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Background Many plants survive fire by resprouting from lignotubers partially or completely buried in the ground. Resprouting success after fire could vary with fire season due to changes in plant carbohydrate reserves, soil moisture, and plant water status, among other factors. It may also depend on plant size, as this affects lignotuber depth in the soil. We investigated the effects of fire season on resprouting success (%) and speed (time to reach 50% of resprouting, t 50 ) in two lignotuberous species ( Erica arborea and Phillyrea angustifolia ) in a shrubland of central Spain. Over three consecutive years, three 13 × 14 m plots were burned early (ES) and late (LS) in the fire season (end of June and early October, respectively). Before fire, individuals were permanently marked after being classified as small, medium, or large. Predawn plant water potential (? pd ) was measured during the fire season. Resprouting was monitored for up to two years. We used repeated measures ANOVA to test the effects of fire season and year (within-subjects effects), and plant size (between-subjects effects) on resprouting success (%) and t 50 . Results Resprouting success in E . arborea increased from ES to LS fires, and with plant size, while P . angustifolia was unaffected by fire season or plant size. The fire year did not directly affect resprouting success but interacted with fire season (for both species) and plant size ( E . arborea ). t 50 was unaffected by fire season in E . arborea , but it significantly affected P . angustifolia , which showed smaller values after ES burns. t 50 varied by fire year in both species, especially when plant water potentials were low. Plant size also affected t 50 in P . angustifolia . ? pd and t 50 for both species combined were negatively related, with a more pronounced effect in LS fires. Conclusions Our results show that the demography of the two resprouters will be differentially affected by fire season, alone or in interaction with the fire year and plant size. E . arborea was highly sensitive to fire season, particularly the smaller individuals, and to ES burns. In contrast, P . angustifolia was highly insensitive. t 50 response among species differed by fire season, fire year, or size. ? pd determined t 50 of both species combined. These responses could impact long-term population structure and community dynamics of this Mediterranean shrubland.

1. Ecological memory, often determined by the extent and type of retained biological legacies present following disturbance, may produce persistent landscape patterns. However, after fire, the persistence or switch to an alternative state may depend on the complex interplay of ecological memory (biological legacies) and potential effects of new external factors influencing the post-fire environment. The current study assesses both the strength of ecological memory resulting from biological legacies of pre-burn vegetation types as well as post-fire effects of livestock. 2. Following a severe fire in 1999, we set up a network of long-term exclosures to examine the effects of legacies and cumulative herbivory by cattle on fuel types, amounts, distribution, flammability and microenvironmental conditions in two post-fire communities representing alternative firedriven states: pyrophobic Nothofagus pumilio subalpine forests and pyrophytic Nothofagus antarctica tall shrublands in northwestern Patagonia, Argentina. 3. Our results show that the retained post-disturbance legacies of tall shrublands and subalpine forests largely determine fuel and flammability traits of the post-fire plant communities 16 years after fire. The importance of biological legacies retained from the unburned plant communities was reflected by the substantially higher amounts of total fine fuel, greater vertical and horizontal fuel continuity and the higher temperatures reached during experimental tissue combustion at post-fire shrubland compared to post-fire forest sites. 4. We show that herbivores may produce antagonistic effects on flammability by decreasing tissue ignitability, total fine fuel and litter depth, and disrupting the vertical and horizontal fine fuel continuity, therefore reducing the probability of fire propagation. However, cattle can increase ratios of dead to live fine fuels, reduce soil moisture, and inhibit tree height growth to canopy size, consequently impeding the development of a closed pyrophobic forest canopy. 5. Synthesis. Our results support the hypothesis that biological legacies, most importantly the dominance by pyrophytic woody plants that resprout vigorously vs. the dominance by pyrophobic obligate seeders, favour fuel and flammability characteristics at the community level which reinforce the mechanisms maintaining pyrophytic shrublands vs. pyrophobic forests. Herbivory by introduced cattle can partially blur sharp pyrophobic/pyrophytic state boundaries by promoting the development of novel post-fire transitional states.

Despite long-time awareness of the importance of the location of buds in plant biology, research on belowground bud banks has been scant. Terms such as lignotuber, xylopodium and sobole, all referring to belowground bud-bearing structures, are used inconsistently in the literature. Because soil efficiently insulates meristems from the heat of fire, concealing buds below ground provides fitness benefits in fire-prone ecosystems. Thus, in these ecosystems, there is a remarkable diversity of bud-bearing structures. There are at least six locations where belowground buds are stored: roots, root crown, rhizomes, woody burls, fleshy swellings and belowground caudexes. These support many morphologically distinct organs. Given their history and function, these organs may be divided into three groups: those that originated in the early history of plants and that currently are widespread (bud-bearing roots and root crowns); those that also originated early and have spread mainly among ferns and monocots (nonwoody rhizomes and a wide range of fleshy underground swellings); and those that originated later in history and are strictly tied to fire-prone ecosystems (woody rhizomes, lignotubers and xylopodia). Recognizing the diversity of belowground bud banks is the starting point for understanding the many evolutionary pathways available for responding to severe recurrent disturbances.

Understanding and predicting plant response to disturbance is of paramount importance in our changing world. Resprouting ability is often considered a simple qualitative trait and used in many ecological studies. Our aim is to show some of the complexities of resprouting while highlighting cautions that need be taken in using resprouting ability to predict vegetation responses across disturbance types and biomes. There are marked differences in resprouting depending on the disturbance type, and fire is often the most severe disturbance because it includes both defoliation and lethal temperatures. In the Mediterranean biome, there are differences in functional strategies to cope with water deficit between resprouters (dehydration avoiders) and nonresprouters (dehydration tolerators); however, there is little research to unambiguously extrapolate these results to other biomes. Furthermore, predictions of vegetation responses to changes in disturbance regimes require consideration not only of resprouting, but also other relevant traits (e.g. seeding, bark thickness) and the different correlations among traits observed in different biomes; models lacking these details would behave poorly at the global scale. Overall, the lessons learned from a given disturbance regime and biome (e.g. crown-fire Mediterranean ecosystems) can guide research in other ecosystems but should not be extrapolated at the global scale.

Tropical grasses fuel the majority of fires on Earth. In fire‐prone landscapes, enhanced flammability may be adaptive for grasses via the maintenance of an open canopy and an increase in spatiotemporal opportunities for recruitment and regeneration. In addition, by burning intensely but briefly, high flammability may protect resprouting buds from lethal temperatures. Despite these potential benefits of high flammability to fire‐prone grasses, variation in flammability among grass species, and how trait differences underpin this variation, remains unknown. By burning leaves and plant parts, we experimentally determined how five plant traits (biomass quantity, biomass density, biomass moisture content, leaf surface‐area‐to‐volume ratio and leaf effective heat of combustion) combined to determine the three components of flammability (ignitability, sustainability and combustibility) at the leaf and plant scales in 25 grass species of fire‐prone South African grasslands at a time of peak fire occurrence. The influence of evolutionary history on flammability was assessed based on a phylogeny built here for the study species. Grass species differed significantly in all components of flammability. Accounting for evolutionary history helped to explain patterns in leaf‐scale combustibility and sustainability. The five measured plant traits predicted components of flammability, particularly leaf ignitability and plant combustibility in which 70% and 58% of variation, respectively, could be explained by a combination of the traits. Total above‐ground biomass was a key driver of combustibility and sustainability with high biomass species burning more intensely and for longer, and producing the highest predicted fire spread rates. Moisture content was the main influence on ignitability, where species with higher moisture contents took longer to ignite and once alight burnt at a slower rate. Biomass density, leaf surface‐area‐to‐volume ratio and leaf effective heat of combustion were weaker predictors of flammability components. Synthesis. We demonstrate that grass flammability is predicted from easily measurable plant functional traits and is influenced by evolutionary history with some components showing phylogenetic signal. Grasses are not homogenous fuels to fire. Rather, species differ in functional traits that in turn demonstrably influence flammability. This diversity is consistent with the idea that flammability may be an adaptive trait for grasses of fire‐prone ecosystems.

North American grasslands have experienced increased relative abundance of shrubs and trees over the last 150 yr. Alterations in herbivore composition, abundance, and grazing pressure along with changes in fire frequency are drivers that can regulate the transition from grassland to shrubland or woodland (a process known as woody encroachment). Historically, North American grasslands had a suite of large herbivores that grazed and/or browsed (i.e., bison, elk, pronghorn, deer), as well as frequent and intense fires. In the tallgrass prairie, many large native ungulates were extirpated by the 1860s, corresponding with increased homesteading (which led to decreased fire frequencies and intensities). Changes in the frequency and intensity of these two drivers (browsing and fire) have coincided with woody encroachment in tallgrass prairie. Within tallgrass prairie, woody encroachment can be categorized in to two groups: non-resprouting species that can be killed with fire and resprouting species that cannot be killed with fire. Resprouting species require additional active management strategies to decrease abundance and eventually be removed from the ecosystem. In this study, we investigated plant cover, ramet density, and physiological effects of continuous simulated browsing and prescribed fire on Cornus drummondii C.A. Mey, a resprouting clonal native shrub species. Browsing reduced C. drummondii canopy cover and increased grass cover. We also observed decreased ramet density, which allowed for more infilling of grasses. Photosynthetic rates between browsed and unbrowsed control shrubs did not increase in 2015 or 2016. In 2017, photosynthetic rates for browsed shrubs were higher in the unburned site than the unbrowsed control shrubs at the end of the growing season. Additionally, after the prescribed fire, browsed shrubs had ~90% decreased cover, ~50% reduced ramet density, and grass cover increased by ~80%. In the roots of browsed shrubs after the prescribed fire, nonstructural carbohydrates (NSC) experienced a twofold reduction in glucose and a threefold reduction in both sucrose and starch. The combined effects of browsing and fire show strong potential as a successful management tool to decrease the abundance of clonal-resprouting woody plants in mesic grasslands and illustrate the potential significance of browsers as a key driver in this ecosystem.