Changes in productivity partitioning induced by precipitation extremes increase inaccuracy of grassland carbon estimation.

The fraction of net primary productivity (NPP) allocated to belowground organs (fBNPP) in grasslands is a critical parameter in global carbon cycle models; moreover, understanding the effect of precipitation changes on this parameter is vital to accurately estimating carbon sequestration in grassland ecosystems. However, how fBNPP responds to temporal precipitation changes along a gradient from extreme drought to extreme wetness, remains unclear, mainly due to the lack of long-term data of belowground net primary productivity (BNPP) and the fact that most precipitation experiments did not have a gradient from extreme drought to extreme wetness. Here, by conducting both a precipitation gradient experiment (100-500 mm) and a long-term observational study (34 years) in the Inner Mongolia grassland, we showed that fBNPP decreased linearly along the precipitation gradient from extreme drought to extreme wetness due to stronger responses in aboveground NPP to drought and wet conditions than those of BNPP. Our further meta-analysis in grasslands worldwide also indicated that fBNPP increased when precipitation decreased, and the vice versa. Such a consistent pattern of fBNPP response suggests that plants increase the belowground allocation with decreasing precipitation, while increase the aboveground allocation with increasing precipitation. Thus, the linearly decreasing response pattern in fBNPP should be incorporated into models that forecast carbon sequestration in grassland ecosystems; failure to do so will lead to underestimation of the carbon stock in drought years and overestimation of the carbon stock in wet years in grasslands.

Application of remote sensing to understand the role of Galician feral horses in the biomass reduction of a shrub-grassland-dominated landscape.

Galician forests in northwestern Spain are subject to frequent wildfires with high environmental and economic costs. In addition, due to the consequences of climate change, these fires are becoming more virulent, occurring throughout the year, and taking place in populated areas, in some cases involving the loss of human life. Therefore, forest fire prevention is even more relevant than mitigating its consequences. Given the costs involved in forestry work, alternative measures to reduce fuel load and create vegetation gaps are needed. One involves grazing by an endemic species of feral horses (Equus ferus atlanticus) that feed on thicket-forming gorse (Ulex europaeus). In a 100-ha forest fenced study area stocked with 11 horses, four 50 m2 enclosed plots prevented the access of these wild animals to the vegetation, with the aim of manipulating their impact on the reduction of forest biomass. The measurement of biomass volumes is an important method that can describe the assessment of wildfire risks, unfortunately, high-resolution data collection at the regional scale is very time-consuming. The best result can be using drones (unmanned aerial vehicles - UAVs) as a method of collecting remotely sensed data at low cost. From September 2018 to November 2020, we collected information about aboveground biomass from these four enclosed plots and their surrounding areas available for horses to forage, via UAV. These data, together with environmental variables from the study site, were used as input for a fire model to assess the differences in the surface rate of spread (SROS) among grazed and ungrazed areas. Our results indicated a consistent but small reduction in the SROS between 0.55 and 3.10 m/min in the ungrazed enclosured plots in comparison to their grazed surrounding areas (which have an SROS between 15 and 25 m/min). The research showed that radar remote sensing (UAV) can be used to map forest aboveground biomass, and emphasized the importance and role of feral horses in Galicia as a prevention tool against wildfires in gorse-dominated landscapes.

Ten years of directing seeding restoration in the Brazilian savanna: Lessons learned and the way forward.

Savannas and grasslands have lost almost 50% of their original cover worldwide. Therefore, the development of methods and information on open-canopy ecosystem restoration is urgent for the inclusion of these ecosystems into global and regional priorities. In the Brazilian savanna, the most diverse savanna in the world, restoration efforts focused on open ecosystems have been virtually absent, but have increased in the last 10 years. Such efforts are frequently threatened by invasive exotic grasses (IEG) that invade and dominate areas excluding native species, oftentimes aided by altered soil conditions. Long-term studies of savanna restoration trajectories are rare. In this study, we surveyed 22 savanna restoration areas established two to ten years before the study with similar restoration methods to assess their current status. We show that the current restoration methods are successful in establishing native species and allowing species turnover but they are threatened by IEG. Restoration success varies and is affected by soil conditions, IEG landscape cover and post-sowing weeding. Despite that, the simultaneous introduction of different plant functional groups allows turnover from fast to slow-growing plants. Establishing savanna native species is possible at an operational scale with current knowledge and techniques. However, native species establishment fails to prevent IEG reinfestation, which needs to be managed in restoration efforts in the Brazilian savanna.

Effects of livestock on arthropod biodiversity in Iberian holm oak savannas revealed by metabarcoding.

Increasing food production while avoiding negative impacts on biodiversity constitutes one of the main challenges of our time. Traditional silvopastoral systems like Iberian oak savannas ("dehesas") set an example, where free-range livestock has been reared for centuries while preserving a high natural value. Nevertheless, factors decreasing productivity need to be addressed, one being acorn losses provoked by pest insects. An increased and focalized grazing by livestock on infested acorns would kill the larvae inside and decrease pest numbers, but increased livestock densities could have undesired side effects on ground arthropod communities as a whole. We designed an experimental setup including areas under trees with livestock exclosures of different ages (short-term: 1-year exclusion, long-term: 10-year exclusion), along with controls (continuous grazing), using DNA metabarcoding (mitochondrial markers COI and 16S) to rapidly assess arthropod communities' composition. Livestock removal quickly increased grass cover and arthropod taxonomic richness and diversity, which was already higher in short-term (1-year exclosures) than beneath the canopies of control trees. Interestingly, arthropod diversity was not highest at long-term exclosures (≥10 years), although their community composition was the most distinct. Also, regardless of treatment, we found that functional diversity strongly correlated with the vegetation structure, being higher at trees beneath which there was higher grass cover and taller herbs. Overall, the taxonomic diversity peak at short term exclosures would support the intermediate disturbance hypothesis, which relates it with the higher microhabitat heterogeneity at moderately disturbed areas. Thus, we propose a rotatory livestock management in dehesas: plots with increased grazing should co-exist with temporal short-term exclosures. Ideally, a few long-term excluded areas should be also kept for the singularity of their arthropod communities. This strategy would make possible the combination of biological pest control and arthropod conservation in Iberian dehesas.

Pyric herbivory decreases soil denitrification despite increased nitrate availability in a temperate grassland.

Pyric herbivory, the combination of controlled burning and targeted grazing, is an effective strategy for restoring abandoned, shrub-encroached rangelands to open ecosystems. This practice may impact soil nitrogen pools by altering soil nitrification and denitrification rates, and may lead to an increase of nitrogen losses through nitrate leaching and N-gas emissions. This research, located in the south-western Pyrenees, investigated the effects of pyric herbivory on soil nitrification and denitrification potentials and mineral nitrogen content in a gorse-encroached temperate rangeland six months after the burning was implemented. The study included three treatments: high-severity burning plus grazing, low-severity burning plus grazing, and unburned and ungrazed areas (control). We measured soil nitrification and denitrification potentials (net and gross), the limitation of denitrifiers by nitrogen or organic carbon, and the abundance of nitrite- and nitrous oxide-reducing bacteria. Additional soil and vegetation data complemented these measurements. Results showed that pyric herbivory did not significantly affect nitrification potential, which was low and highly variable. However, it decreased gross denitrification potential and nitrous oxide reduction to dinitrogen in high-severely burned areas compared to the control. Denitrification rates directly correlated with microbial biomass nitrogen, soil organic carbon, soil water content and abundance of nirS-harbouring bacteria. Contrary to the expected, soil nitrate availability did not directly influence denitrification despite being highest in burned areas. Overall, the study suggests that pyric herbivory does not significantly affect mid-term nitrification rates in temperate open ecosystems, but may decrease denitrification rates in intensely burned areas. These findings highlight the importance of assessing the potential impacts of land management practices, such as pyric herbivory, on soil nutrient cycling and ecosystem functioning.

The impact of compound drought and heatwave events from 1982 to 2022 on the phenology of Central Asian grasslands.

In the context of global warming, the occurrence and severity of extreme events like atmospheric drought (AD) and warm spell duration index (WSDI) have increased, causing significant impacts on terrestrial ecosystems in Central Asia's arid regions. Previous research has focused on single extreme events such as AD and WSDI, but the effect of compound hot and dry events (CHWE) on grassland phenology in the arid regions of Central Asia remains unclear. This study utilized structural equation modeling (SEM) and the Pettitt breakpoint test to quantify the direct and indirect responses of grassland phenology (start of season - SOS, length of season - LOS, and end of season - EOS) to AD, WSDI, and CHWE. Furthermore, this research investigated the threshold of grassland phenology response to compound hot and dry events. The research findings indicate a significant increasing trend in AD, WSDI, and CHWE in the arid regions of Central Asia from 1982 to 2022 (0.51 day/year, P < 0.01; 0.25 day/year, P < 0.01; 0.26 day/year, P < 0.01). SOS in the arid regions of Central Asia showed a significant advancement trend, while EOS exhibited a significant advance. LOS demonstrated an increasing trend (-0.23 day/year, P < 0.01; -0.12 day/year, P < 0.01; 0.56 day/year). The temperature primarily governs the variation in SOS. While higher temperatures promote an earlier SOS, they also offset the delaying effect of CHWE on SOS. AD, temperature, and CHWE have negative impacts on EOS, whereas WSDI has a positive effect on EOS. AD exhibits the strongest negative effect on EOS, with an increase in AD leading to an earlier EOS. Temperature and WSDI are positively correlated with LOS, indicating that higher temperatures and increased WSDI contribute to a longer LOS. The threshold values for the response of SOS, EOS, and LOS to CHWE are 16.14, 18.49, and 16.61 days, respectively. When CHWE exceeds these critical thresholds, there are significant changes in the response of SOS, EOS, and LOS to CHWE. These findings deepen our understanding of the mechanisms by which extreme climate events influence grassland phenology dynamics in Central Asia. They can contribute to better protection and management of grassland ecosystems and help in addressing the impacts of global warming and climate change in practice.

Evaluation of an autonomous acoustic surveying technique for grassland bird communities in Nebraska.

Monitoring trends in wildlife communities is integral to making informed land management decisions and applying conservation strategies. Birds inhabit most niches in every environment and because of this they are widely accepted as an indicator species for environmental health. Traditionally, point counts are the common method to survey bird populations, however, passive acoustic monitoring approaches using autonomous recording units have been shown to be cost-effective alternatives to point count surveys. Advancements in automatic acoustic classification technologies, such as BirdNET, can aid in these efforts by quickly processing large volumes of acoustic recordings to identify bird species. While the utility of BirdNET has been demonstrated in several applications, there is little understanding of its effectiveness in surveying declining grassland birds. We conducted a study to evaluate the performance of BirdNET to survey grassland bird communities in Nebraska by comparing this automated approach to point count surveys. We deployed ten autonomous recording units from March through September 2022: five recorders in row-crop fields and five recorders in perennial grassland fields. During this study period, we visited each site three times to conduct point count surveys. We compared focal grassland bird species richness between point count surveys and the autonomous recording units at two different temporal scales and at six different confidence thresholds. Total species richness (focal and non-focal) for both methods was also compared at five different confidence thresholds using species accumulation curves. The results from this study demonstrate the usefulness of BirdNET at estimating long-term grassland bird species richness at default confidence scores, however, obtaining accurate abundance estimates for uncommon bird species may require validation with traditional methods.

Climate change will lead to range shifts and genetic diversity losses of dung beetles in the Gobi Desert and Mongolian Steppe.

Desertification is known to be a major threat to biodiversity, yet our understanding of the consequent decline in biodiversity remains insufficient. Here, we predicted climate change-induced range shifts and genetic diversity losses in three model dung beetles: Colobopterus erraticus, Cheironitis eumenes, and Gymnopleurus mopsus, distributed across the Gobi Desert and Mongolian Steppe, areas known for desertification. Phylogeographic analyses of mitochondrial COI sequences and species distribution modeling, based on extensive field investigations spanning 14 years, were performed. Species confined to a single biome were predicted to contract and shift their distribution in response to climate change, whereas widespread species was predicted to expand even if affected by range shifts. We indicated that all species are expected to experience significant haplotype losses, yet the presence of high singleton frequencies and low genetic divergence across geographic configurations and lineages mitigate loss of genetic diversity. Notably, Cheironitis eumenes, a desert species with low genetic diversity, appears to be the most vulnerable to climate change due to the extensive degradation in the Gobi Desert. This is the first study to predict the response of insects to desertification in the Gobi Desert. Our findings highlight that dung beetles in the Gobi Desert and Mongolian Steppe might experience high rates of occupancy turnover and genetic loss, which could reshuffle the species composition.

Sucrose-Phosphate Synthase and Sucrose Synthase contribute to refoliation in ryegrass, a grassland fructan-accumulating species.

The perennity of grassland species such as Lolium perenne greatly depends on their ability to regrow after cutting or grazing. Refoliation largely relies on the mobilization of fructans in the remaining tissues and on the associated sucrose synthesis and transport towards the basal leaf meristems. However, nothing is known yet about the sucrose synthesis pathway. Sucrose Phosphate Synthase (SPS) and Sucrose Synthase (SuS) activities, together with their transcripts, were monitored during the first hours after defoliation along the leaf axis of mature leaf sheaths and elongating leaf bases (ELB) where the leaf meristems are located. In leaf sheaths, which undergo a sink-source transition, fructan and sucrose contents declined while SPS and SuS activities increased, along with the expression of LpSPSA, LpSPSD.2, LpSuS1, LpSuS2, and LpSuS4. In ELB, which continue to act as a strong carbon sink, SPS and SuS activities increased to varying degrees while the expression of all the LpSPS and LpSuS genes decreased after defoliation. SPS and SuS both contribute to refoliation but are regulated differently depending on the source or sink status of the tissues. Together with fructan metabolism, they represent key determinants of ryegrass perennity and, more generally, of grassland sustainability.

Plant roots affect free-living diazotroph communities in temperate grassland soils despite decades of fertilization.

Fixation of atmospheric N2 by free-living diazotrophs accounts for an important proportion of nitrogen naturally introduced to temperate grasslands. The effect of plants or fertilization on the general microbial community has been extensively studied, yet an understanding of the potential combinatorial effects on the community structure and activity of free-living diazotrophs is lacking. In this study we provide a multilevel assessment of the single and interactive effects of different long-term fertilization treatments, plant species and vicinity to roots on the free-living diazotroph community in relation to the general microbial community in grassland soils. We sequenced the dinitrogenase reductase (nifH) and the 16S rRNA genes of bulk soil and root-associated compartments (rhizosphere soil, rhizoplane and root) of two grass species (Arrhenatherum elatius and Anthoxanthum odoratum) and two herb species (Galium album and Plantago lanceolata) growing in Austrian grassland soils treated with different fertilizers (N, P, NPK) since 1960. Overall, fertilization has the strongest effect on the diazotroph and general microbial community structure, however with vicinity to the root, the plant effect increases. Despite the long-term fertilization, plants strongly influence the diazotroph communities emphasizing the complexity of soil microbial communities' responses to changing nutrient conditions in temperate grasslands.

Assessing changes in the ecosystem service value in response to land use and land cover dynamics in Malawi.

Land use and land cover (LULC) changes are inevitable outcomes of socioeconomic changes and greatly affect ecosystem services. Our study addresses the critical gap in the existing literature by providing the first comprehensive national analysis of LULC changes and their impacts on ecosystem service values (ESVs) in Malawi. We assessed changes in ecosystem service values (ESVs) in response to LULC changes using the benefit transfer method in ArcGIS 10.6 software. Our findings revealed a significant increase in grasslands, croplands, and urban areas and a notable decline in forests, shrubs, wetlands, and water bodies. Grassland, cropland, and built-up areas expanded by 52%, 1%, and 23.2%, respectively. In contrast, permanent wetlands, barren land, and water bodies declined by 27.6%, 34.3%, and 1%, respectively. The ESV declined from US$90.87 billion in 2001 to US$85.60 billion in 2022, marking a 5.8% reduction. Provisioning services increased by 0.5% while regulating, supporting, and cultural ecosystem service functions declined by 12.2%, 3.16%, and 3.22%, respectively. The increase in provisioning services was due to the expansion of cropland. However, the loss of regulating, supporting, and cultural services was mainly due to the loss of natural ecosystems. Thus, environmental policy should prioritise the conservation and restoration of natural ecosystems to enhance the ESV of Malawi.

Fast grassland recovery from viable propagules after reintroducing traditional mowing management on a steep slope.

Semi-natural grasslands on steep slopes often show high plant species diversity. These grasslands were traditionally maintained through mowing and/or grazing. The traditional management practices help to maintain species diversity, whereas land abandonment reduces diversity by increasing competition from dominant species and reducing seedling recruitment. The reintroduction of management can reverse species diversity declines, but suitable grassland restoration programs are scarce in Japan. To study the effect of short-term abandonment on seedling ecology, we monitored the vegetation of a Susogari grassland that had been abandoned for 3 years; the grassland occupies a steep slope (ca. 50°) on a hillside above paddy fields, and was traditionally mown. We monitored the vegetation before abandonment, in the 3rd year of abandonment, and in the 1st and 2nd years after restoration of mowing management. Emergence and survival of seedlings was monitored for 18 months after reintroduction of management. We monitored 1,183 seedlings of grassland species and non-target annuals in ten 1-m2 plots. After mowing was reintroduced, most grassland species reappeared or increased in the first and second years. Few seedlings of perennial plants and no seedlings of annuals flowered. An exotic species, Solidago altissima, had a lower survival rate (10%) than grassland species (>30%), and all but two grassland species survived over the 18-month period. Although vegetation composition was not fully recovered, our findings suggest that a steep slope acts as a strong filter that inhibits the establishment of non-target species while enhancing persistence of target grassland species.

Size exclusion experiment in a grassland field unravels top-down control of the soil fauna on microbial community assembly.

Highly diverse and abundant organisms coexist in soils. However, the contribution of biotic interactions between soil organisms to microbial community assembly remains to be explored. Here, we assess the extent to which soil fauna can shape microbial community assembly using an exclusion experiment in a grassland field to sort soil biota based on body size. After 1 year, the exclusion of larger fauna favoured phagotrophic protists, with increases up to 32% in their proportion compared to the no-mesh treatment. In contrast, members of the bacterial community and to a lesser extent of the fungal community were negatively impacted. Shifts in bacterial but not in fungal communities were best explained by the response of the protistan community to exclusion. Our findings provide empirical evidence of top-down control on the soil microbial communities and underline the importance of integrating higher trophic levels for a better understanding of the soil microbiome assembly.

Natural forest regeneration through fire protection is a less imminent threat for truly stable savannas than afforestation.

African bistable savannas have important biodiversity value and merit conservation. At the same time, forest restoration is a nature-based solution that can be used to increase biodiversity, carbon stocks, and human well-being. Here we describe an experiment based on natural forest regeneration through the exclusion of anthropogenic fire. We show that it is easier to let nature do its work instead of channeling it into an artificial man-made ecosystem through human-induced burning or planting. We emphasize that nature-based solutions must be biome-appropriate and the choice between restoring forests or protecting savannas requires a thorough understanding of the local context.

Analysis of rotational grazing management for sheep in mixed grassland.

Sown mixed grassland is rarely used for livestock raising and grazing; however, different forages can provide various nutrients for livestock, which may be beneficial to animal health and welfare. We established a sown mixed grassland and adopted a rotational grazing system, monitored the changes in aboveground biomass and sheep weights during the summer grazing period, measured the nutrients of forage by near-infrared spectroscopy, tested the contents of medium- and long-chain fatty acids by gas chromatography, and explored an efficient sheep fattening system that is suitable for agro-pastoral interlacing areas. The results showed that the maximum forage supply in a single grazing paddock was 4.6 kg DM/d, the highest dry matter intake (DMI) was 1.80 kg DM/ewe/d, the average daily weight gain (ADG) was 193.3 g, the DMI and ADG were significantly correlated (P < 0.05), and the average feed weight gain ratio (F/G) reached 8.02. The average crude protein and metabolizable energy intake by sheep were 286 g/ewe/d and 18.5 MJ/ewe/d respectively, and the n-6/n-3 ratio of polyunsaturated fatty acids in mutton was 2.84. The results indicated that the sheep fattening system had high feed conversion efficiency, could improve the yield and quality of sheep, and could be promoted in suitable regions.

Tibetan Plateau grasslands might increase sequestration of microbial necromass carbon under future warming.

Microbial necromass carbon (MNC) can reflect soil carbon (C) sequestration capacity. However, changes in the reserves of MNC in response to warming in alpine grasslands across the Tibetan Plateau are currently unclear. Based on large-scale sampling and published observations, we divided eco-clusters based on dominant phylotypes, calculated their relative abundance, and found that their averaged importance to MNC was higher than most other environmental variables. With a deep learning model based on stacked autoencoder, we proved that using eco-cluster relative abundance as the input variable of the model can accurately predict the overall distribution of MNC under current and warming conditions. It implied that warming could lead to an overall increase in the MNC in grassland topsoil across the Tibetan Plateau, with an average increase of 7.49 mg/g, a 68.3% increase. Collectively, this study concludes that alpine grassland has the tendency to increase soil C sequestration capacity on the Tibetan Plateau under future warming.

Grazing herbivores reduce herbaceous biomass and fire activity across African savannas.

Fire and herbivory interact to alter ecosystems and carbon cycling. In savannas, herbivores can reduce fire activity by removing grass biomass, but the size of these effects and what regulates them remain uncertain. To examine grazing effects on fuels and fire regimes across African savannas, we combined data from herbivore exclosure experiments with remotely sensed data on fire activity and herbivore density. We show that, broadly across African savannas, grazing herbivores substantially reduce both herbaceous biomass and fire activity. The size of these effects was strongly associated with grazing herbivore densities, and surprisingly, was mostly consistent across different environments. A one-zebra increase in herbivore biomass density (~100 kg/km2 of metabolic biomass) resulted in a ~53 kg/ha reduction in standing herbaceous biomass and a ~0.43 percentage point reduction in burned area. Our results indicate that fire models can be improved by incorporating grazing effects on grass biomass.

Mowing aggravates the adverse effects of nitrogen addition on soil acid neutralizing capacity in a meadow steppe.

Soil acidification induced by reactive nitrogen (N) inputs is a major environmental issue in grasslands, as it lowers the acid neutralizing capacity (ANC). The specific impacts of different N compound forms on ANC remain unclear. Grassland management practices like mowing and grazing can remove a considerable amount of soil N and other nutrients, potentially mitigating soil acidification by removing N from the ecosystem or aggravating it by removing base cations. However, empirical evidence regarding the joint effects of adding different forms of N compounds and mowing on ANC changes in different-sized soil aggregates is still lacking. This study aimed to address this knowledge gap by examining the effects of three N compounds (urea, ammonium nitrate, and ammonium sulfate) combined with mowing (mown vs. unmown) on soil ANC in different soil aggregate sizes (>2000 µm, 250-2000 µm, and <250 µm) through a 6-year field experiment in Inner Mongolia grasslands. We found that the average decline in soil ANC caused by ammonium sulfate (AS) addition (-78.9%) was much greater than that by urea (-25.0%) and ammonium nitrate (AN) (-52.1%) as compared to control. This decline was attributed to increased proton (H+) release from nitrification and the leaching of exchangeable Ca2+ and Mg2+. Mowing aggravated the adverse effects of urea and AN on ANC, primarily due to the reduction in soil organic matter (SOM) contents and the removal of exchangeable Ca2+, K+, and Na + via plant biomass harvest. This pattern was consistent across all aggregate fractions. The lack of variation in soil ANC among different soil aggregate fractions is likely due to the contrasting trend in the distribution of exchangeable Ca2+ and Mg2+. Specifically, the concentration of exchangeable Ca2+ increased with increasing aggregate size, while the opposite was true for that of exchangeable Mg2+. These findings underscore the importance of considering the forms of N compounds when assessing the declines of ANC induced by N inputs, which also calls for an urgent need to reduce N emissions to ensure the sustainable development of the meadow ecosystems.

Beyond fire: Flower production naturally occurs and is also influenced by leaf removal in a Neotropical savanna herb.

Several herbaceous species exhibit mass flowering after fires in Neotropical savannas. However, unequivocal evidence of fire dependency and the consequences for plant reproduction are lacking. In nutrient-poor fire-prone savannas, the damage caused by fire and by other means (e.g., leaf removal, but not necessarily having a negative impact) constrains the maintenance and expansion of plant population by affecting the ability of individuals to recover. Therefore, the compensatory responses of plants to both damages should be convergent in such environments. Using Bulbostylis paradoxa-reported to be fire-dependent to flower-as a model, we investigated the role of fire and leaf removal in anticipating the flowering and reproduction periods, and its possible consequences on seedling establishment. We monitored 70 burned individuals, 70 damaged/clipped, and 35 without damage to estimate time for flowering, seed quality and germination parameters. To expand our sampling coverage, we examined high-resolution images from herbarium collections in the SpeciesLink database. For each herbarium image, we recorded the presence or absence of a fire scar, the month of flowering, and the number of flowering stalks. Bulbostylis paradoxa was fire-stimulated but not dependent on fire to flower, with 65.7% of the individuals flowering in the burned area, 48.6% in the clipped, and 11.4% in the control. This was consistent with the analysis of the herbarium images in which 85.7% of the specimens with flowers had fire scars and 14.3% did not. Burned individuals synchronized flowering and produced more viable seeds. However, the seeds might face a period of unsuitable ecological conditions after early to mid-dry season fires. Flowering of unburned plants was synchronized with the onset of the rainy season. Flexibility in flowering and vegetative reproduction by fragmentation confer to this species, and most likely other plants from the herbaceous layer, the capability of site occupation and population persistence in burned and unburned savanna sites.

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The natural abundance of stable carbon and nitrogen isotopes (δ13C and δ15N) in leaves can provide comprehensive information on the physiological and ecological processes of plants and has been widely used in ecological research. However, recent studies on leaf δ13C and δ15N have focused mainly on woody species, few studies have been conducted on herbs in different vegetation types, and their differences and driving factors are still unclear. In this study, we focused on the herbs in subalpine coniferous forests, alpine shrublands, and alpine mea-dows on the eastern Qinghai-Tibet Plateau, and investigated the differences in leaf δ13C and δ15N of herbs and the driving factors. The results showed that there were significant differences in leaf δ13C and δ15N values of herbs among different vegetation types, with the highest δ13C and δ15N values in alpine meadows, followed by alpine shrublands, and the lowest in subalpine coniferous forests. Using variation partitioning analysis, we revealed that differences in leaf δ13C and δ15N of herbs among various vegetation types were driven by both leaf functional traits and climate factors, with the contribution of leaf functional traits being relatively higher than that of climate factors. Hierarchical partitioning results indicated that mean annual temperature (MAT), chlorophyll content index, leaf nitrogen content per unit area (Narea), and leaf mass per area were the main drivers of leaf δ13C variations of herbs across different vegetation types, while the relative importance of Narea and MAT for variation in leaf δ15N of herbs was much higher than those other variables. There was a strong coupling relationship between leaf δ13C and δ15N as indicated by the result of the ordinary least squares regression. Our findings could provide new insights into understanding the key drivers of leaf δ13C and δ15N variations in herbs across different vegetation types.