundation models, and interdisciplinary collaboration to align KGML innovations with sustainability goals. By uniting ecological theory with AI advances, KGML offers a robust pathway to encompass ecosystem responses to global change, fostering scientific discovery and actionable solutions.
Water availability strongly constrains terrestrial plant growth and distribution, with water-use efficiency (WUE) reflecting plant adaptive strategies. Leaf pH is an important but poorly explored functional trait whose ecological significance remains unclear, although its geographical pattern appears closely linked to water conditions. We compiled datasets of leaf pH and leaf delta 13C across China to examine how leaf pH is related to water-use efficiency at different scales and to identify the climatic drivers underlying their spatial variation. We found that leaf pH was positively correlated with intrinsic water-use efficiency (iWUE) at species, functional group and geographical levels, with precipitation being the dominant factor shaping their geographic distributions. Leaf pH and iWUE were tightly coupled geographically, exhibiting consistent correlations with other traits related to water-carbon balance. Our results suggest that plant adaptation to water availability underlies the association between leaf pH and iWUE, resulting in their strong geographical coupling. This study highlights the potential eco-physiological and eco-geographical significance of leaf pH and introduces it into the framework of plant water-carbon balance research.
Aim Global climate change has intensified the extinction risk of plant species in tropical forests. Yet, empirical validation of how individual species respond to climate change remains lacking, and the demographic mechanisms linking individual mortality to long-term extinction risk are poorly understood. Location Panama, Central America. Time Period 1982-2100.Major Taxa Studies Woody plants. Methods Using replicated tree census data from 44 forest plots in Central American tropical forests, comprising > 400,000 individuals and > 130,000 records of rare species survival, we examined the mortality dynamics and extinction risk of rare plants under climate change through a spatially explicit, trait-based forest dynamics model. Results We found that rare species exhibited consistently higher mortality than common species, driven jointly by climatic stress and biotic interactions. Specifically, mortality increased with warming, vapour pressure deficit and neighbourhood density, but decreased with individual basal area. Functional traits, particularly those related to hydraulics and structures, modulated species-specific responses and vulnerability. Over the last two decades, rare species experienced an average loss of 16.8% per plot, with climate-driven changes accounting for approximately one-quarter of this loss. By 2100, depending on the climate scenarios, 53.3%-64.2% of native rare species are projected to be extirpated locally, and 9.4%-27.8% to become regionally extinct. Main Conclusions Our findings demonstrate that rare species face elevated mortality and accelerated extinction risk under climate warming, with functional traits playing an important role in shaping species-specific responses. By linking individual demography, species traits and long-term forest dynamics, this study provides a mechanistic framework for extinction risk assessments and highlights the need for trait-informed conservation strategies to safeguard tropical biodiversity under climate change.
Anthropogenic nitrogen (N) and phosphorus (P) inputs have significant effects on plant community composition and diversity in forest ecosystems. Compared to canopy layers, understory communities are more sensitive to exogenous nutrient inputs. Although some experimental evidence exists on the effects of N addition on understory communities, knowledge about the impacts of P addition and its interaction with N, as well as the underlying mechanisms, remains limited. Here, we conducted two nutrient manipulation experiments in tropical primary and secondary montane rainforests, and synthesized 33 peer-reviewed studies to evaluate the impacts of N and P enrichments on species richness, density, and cover of understory plants in forests. Integrated evidence from field experiments and meta-analyses consistently demonstrated that nutrient enrichment with N, P, or both significantly reduced the species richness, density, and cover of understory plant communities. The negative impacts intensified with experimental duration and addition rate. Mechanistically, alteration in soil conditions including excessive accumulations of N and P as well as N-induced soil acidification mainly caused the loss of understory plant species. Our study underscores the negative influences of anthropogenic nutrient inputs on understory community diversity and coverage, thereby altering forest structure and functioning in the future.
Bamboo forests substantially contribute to the biogeochemical cycling of carbon (C) and help mitigate climate change. Moso bamboo (Phyllostachys edulis), which occupies the largest bamboo forest area globally, is widely distributed across China. However, the C stocks of these forests and their controlling factors remain poorly quantified due to a lack of large-scale field data. Here, we conducted a nationwide survey of 322 plots and 1,245 soil samples throughout the full distribution range of moso bamboo forests in China. We estimated a total ecosystem C stock of 511.0 +/- 9.9 Tg C, with 29% stored in vegetation and 71% in soil (0-50 cm). Vegetation C density was weakly influenced by climate but increased under moderate human disturbance. Soil C density was shaped by both climate and human activity: higher temperature, precipitation, and wetness index promoted soil C accumulation, whereas intensive disturbance indirectly reduced soil C by decreasing soil moisture and nitrogen content. These findings suggest that increasing drought and intensified human activity may reduce soil C sequestration in bamboo ecosystems. Our study provides a new field-based estimate of C stocks in China's bamboo forests and offers insights to improve biogeochemical models and inform C sink management.
Plant-insect herbivore interactions are essential in shaping forest ecosystem health. The resource availability hypothesis (RAH) and the leaf economics spectrum (LES) theory predict that species in high-resource environments tend to adopt a 'fast' strategy but are more susceptible to herbivory. However, this contradicts the reports of increased insect herbivory in the context of global drought intensification and hinders accurate prediction about how different plant species respond to herbivorous insect feeding. To fill this knowledge gap, we conducted an observational study in two temperate forests dominated by Quercus mongolica and Betula platyphylla in eastern China to compare their leaf herbivory patterns and explore possible mechanisms. We measured three leaf herbivory proxies (consumed leaf area, percent consumed and herbivory frequency), some leaf traits (leaf area [LA], specific leaf area [SLA], leaf water content [LWC], leaf nitrogen, phosphorus and non-structural carbohydrate contents) and soil properties (pH, soil water content [SWC], soil organic carbon content, soil nitrogen and phosphorus contents). We found that Q. mongolica, growing in poorer soil environments with lower water and nutrient contents, experienced higher leaf herbivory than B. platyphylla. Regarding leaf traits, Q. mongolica had a higher LA and non-structural carbohydrate content, but lower SLA, leaf nutrient and water contents, than B. platyphylla. At the leaf level, LA, rather than SLA, of both tree species was positively correlated with leaf herbivory. At the tree level, species-specific patterns emerged, that is, leaf herbivory of B. platyphylla was positively related to LA and negatively related to leaf nitrogen and water contents and soil phosphorus content, whereas that of Q. mongolica was only positively affected by soil phosphorus content. These findings challenge the predictions of RAH and LES theory, as Q. mongolica that grows in resource-poor soil environments with a conservative strategy suffers higher leaf herbivory than B. platyphylla, shedding some light on the proverb that trouble follows the needy. Moreover, water-related factors (i.e. LWC and SWC) and LA showed an important effect on driving interspecific and intraspecific leaf herbivory variations here, implying that climate-induced droughts may exacerbate herbivore pressure in temperate forests.Read the free Plain Language Summary for this article on the Journal blog.
Plant biomass and its allocation are fundamental for understanding biospheric matter production. However, the impacts of atmospheric phosphorus (P) deposition on species-specific biomass and its allocation in global terrestrial plants remain unclear. By synthesizing 5548 observations of plant biomass and its allocation related to P addition worldwide, we find that P addition increases plant biomass by an average of 35% globally. This increase varies across plant functional groups, with stronger responses in deciduous (45%), C3 (36%), and N2-fixing plants (54%) than in evergreen (28%), C4 (19%), and non-N2-fixing plants (31%), respectively. Plants possessing traits indicative of an acquisitive strategy, such as higher nutrient concentrations and specific leaf area, faster photosynthetic rates and shorter leaf lifespan, are particularly responsive to P addition. Furthermore, P addition promotes a greater allocation of biomass to aboveground than belowground organs, resulting in a 5% decrease in root-to-shoot ratio. Our findings provide global-scale quantifications of how P addition regulates biomass accumulation and allocation strategies in terrestrial plants, offering critical insights for predicting the response of terrestrial carbon storage to rising atmospheric P deposition.
Ecological stoichiometry in the plant-soil-microbial systems is crucial for regulating nutrient flow and sustaining ecosystem functions in forests. However, how rising atmospheric nitrogen (N) deposition induced by human activities impacts the stoichiometric relationships across ecosystem compartments remains poorly understood. We therefore conducted a 13-year N-addition experiment across eight forests from tropical to boreal zones in China, to assess the shifts in carbon:nitrogen:phosphorus (C:N:P) stoichiometry across live plant tissues, detritus, soil, and microbes. The N addition significantly elevated N concentrations in green leaves (+7.2% to 10.1%), twigs (+8.4%), fine roots (+20.2%), and leaf litter (+5.9%), while decreasing microbial N (-16.0%) and P concentrations (-20.2% to - 12.1%) across compartments. These changes triggered compartment-specific responses, with greater stoichiometric variability in plants than in soils, particularly in high-latitude forests. However, N addition did not significantly alter the intrinsic scaling relationships of P concentrations and N:P ratios among compartments or the N-P scaling patterns within individual compartments. Our findings provide new insights into the resilience and complexity of nutrient regulation in forests under chronic N deposition, with important implications for predicting long-term ecosystem responses and developing sustainable nutrient management strategies.
China represents a significant global hotspot for species in the family Fagaceae, which are widely distributed across the country and play a crucial role in various ecological and social systems. As the global cliamte is changing rapidly, predicting the future distribution and richness of these species in China holds substantial importance. This study presents the first national-scale assessment of the future distribution of 243 Fagaceae species in China, utilizing ensemble species distribution models (SDMs) for the 2050s and 2070s under various climate change scenarios. The SDM projections indicate notable changes in the distribution of Fagaceae species, characterizing with an overall decline in the distribution area, an upward migration in elevation and a northeastward shift in their range. These changes are expected to significantly alter the spatial pattern of species richness, creating possible refugia in the southwestern mountainous regions and the western Qinling Mountains. We further revealed that a considerable amount of China's natural reserves will show decreased richness of Fagaceae under climate change. Our study systematically evaluates the impact of future climate change on the distribution of Fagaceae species in China, potentially helpful for conservation planning of these species in China. (sic)(sic) (sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic) (sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic),(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic);(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic).(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic),(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic).(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)2050,2070(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic),(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)243(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic).(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic),(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic),(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic).(sic)(sic)(sic)(sic),(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic),(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic).(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic),(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic).(sic)(sic)(sic)(sic)(sic)(sic)(sic),(sic)(sic)(sic)(sic)(sic)(sic)(sic),(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic).(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic)(sic).