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Different ecosystems evolved and are maintained by fire, with their vegetation hosting species with a wide diversity of persistence strategies allowing them to insulate their body and resprout new branches after fire disturbance. Changes in fire regime are predicted due to climate change, either by promoting more frequent and/or severe fires or by reducing the number of fire events due to the limitation of fuel load. Predicting the future of fire-driven ecosystems is a complex task as species' survival depends on many factors that vary in space and time. Since plants are constantly experiencing new environments as they grow through meristem development, woody plant modularity, modules morpho-physiological aspects and their integration should be considered when investigating species strategies in fire-prone ecosystems: according to their position and their tissue composition, plants' modules experience fire differently and will contribute differently to other modules and the whole plant survival, with consequences cascading over the overall vegetation structure. Growth modules may hold the key to understanding how fast plants can get protected from fire, ultimately helping us to predict which species will persist across changing fire regimes. We present an empirical example showing how different fire-return intervals translate into distinct pressures on the timing, protection and location of modules, and discuss how these can translate into modifications in the vegetation structure due to climate change., Competing Interests: None declared., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company.)

Background and Aims: Bark allows species to survive fire, protecting their inner tissues and allowing new branches to resprout from aerial buds. Thus, bark production is likely to be selected with aerial bud protection in fire-prone ecosystems. By considering the coexistence of fire-prone and fire-free ecosystems, in addition to the different impacts of flames on different growth forms, in this study we tested whether: (1) species from areas with higher fire frequencies have a faster bark production; (2) bark growth rate differs between trees and shrubs; (3) generalists adjust their bark production according to their environment (fire-prone or fire-free ecosystems); and (4) fast bark production results in better aerial bud protection., Methods: We sampled two different types of forests and savannas in the Cerrado and registered every woody individual with height between 1.5 and 3 m tall (directly exposed to the flames). For the 123 species registered, we sampled three different individuals in each vegetation type where the species occurred to assess their bark production and aerial bud protection. We then checked, for each species, their preferred habitat (savanna and forest specialists or generalists) and their predominant growth form., Key Results: A minimal threshold of 0.13 mm per growth unit of bark production differentiated woody communities from savannas and forests. Shrubs and trees did not differ in terms of bark growth rate, despite being exposed to the flames in a different manner. Generalist species in savannas were able to produce bark above the threshold. However, when these species were in forests they produced bark below the threshold. Finally, a higher bark growth rate accounted for a better aerial bud protection., Conclusions: Generalist species are likely to be capable of displaying plasticity in their bark production, which could be important for their success in contrasting ecosystems. The relationship between aerial bud protection and bark growth rate suggests that bark production plays an important role in protecting the dormant buds, in addition to being selected in fire-prone ecosystems., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

If trees minimize self-shading, new foliage in shaded parts of the crown should remain minimal. However, many species have abundant foliage on short shoots inside their crown. In this paper, we test the hypothesis that short shoots allow trees to densify their foliage in self-shaded parts of the crown thanks to reduced costs. Using 30 woody species in Mediterranean and tropical biomes, we estimated the contribution of short shoots to total plant foliage, calculated their costs relative to long shoots including wood cost and used 3D plant simulations calibrated with field measurements to quantify their light interception, self-shading and yield. In species with short shoots, leaves on short shoots account for the majority of leaf area. The reduced cost of short stems enables the production of leaf area with 36% less biomass. Simulations show that although short shoots are more self-shaded, they benefit the plant because they cost less. Lastly, the morphological properties of short shoots have major implications for whole plant architecture. Taken together, our results question the validity of only assessing leaf costs to understand leaf economics and call for more integrated observations at the crown scale to understand light capture strategies in woody plants., (© 2022 The Authors New Phytologist © 2022 New Phytologist Foundation.)

Background and Aims: The defensive role of spines has previously been related to leaves, young shoots and reproductive organs. However, some woody species harbour spines on their trunks where none of those organs are present. Several explanations are plausible: they could be (1) climbing aids, (2) remnants from defence of leaves or reproductive organs during an earlier development phase, or (3) an as-yet undescribed defence. Here we investigate whether they could play a role against either bark feeding or preventing climbing animals accessing food resources in the tree canopy., Methods: We described 31 woody species with spines on their trunk, growing in a botanical garden, to test whether morphological strategies could be identified and suggest what could be their most likely function. As testing their function is difficult experimentally for large pools of species, we performed virtual experiments to evaluate the potential roles of trunk spines against bark removal and climbing animals of different sizes. We then compared for each species and their confamilial non-spiny species the nutritional profiles of leaf, bark and reproductive organs to test whether trunk spines were associated with a nutritious organ (more likely targeted by herbivores)., Key Results: We identified four morphological syndromes of trunk spines. Two corresponded to already known functions (anchorage for lianas and crown defence against large ground mammals), and two strategies are newly described trait syndromes with traits suggesting a defence against bark feeding and climbing mammals. By simulation, we show how each strategy could translate into defence against debarking and prevent herbivores from climbing., Conclusions: We identified trunk spine strategies and the criteria to classify them, their most likely function and the likely feeding mode and size of animal against which different trunk spine strategies may be effective. We discuss further perspectives for testing their function and their ecological significance., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Plant architecture strongly influences ecological performance, yet its role in plant evolution has not been explored in depth. By testing both phylogenetic and environmental signals, it is possible to separate architectural traits into four categories: development constraints (phylogenetic signal only); convergences (environmental dependency only); key confluences to the environmental driver (both); unknown (neither). We analysed the evolutionary history of the genus Euphorbia, a model clade with both high architectural diversity and a wide environmental range. We conducted comparative analyses of 193 Euphorbia species world-wide using 73 architectural traits, a dated phylogeny, and climate data. We identified 14 architectural types in Euphorbia based on trait combinations. We found 22 traits and three types representing convergences under climate groups, 21 traits and four types showing phylogenetic signal but no relation to climate, and 16 traits and five types with both climate and phylogenetic signals. Major drivers of architectural trait evolution likely include water stress in deserts (selected for succulence, continuous branching), frost disturbance in temperate systems (selected for simple, prostrate, short-lived shoots) and light competition (selected for arborescence). Simple architectures allowed resilience to disturbance, and frequent transitions into new forms. Complex architectures with functional specialisation developed under stable climates but have low evolvability., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

Premise: To further advance the understanding of the species-rich, economically and ecologically important angiosperm order Myrtales in the rosid clade, comprising nine families, approximately 400 genera and almost 14,000 species occurring on all continents (except Antarctica), we tested the Angiosperms353 probe kit., Methods: We combined high-throughput sequencing and target enrichment with the Angiosperms353 probe kit to evaluate a sample of 485 species across 305 genera (76% of all genera in the order)., Results: Results provide the most comprehensive phylogenetic hypothesis for the order to date. Relationships at all ranks, such as the relationship of the early-diverging families, often reflect previous studies, but gene conflict is evident, and relationships previously found to be uncertain often remain so. Technical considerations for processing HTS data are also discussed., Conclusions: High-throughput sequencing and the Angiosperms353 probe kit are powerful tools for phylogenomic analysis, but better understanding of the genetic data available is required to identify genes and gene trees that account for likely incomplete lineage sorting and/or hybridization events., (© 2021 The Authors. American Journal of Botany published by Wiley Periodicals LLC on behalf of Botanical Society of America.)

Ideas on hominin evolution have long invoked the emergence from forests into open habitats as generating selection for traits such as bipedalism and dietary shifts. Though controversial, the savanna hypothesis continues to motivate research into the palaeo-environments of Africa. Reconstruction of these ancient environments has depended heavily on carbon isotopic analysis of fossil bones and palaeosols. The sparsity of the fossil record, however, imposes a limit to the strength of inference that can be drawn from such data. Time-calibrated phylogenies offer an additional tool for dating the spread of savanna habitat. Here, using the evolutionary ages of African savanna trees, we suggest an initial tropical or subtropical expansion of savanna between 10 and 15 Ma, which then extended to higher latitudes, reaching southern Africa ca. 3 Ma. Our phylogenetic estimates of the origin and latitudinal spread of savannas broadly correspond with isotopic age estimates and encompass the entire hominin fossil record. Our results are consistent with the savanna hypothesis of early hominin evolution and reignite the debate on the drivers of savanna expansion. Our analysis demonstrates the utility of phylogenetic proxies for dating major ecological transitions in geological time, especially in regions where fossils are rare or absent or occur in discontinuous sediments.

Background and Aims: Herbivory by large mammals imposes a critical recruitment bottleneck on plants in many systems. Spines defend plants against large herbivores, and how early they emerge in saplings may be one of the strongest predictors of sapling survival in herbivore-rich environments. Yet little effort has been directed at understanding the variability in spine emergence across saplings., Methods: We present a multispecies study examining whether and how sapling size, spine type and species' environmental niche (light and precipitation environment) influence early emergence and biomass investment in spines. A phylogenetically diverse pool of 45 species possessing different spine types (spines, prickles and thorns; that are derived from distinct plant organs: leaf, epidermis or cortex, and branch, respectively), were grown under common-garden conditions, and patterns of spine emergence and biomass allocation to spines at 5 and 15 weeks after transplanting were characterized., Key Results: Spine type and species' resource niche were the main factors driving early emergence and investment patterns. Spines emerged earliest in leaf spine-bearing species, and latest in thorn-bearing species. The probability of early spine emergence increased with decreasing precipitation, and was greater in species from open than from closed habitats. Sapling investment in spines changed with plant mass but was contingent on spine type and habitat type., Conclusions: Different spine types have strikingly different timing of expression, suggesting that developmental origins of spines play a critical role in sapling defences. Furthermore, species from different precipitation and light environments (open vs. closed habitats) showed contrasting patterns of early spine expression, suggesting that resource limitation in their native range may have driven divergent evolution of early defence expression., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Tropical savannas have a ground cover dominated by C 4 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 CO 2 . 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., (© 2018 The Authors New Phytologist © 2018 New Phytologist Trust.)

Shade cast by trees, which suppresses grass growth, and fire fuelled by grass biomass, which prevents tree sapling establishment, are mutually exclusive and self-reinforcing drivers of biome distribution in savanna-forest mosaics. We investigated how shade depth, represented by canopy leaf area index (LAI), is generated by adult trees across savanna-forest boundaries and how a shade gradient filters tree functioning, and grass composition and biomass. Forest trees exerted greater shading through increased stem density and greater light interception per unit biomass. A critical transition at LAI c. 1.5 was linked to tree shifts from savanna to forest species, functional shifts from fire-tolerant to light-competitive species, and grass composition shifts from C 4 to C 3 pathways. A second transition to grass fuel loads too low to support fires, occurred at a lower canopy density (LAI > 0.5), accompanied by shifts in C 4 subtype dominance. This pattern suggests that shade suppression of grass biomass is an essential first step for the maintenance of alternative stable states., (© 2018 The Authors. New Phytologist © 2018 New Phytologist Trust.)

Disease emergence events regularly result from human activities such as agriculture, which frequently brings large populations of genetically uniform hosts into contact with potential pathogens. Although viruses cause nearly 50% of emerging plant diseases, there is little systematic information about virus distribution across agro-ecological interfaces and large gaps in understanding of virus diversity in nature. Here we applied a novel landscape-scale geometagenomics approach to examine relationships between agricultural land use and distributions of plant-associated viruses in two Mediterranean-climate biodiversity hotspots (Western Cape region of South Africa and Rhône river delta region of France). In total, we analysed 1725 geo-referenced plant samples collected over two years from 4.5 × 4.5 km 2 grids spanning farmlands and adjacent uncultivated vegetation. We found substantial virus prevalence (25.8-35.7%) in all ecosystems, but prevalence and identified family-level virus diversity were greatest in cultivated areas, with some virus families displaying strong agricultural associations. Our survey revealed 94 previously unknown virus species, primarily from uncultivated plants. This is the first effort to systematically evaluate plant-associated viromes across broad agro-ecological interfaces. Our findings indicate that agriculture substantially influences plant virus distributions and highlight the extent of current ignorance about the diversity and roles of viruses in nature.

Savannas first began to spread across Africa during the Miocene. A major hypothesis for explaining this vegetation change is the increase in C4 grasses, promoting fire. We investigated whether mammals could also have contributed to savanna expansion by using spinescence as a marker of mammal herbivory. Looking at the present distribution of 1,852 tree species, we established that spinescence is mainly associated with two functional types of mammals: large browsers and medium-sized mixed feeders. Using a dated phylogeny for the same tree species, we found that spinescence evolved at least 55 times. The diversification of spiny plants occurred long after the evolution of Afrotherian proboscideans and hyracoids. However, it is remarkably congruent with diversification of bovids, the lineage including the antelope that predominantly browse these plants today. Our findings suggest that herbivore-adapted savannas evolved several million years before fire-maintained savannas and probably, in different environmental conditions. Spiny savannas with abundant mammal herbivores occur in drier climates and on nutrient-rich soils, whereas fire-maintained savannas occur in wetter climates on nutrient-poor soils., Competing Interests: The authors declare no conflict of interest.

Tropical savannas are hypothesized to be hot spots of nitrogen-fixer diversity and activity because of the high disturbance and low nitrogen characteristic of savanna landscapes. Here we compare the abundances of nitrogen-fixing and non-fixing trees in both tropical savannas and tropical forests under climatically equivalent conditions, using plant inventory studies across 566 plots in South America and Africa. A single factor, aridity, explained 19-54% of the variance in fixer abundance, and unexpectedly was more important than fire frequency, biome, and continent. Nitrogen fixers were more abundant in arid environments; as a result, African savannas, which tend to be drier, were richer in nitrogen fixers than South American savannas. Fixer abundance converged on similar levels in forests in both continents. We conclude that climate plays a greater role than fire in determining the distribution of nitrogen fixers across tropical savanna and forest biomes., (© 2016 by the Ecological Society of America.)

Contrasting fire regimes maintain patch mosaics of savanna, thicket and forest biomes in many African subtropical landscapes. Species dominating each biome are thus expected to display distinct fire-related traits, commonly thought to be bark related. Recent Australian savanna research suggests that bud position, not bark protection alone, determines fire resilience via resprouting. We tested first how bud position influences resprouting ability in 17 tree species. We then compared the effect of both bark-related protection and bud position on the distribution of 63 tree species in 253 transects in all three biomes. Tree species with buds positioned deep under bark had a higher proportion of post-fire aboveground shoot resprouting. Species with low bud protection occurred in fire-prone biomes only if they could root-sucker. The effect of bud protection was supported by a good relationship between species bud protection and distribution across a gradient of fire frequency. Bud protection and high bark production are required to survive frequent fires in savanna. Forests are fire refugia hosting species with little or no bud protection and thin bark. Root-suckering species occur in the three biomes, suggesting that fire is not the only factor filtering this functional type., (© 2015 The Authors. New Phytologist © 2015 New Phytologist Trust.)

In the Neotropics, the genus Hydrangea of the popular ornamental hortensia family is represented by climbing species that strongly cling to their support surface by means of adhesive roots closely positioned along specialized anchoring stems. These root-climbing hortensia species belong to the nearly exclusive American Hydrangea section Cornidia and generally are long lianescent climbers that mostly flower and fructify high in the host tree canopy. The Mexican species Hydrangea seemannii, however, encompasses not only long lianescent climbers of large vertical rock walls and coniferous trees, but also short 'shrub-like' climbers on small rounded boulders. To investigate growth form plasticity in root-climbing hortensia species, we tested the hypothesis that support variability (e.g. differences in size and shape) promotes plastic responses observable at the mechanical, structural and anatomical level. Stem bending properties, architectural axis categorization, tissue organization and wood density were compared between boulder and long-vertical tree-climbers of H. seemannii. For comparison, the mechanical patterns of a closely related, strictly long-vertical tree-climbing species were investigated. Hydrangea seemannii has fine-tuned morphological, mechanical and anatomical responses to support variability suggesting the presence of two alternative root-climbing strategies that are optimized for their particular environmental conditions. Our results suggest that variation of some stem anatomical traits provides a buffering effect that regulates the mechanical and hydraulic demands of two distinct plant architectures. The adaptive value of observed plastic responses and the importance of considering growth form plasticity in evolutionary and conservation studies are discussed., (© 2014 The Author(s) Published by the Royal Society. All rights reserved.)