Combined effects of fire and drought are not sufficient to slow shrub encroachment in tallgrass prairie.

Woody encroachment-the spread of woody vegetation in open ecosystems-is a common threat to grasslands worldwide. Reversing encroachment can be exceedingly difficult once shrubs become established, particularly clonal species that resprout following disturbance. Single stressors are unlikely to reverse woody encroachment, but using multiple stressors in tandem could be successful in slowing or reversing encroachment. We explored whether increasing fire frequency in conjunction with multi-year drought could reduce growth and survival of encroaching shrubs in a tallgrass prairie in northeastern Kansas, USA. Passive rainout shelters (~ 50% rainfall reduction) were constructed over mature clonal shrubs (Cornus drummondii) and co-existing C4 grasses in two fire treatments (1-year and 4-year burn frequency). Leaf- and whole-plant level physiological responses to drought and fire frequency were monitored in shrubs and grasses from 2019 to 2022. Shrub biomass and stem density following fire were unaffected by five years of consecutive drought treatment, regardless of fire frequency. The drought treatment had more negative effects on grass leaf water potential and photosynthetic rates compared to shrubs. Shrub photosynthetic rates were remarkably stable across each growing season. Overall, we found that five consecutive years of moderate drought in combination with fire was not sufficient to reduce biomass production or stem density in an encroaching clonal shrub (C. drummondii). These results suggest that moderate but chronic press-drought events do not sufficiently stress encroaching clonal shrubs to negatively impact their resilience following fire events, even when fire frequency is high.

Contrasting shrub and grass hydraulic responses to experimental drought.

Whole-plant hydraulics provide important information about responses to water limitation and can be used to understand how plant communities may change in a drier climate when measured on multiple species. Here, we measured above- and belowground hydraulic traits in Cornus drummondii, an encroaching shrub within North American tallgrass prairies, and Andropogon gerardii, a dominant C4 grass, to assess the potential hydraulic responses to future drought as this region undergoes woody expansion. Shelters that reduced precipitation by 50% and 0% were built over shrubs and grasses growing in sites that are burned at 1-year and 4-year frequencies. We then measured aboveground (Kshoot), belowground (Kroot), and whole-plant maximum hydraulic conductance (Kplant) in C. drummondii and Kroot in A. gerardii. We also measured vulnerability to embolism (P50) in C. drummondii stems. Overall, we show that: (1) A. gerardii had substantially greater Kroot than C. drummondii; (2) belowground hydraulic functioning was linked with aboveground processes; (3) above- and belowground C. drummondii hydraulics were not negatively impacted by the rainfall reductions imposed here. These results suggest that a multi-year drought will not ameliorate rates of woody expansion and highlight key differences in aboveground and belowground hydraulics for dominant species within the same ecosystem.

Seedling responses to soil moisture amount versus pulse frequency in a successfully encroaching semi-arid shrub.

Rainfall timing, frequency, and quantity is rapidly changing in dryland regions, altering dryland plant communities. Understanding dryland plant responses to future rainfall scenarios is crucial for implementing proactive management strategies, particularly in light of land cover changes concurrent with climate change. One such change is woody plant encroachment, an increasing abundance of woody plants in areas formerly dominated by grasslands or savannas. Continued woody plant encroachment will depend, in part, on seedling capacity to establish and thrive under future climate conditions. Seedling performance is primarily impacted by soil moisture conditions governed by precipitation amount (quantity) and frequency. We hypothesized that (H1) seedling performance would be enhanced by both greater soil moisture and pulse frequency, such that seedlings with similar mean soil moisture would perform best under high pulse frequency. Alternatively, (H2) mean soil moisture would have greater influence than pulse frequency, such that a given pulse frequency would have little influence on seedling performance. The hypotheses were tested with Prosopis velutina, a shrub native to the United States that has encroached throughout its range and is invasive in other continents. Seedlings were grown in a greenhouse under two soil moisture treatments, each which was maintained by two pulse frequency treatments. Contrary to H1, mean soil moisture had greater impact than pulse frequency on seedling growth, photosynthetic gas exchange, leaf chemistry, and biomass allocation. These results indicate that P. velutina seedlings may be more responsive to rainfall amount than frequency, at least within the conditions seedlings experienced in this experimental manipulation.

Fire frequency, state change and hysteresis in tallgrass prairie.

Hysteresis is a fundamental characteristic of alternative stable state theory, yet evidence of hysteresis is rare. In mesic grasslands, fire frequency regulates transition from grass- to shrub-dominated system states. It is uncertain, however, if increasing fire frequency can reverse shrub expansion, or if grass-shrub dynamics exhibit hysteresis. We implemented annual burning in two infrequently burned grasslands and ceased burning in two grasslands burned annually. With annual fires, grassland composition converged on that of long-term annually burned vegetation due to rapid recovery of grass cover, although shrubs persisted. When annual burning ceased, shrub cover increased, but community composition did not converge with a long-term infrequently burned reference site because of stochastic and lagged dispersal by shrubs, reflecting hysteresis. Our results demonstrated that annual burning can slow, but not reverse, shrub encroachment. In addition, reversing fire frequencies resulted in hysteresis because vegetation trajectories from grassland to shrubland differed from those of shrubland to grassland.

Biogeography of global drylands.

Despite their extent and socio-ecological importance, a comprehensive biogeographical synthesis of drylands is lacking. Here we synthesize the biogeography of key organisms (vascular and nonvascular vegetation and soil microorganisms), attributes (functional traits, spatial patterns, plant-plant and plant-soil interactions) and processes (productivity and land cover) across global drylands. These areas have a long evolutionary history, are centers of diversification for many plant lineages and include important plant diversity hotspots. This diversity captures a strikingly high portion of the variation in leaf functional diversity observed globally. Part of this functional diversity is associated with the large variation in response and effect traits in the shrubs encroaching dryland grasslands. Aridity and its interplay with the traits of interacting plant species largely shape biogeographical patterns in plant-plant and plant-soil interactions, and in plant spatial patterns. Aridity also drives the composition of biocrust communities and vegetation productivity, which shows large geographical variation. We finish our review by discussing major research gaps, which include: studying regular vegetation spatial patterns; establishing large-scale plant and biocrust field surveys assessing individual-level trait measurements; knowing whether the impacts of plant-plant and plant-soil interactions on biodiversity are predictable; and assessing how elevated CO2 modulates future aridity conditions and plant productivity.

Woody encroachment happens via intensification, not extensification, of species ranges in an African savanna.

Widespread woody encroachment is a prominent concern for savanna systems as it is often accompanied by losses in productivity and biodiversity. Extensive ecosystem-level work has advanced our understanding of its causes and consequences. However, there is still debate over whether local management can override regional and global drivers of woody encroachment, and it remains largely unknown how encroachment influences woody community assemblages. Here, we examined species-level changes in woody plant distributions and size structure from the late 1980s to the late 2000s based on spatially intensive ground-based surveys across Kruger National Park, South Africa. This study region spans broad gradients in rainfall, soil texture, fire frequency, elephant density, and other topographic variables. Species-level changes in frequency of occurrence and size class proportion reflected widespread woody encroachment primarily by Dichrostachys cinerea and Combretum apiculatum, and a loss of large trees mostly of Sclerocarya birrea and Acacia nigrescens. Environmental variables determining woody species distributions across Kruger varied among species but did not change substantially between two sampling times, indicating that woody encroachers were thickening within their existing ranges. Overall, more areas across Kruger were found to have an increased number of common woody species through time, which indicated an increase in stem density. These areas were generally associated with decreasing fire frequency and rainfall but increasing elephant density. Our results suggest that woody encroachment is a widespread but highly variable trend across landscapes in Kruger National Park and potentially reflects an erosion of local heterogeneity in woody community assemblages. Many savanna managers, including in Kruger, aim to manage for heterogeneity in order to promote biodiversity, where homogenization of vegetation structure counters this specific goal. Increasing fire frequency has some potential as a local intervention. However, many common species increased in commonness even under near-constant disturbance conditions, which likely limits the potential for managing woody encroachment in the face of drivers beyond the scope of local control. Regular field sampling coupled with targeted fire management will enable more accurate monitoring of the rate of encroachment intensification.

Fire and browsing interact to alter intra-clonal stem dynamics of an encroaching shrub in tallgrass prairie.

The expansion of woody species into grasslands has altered community structure and ecosystem function of grasslands worldwide. In tallgrass prairie of the Central Great Plains, USA, decreased fire frequency and intensity have increased the cover and abundance of woody species. In particular, clonal shrub cover has increased at accelerated rates due to vegetative reproduction and resprouting after disturbance. We measured the intra-clonal stem demography and relative growth rates (estimated change in woody biomass) of the shrub Cornus drummondii in response to fire frequency (4 vs 20 year burn intervals) and simulated browsing during the 2018 and 2019 growing seasons at Konza Prairie Biological Station (Manhattan, Kansas). Overall, infrequent fire (4 year burn interval) increased intra-clonal stem relative growth rates and shrub relative growth rates. Intra-clonal stem relative growth rates were reduced in unbrowsed clones in 2018 due to drought and simulated browsing reduced intra-clonal stem relative growth rates in 2019. Additionally, simulated browsing nearly eliminated flower production within clones but did not affect intra-clonal stem mortality or recruitment within a growing season. Fire in conjunction with simulated browsing reduced estimated relative growth rates for entire shrub clones. Browsed shrubs that experienced prescribed fire in 2017 had reduced intra-clonal stem densities compared to unbrowsed shrubs and stem densities of browsed shrubs did not recover in 2018 or 2019. These results illustrate that infrequent fire alone promotes the expansion of clonal shrubs in tallgrass prairie and multiple interacting disturbances (e.g., fire and browsing) are required to control the spread of clonal shrubs into grasslands.

Belowground impacts of alpine woody encroachment are determined by plant traits, local climate, and soil conditions.

Global climate and land use change are causing woody plant encroachment in arctic, alpine, and arid/semi-arid ecosystems around the world, yet our understanding of the belowground impacts of this phenomenon is limited. We conducted a globally distributed field study of 13 alpine sites across four continents undergoing woody plant encroachment and sampled soils from both woody encroached and nearby herbaceous plant community types. We found that woody plant encroachment influenced soil microbial richness and community composition across sites based on multiple factors including woody plant traits, site level climate, and abiotic soil conditions. In particular, root symbiont type was a key determinant of belowground effects, as Nitrogen-fixing woody plants had higher soil fungal richness, while Ecto/Ericoid mycorrhizal species had higher soil bacterial richness and symbiont types had distinct soil microbial community composition. Woody plant leaf traits indirectly influenced soil microbes through their impact on soil abiotic conditions, primarily soil pH and C:N ratios. Finally, site-level climate affected the overall magnitude and direction of woody plant influence, as soil fungal and bacterial richness were either higher or lower in woody encroached versus herbaceous soils depending on mean annual temperature and precipitation. All together, these results document global impacts of woody plant encroachment on soil microbial communities, but highlight that multiple biotic and abiotic pathways must be considered to scale up globally from site- and species-level patterns. Considering both the aboveground and belowground effects of woody encroachment will be critical to predict future changes in alpine ecosystem structure and function and subsequent feedbacks to the global climate system.

Soil organic carbon in drylands: shrub encroachment and vegetation management effects dwarf those of livestock grazing.

Dryland ecosystems occur worldwide and play a prominent, but potentially shifting, role in global biogeochemical cycling. Widespread woody plant proliferation, often associated with declines in palatable grasses, has jeopardized livestock production in drylands and prompted attempts to reduce woody cover by chemical or mechanical means. Woody encroachment also has the potential to significantly alter terrestrial carbon storage. However, little is known of the long-term biogeochemical consequences of woody encroachment in the broader context of its interaction with common dryland land uses, including "brush management" (woody plant clearing) and livestock grazing. Present assessments exhibit considerable variation in the consequences of these land use/land cover changes, with evidence that brush management may counteract sizeable impacts of shrub encroachment on soil biogeochemical pools. A challenge to assessing the net effects of brush management in shrub-encroached grasslands on soil organic carbon (SOC) and total nitrogen (N) pools is that land management practices are typically considered in isolation, when they are co-occurring phenomena. Furthermore, few studies have assessed spatial patterns in brush management and how these are affected in decades following treatment on sites with contrasting grazing histories. To address these uncertainties and interactions, we quantified the impacts of shrub encroachment and their subsequent mortality resulting from brush management (herbicide application) on SOC and N pools in a Sonoran Desert grassland where long-term grazing manipulations (>100 yr) co-occur with shrub encroachment and brush management. Pools of SOC and N associated with herbicided shrubs declined markedly over ~40 yr, offsetting 66% of the increases from shrub encroachment. However, spatial patterns in SOC induced by shrubs persisted over the decades following brush management. Century-long protection from grazing did little to change SOC and N pools. Accordingly, shrub encroachment and shrub mortality from brush management each far outweighed livestock grazing impacts. Consideration of the patterns of SOC and N through space (e.g., bole-to-dripline gradients), time (e.g., shrub age/size), land use (e.g., livestock grazing and brush management), and their interactions will position us to improve predictions of SOC and N responses to land use/land cover change, inform C-based management decisions, and objectively evaluate trade-offs with other ecosystem services.

Elephants limit aboveground carbon gains in African savannas.

Understanding the drivers of vegetation carbon dynamics is essential for climate change mitigation and effective policy formulation. However, most efforts focus on abiotic drivers of plant biomass change, with little consideration for functional roles performed by animals, particularly at landscape scales. We combined repeat airborne Light Detection and Ranging with measurements of elephant densities, abiotic factors, and exclusion experiments to determine the relative importance of drivers of change in aboveground woody vegetation carbon stocks in Kruger National Park, South Africa. Despite a growing elephant population, aboveground carbon density (ACD) increased across most of the landscape over the 6-year study period, but at fine scales, bull elephant density was the most important factor determining carbon stock change, with ACD losses recorded only where bull densities exceeded 0.5 bulls/km2 . Effects of bull elephants were, however, spatially restricted and landscape dependent, being especially pronounced along rivers, at mid-elevations, and on steeper slopes. In contrast, elephant herds and abiotic drivers had a comparatively small influence on the direction or magnitude of carbon stock change. Our findings demonstrate that animals can have a substantive influence on regional-scale carbon dynamics and warrant consideration in carbon cycling models and policy formulation aimed at carbon management and climate change mitigation.

Human impacts in African savannas are mediated by plant functional traits.

Tropical savannas have a ground cover dominated by C4 grasses, with fire and herbivory constraining woody cover below a rainfall-based potential. The savanna biome covers 50% of the African continent, encompassing diverse ecosystems that include densely wooded Miombo woodlands and Serengeti grasslands with scattered trees. African savannas provide water, grazing and browsing, food and fuel for tens of millions of people, and have a unique biodiversity that supports wildlife tourism. However, human impacts are causing widespread and accelerating degradation of savannas. The primary threats are land cover-change and transformation, landscape fragmentation that disrupts herbivore communities and fire regimes, climate change and rising atmospheric CO2 . The interactions among these threats are poorly understood, with unknown consequences for ecosystem health and human livelihoods. We argue that the unique combinations of plant functional traits characterizing the major floristic assemblages of African savannas make them differentially susceptible and resilient to anthropogenic drivers of ecosystem change. Research must address how this functional diversity among African savannas differentially influences their vulnerability to global change and elucidate the mechanisms responsible. This knowledge will permit appropriate management strategies to be developed to maintain ecosystem integrity, biodiversity and livelihoods.

Woody plant encroachment amplifies spatial heterogeneity of soil phosphorus to considerable depth.

The geographically extensive phenomenon of woody plant encroachment into grass-dominated ecosystems has strong potential to influence biogeochemical cycles at ecosystem to global scales. Previous research has focused almost exclusively on quantifying pool sizes and flux rates of soil carbon and nitrogen (N), while few studies have examined the impact of woody encroachment on soil phosphorus (P) cycling. Moreover, little is known regarding the impact of woody encroachment on the depth distribution of soil total P at the landscape scale. We quantified patterns of spatial heterogeneity in soil total P along a soil profile by taking spatially explicit soil cores to a depth of 120 cm across a subtropical savanna landscape that has undergone encroachment by Prosopis glandulosa (an N2 -fixer) and other tree/shrub species during the past century. Soil total P increased significantly following woody encroachment throughout the entire 120-cm soil profile. Large groves (>100 m2 ) and small discrete clusters (<100 m2 ) accumulated 53 and 10 g P/m2 more soil P, respectively, compared to grasslands. This P accumulation in soils beneath woody patches is most likely attributable to P uplift by roots located deep in the soil profile (>120 cm) and transfer to upper portions of the profile via litterfall and root turnover. Woody encroachment also altered patterns of spatial heterogeneity in soil total P in the horizontal plane, with highest values at the centers of woody patches, decreasing toward the edges, and reaching lowest values in the surrounding grassland matrix. These spatial patterns were evident throughout the upper 1.2 m of the soil profile, albeit at reduced magnitude deeper in the soil profile. Spatial generalized least squares models indicated that fine root biomass explained a significant proportion of the variation in soil total P both across the landscape and throughout the profile. Our findings suggest that transfer of P from deeper soil layers enlarges the P pool in upper soil layers where it is more actively cycled may be a potential strategy for encroaching woody species to satisfy their P demands.

Soil phosphorus does not keep pace with soil carbon and nitrogen accumulation following woody encroachment.

Soil carbon, nitrogen, and phosphorus cycles are strongly interlinked and controlled through biological processes, and the phosphorus cycle is further controlled through geochemical processes. In dryland ecosystems, woody encroachment often modifies soil carbon, nitrogen, and phosphorus stores, although it remains unknown if these three elements change proportionally in response to this vegetation change. We evaluated proportional changes and spatial patterns of soil organic carbon (SOC), total nitrogen (TN), and total phosphorus (TP) concentrations following woody encroachment by taking spatially explicit soil cores to a depth of 1.2 m across a subtropical savanna landscape which has undergone encroachment by Prosopis glandulosa (an N2 fixer) and other woody species during the past century in southern Texas, USA. SOC and TN were coupled with respect to increasing magnitudes and spatial patterns throughout the soil profile following woody encroachment, while TP increased slower than SOC and TN in topmost surface soils (0-5 cm) but faster in subsurface soils (15-120 cm). Spatial patterns of TP strongly resembled those of vegetation cover throughout the soil profile, but differed from those of SOC and TN, especially in subsurface soils. The encroachment of woody species dominated by N2 -fixing trees into this P-limited ecosystem resulted in the accumulation of proportionally less soil P compared to C and N in surface soils; however, proportionally more P accrued in deeper portions of the soil profile beneath woody patches where alkaline soil pH and high carbonate concentrations would favor precipitation of P as relatively insoluble calcium phosphates. This imbalanced relationship highlights that the relative importance of biotic vs. abiotic mechanisms controlling C and N vs. P accumulation following vegetation change may vary with depth. Our findings suggest that efforts to incorporate effects of land cover changes into coupled climate-biogeochemical models should attempt to represent C-N-P imbalances that may arise following vegetation change.

Savanna woody encroachment is widespread across three continents.

Tropical savannas are a globally extensive biome prone to rapid vegetation change in response to changing environmental conditions. Via a meta-analysis, we quantified savanna woody vegetation change spanning the last century. We found a global trend of woody encroachment that was established prior the 1980s. However, there is critical regional variation in the magnitude of encroachment. Woody cover is increasing most rapidly in the remaining uncleared savannas of South America, most likely due to fire suppression and land fragmentation. In contrast, Australia has experienced low rates of encroachment. When accounting for land use, African savannas have a mean rate annual woody cover increase two and a half times that of Australian savannas. In Africa, encroachment occurs across multiple land uses and is accelerating over time. In Africa and Australia, rising atmospheric CO2 , changing land management and rainfall are likely causes. We argue that the functional traits of each woody flora, specifically the N-fixing ability and architecture of woody plants, are critical to predicting encroachment over the next century and that African savannas are at high risk of widespread vegetation change.

Climate and landscape drive the pace and pattern of conifer encroachment into subalpine meadows.

Mountain meadows have high biodiversity and help regulate stream water release following the snowmelt pulse. However, many meadows are experiencing woody plant encroachment, threatening these ecosystem services. While there have been field surveys of individual meadows and remote sensing-based landscape-scale studies of encroachment, what is missing is a broad-scale, ground-based study to understand common regional drivers, especially at high elevations, where land management has often played a less direct role. With this study, we ask: What are the climate and landscape conditions conducive to woody plant encroachment at the landscape scale, and how has historical climate variation affected tree recruitment in subalpine meadows over time? We measured density of encroaching trees across 340 subalpine meadows in the central Sierra Nevada, California, USA, and used generalized additive models (GAMs) to determine the relationship between landscape-scale patterns of encroachment and meadow environmental properties. We determined ages of trees in 30 survey meadows, used observed climate and GAMs to model the relationship between timing of recruitment and climate since the early 1900s, and extrapolated recruitment patterns into the future using downscaled climate scenarios. Encroachment was high among meadows with lodgepole pine (Pinus contorta Douglas ex Loudon var. murrayana (Balf.) Engelm.) in the immediate vicinity, at lower elevations, with physical conditions favoring strong soil drying, and with maximum temperatures above or below average. Climatic conditions during the year of germination were unimportant, with tree recruitment instead depending on a 3-yr seed production period prior to germination and a 6-yr seedling establishment period following germination. Recruitment was high when the seed production period had high snowpack, and when the seedling establishment period had warm summer maximum temperatures, high summer precipitation, and high snowpack. Applying our temporal model to downscaled output from four global climate models indicated that the average meadow will shift to forest by the end of the 21st century. Sierra Nevada meadow encroachment by conifers is ubiquitous and associated with climate conditions increasingly favorable for tree recruitment, which will lead to substantial changes in subalpine meadows and the ecosystem services they provide.

Rainfall variability counteracts N addition by promoting invasive Lonicera maackii and extending phenology in prairie.

Although encroaching woody plants have reduced the global extent of grasslands, continuing increases in soil nitrogen availability could slow this trend by favoring resident herbaceous species. At the same time, projected increases in rainfall variability could promote woody encroachment by aligning spatiotemporal patterns of soil moisture availability with the needs of woody species. We evaluated the responses of two deciduous woody species to these simulated environmental changes by planting seedlings of Quercus palustris and Lonicera maackii into tallgrass prairie communities grown under a factorial combination of increased rainfall variability and nitrogen addition. Lonicera maackii growth was reduced 20% by nitrogen addition, and increased rainfall variability led to 33% larger seedlings, despite greater competition for light and soil resources. In contrast, Q. palustris growth showed little response to either treatment. Increased rainfall variability allowed both species to retain their leaves for an additional 6.5 d in autumn, potentially in response to wetter end-of-season shallow soils. Our findings suggest increases in rainfall variability will counteract the inhibitory effect of nitrogen deposition on growth of L. maackii, extend autumn phenology, and promote the encroachment of some woody species into grasslands.

Trade-offs between savanna woody plant diversity and carbon storage in the Brazilian Cerrado.

Incentivizing carbon storage can be a win-win pathway to conserving biodiversity and mitigating climate change. In savannas, however, the situation is more complex. Promoting carbon storage through woody encroachment may reduce plant diversity of savanna endemics, even as the diversity of encroaching forest species increases. This trade-off has important implications for the management of biodiversity and carbon in savanna habitats, but has rarely been evaluated empirically. We quantified the nature of carbon-diversity relationships in the Brazilian Cerrado by analyzing how woody plant species richness changed with carbon storage in 206 sites across the 2.2 million km(2) region at two spatial scales. We show that total woody plant species diversity increases with carbon storage, as expected, but that the richness of endemic savanna woody plant species declines with carbon storage both at the local scale, as woody biomass accumulates within plots, and at the landscape scale, as forest replaces savanna. The sharpest trade-offs between carbon storage and savanna diversity occurred at the early stages of carbon accumulation at the local scale but the final stages of forest encroachment at the landscape scale. Furthermore, the loss of savanna species quickens in the final stages of forest encroachment, and beyond a point, savanna species losses outpace forest species gains with increasing carbon accumulation. Our results suggest that although woody encroachment in savanna ecosystems may provide substantial carbon benefits, it comes at the rapidly accruing cost of woody plant species adapted to the open savanna environment. Moreover, the dependence of carbon-diversity trade-offs on the amount of savanna area remaining requires land managers to carefully consider local conditions. Widespread woody encroachment in both Australian and African savannas and grasslands may present similar threats to biodiversity.

Shifts in functional traits elevate risk of fire-driven tree dieback in tropical savanna and forest biomes.

Numerous predictions indicate rising CO2 will accelerate the expansion of forests into savannas. Although encroaching forests can sequester carbon over the short term, increased fires and drought-fire interactions could offset carbon gains, which may be amplified by the shift toward forest plant communities more susceptible to fire-driven dieback. We quantify how bark thickness determines the ability of individual tree species to tolerate fire and subsequently determine the fire sensitivity of ecosystem carbon across 180 plots in savannas and forests throughout the 2.2-million km(2) Cerrado region in Brazil. We find that not accounting for variation in bark thickness across tree species underestimated carbon losses in forests by ~50%, totaling 0.22 PgC across the Cerrado region. The lower bark thicknesses of plant species in forests decreased fire tolerance to such an extent that a third of carbon gains during forest encroachment may be at risk of dieback if burned. These results illustrate that consideration of trait-based differences in fire tolerance is critical for determining the climate-carbon-fire feedback in tropical savanna and forest biomes.

Woody encroachment induced earlier and extended growing season in boreal wetland ecosystems.

Woody plant encroachment (WPE), a widespread ecological phenomenon globally, has significant impacts on ecosystem structure and functions. However, little is known about how WPE affects phenology in wetland ecosystems of middle and high latitudes. Here, we investigated the regional-scale effects of WPE on the start (SOS), peak (POS), end (EOS), and length (GSL) of the growing season in boreal wetland ecosystems, and their underlying mechanisms, using remote sensing dataset during 2001-2016. Our results showed that WPE advanced the annual SOS and POS, while delaying EOS and extending GSL in boreal wetlands with these impacts increasing over time. When boreal wetland ecosystems were fully encroached by woody plants, the SOS and POS were advanced by 12.17 and 5.65 days, respectively, the EOS was postponed by 2.74 days, and the GSL was extended by 15.21 days. We also found that the impacts of WPE on wetland SOS were predominantly attributed to the increased degree of WPE (α), while climatic factors played a more significant role in controlling the POS and EOS responses to WPE. Climate change not only directly influenced phenological responses of wetlands to WPE but also exerted indirect effects by regulating soil moisture and α. Winter precipitation and spring temperature primarily determined the effects of WPE on SOS, while its impacts on POS were mainly controlled by winter precipitation, summer temperature, and precipitation, and the effects on EOS were mainly determined by winter precipitation, summer temperature, and autumn temperature. Our findings offer new insights into the understanding of the interaction between WPE and wetland ecosystems, emphasizing the significance of considering WPE effects to ensure accurate assessments of phenology changes.

Eastern redcedar roots create legacy effects that suppresses growth of prairie species.

The expansion of woody species from their historical ranges into grasslands is a global problem. Understanding the mechanisms that enable species to successfully establish and then re-encroach following their removal is critical to effectively managing problem species. Legacy effects are a mechanism that could be critical to the reestablishment of woody encroachers following their removal. Legacy effects occur when a species alters the biotic and abiotic environment in a way that affects communities that establish subsequently. In this study, we assess whether the eastern redcedar (Juniperus virginiana), a North American woody encroacher, generates legacy effects that affect communities that establish following removal of this species from an experimental grass community. We conducted a series of experiments to evaluate the effects of J. virginiana, roots on the germination and growth of grasses and to determine if the effects of root-addition treatments were derived from a microbial or allelopathic origin. Aqueous extracts of J. virginiana roots were found to inhibit the germination of grasses. We found escalating suppression of overall community biomass and the biomass of each individual species with increasing root treatments. Finally, we determined the origin of the observed suppressive effect is unlikely to be of microbial origin. Synthesis: Our results suggest that J. virginiana exudes an allelochemical into soils that inhibits the growth of certain grasses and thus has the potential to have legacy effects on future occupants. We suggest that the inhibition of the development of grasses in areas where J. virginiana has been removed is a mechanism that may favor the reestablishment of J. virginiana. Our results indicate the legacy effects of J. virginiana must be considered when conducting removal and restoration of J. virginiana infested lands.