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Narratives of landscape degradation are often linked to unsustainable fire use by local communities. Madagascar is a case in point: the island is considered globally exceptional, with its remarkable endemic biodiversity viewed as threatened by unsustainable anthropogenic fire. Yet, fire regimes on Madagascar have not been empirically characterised or globally contextualised. Here, we contribute a comparative approach to determining relationships between regional fire regimes and global patterns and trends, applied to Madagascar using MODIS remote sensing data (2003–2019). Rather than a global exception, we show that Madagascar's fire regimes are similar to 88% of tropical burned area with shared climate and vegetation characteristics, and can be considered a microcosm of most tropical fire regimes. From 2003–2019, landscape‐scale fire declined across tropical grassy biomes (17%–44% excluding Madagascar), and on Madagascar at a relatively fast rate (36%–46%). Thus, high tree loss anomalies on the island (1.25–4.77× the tropical average) were not explained by any general expansion of landscape‐scale fire in grassy biomes. Rather, tree loss anomalies centred in forests, and could not be explained by landscape‐scale fire escaping from savannas into forests. Unexpectedly, the highest tree loss anomalies on Madagascar (4.77×) occurred in environments without landscape‐scale fire, where the role of small‐scale fires (<21 h [0.21 km2]) is unknown. While landscape‐scale fire declined across tropical grassy biomes, trends in tropical forests reflected important differences among regions, indicating a need to better understand regional variation in the anthropogenic drivers of forest loss and fire risk. Our new understanding of Madagascar's fire regimes offers two lessons with global implications: first, landscape‐scale fire is declining across tropical grassy biomes and does not explain high tree loss anomalies on Madagascar. Second, landscape‐scale fire is not uniformly associated with tropical forest loss, indicating a need for socio‐ecological context in framing new narratives of fire and ecosystem degradation. [ABSTRACT FROM AUTHOR]

Woody encroachment is ubiquitous in grassy ecosystems worldwide, but its global impacts on the diversity of herbaceous plants that characterise and define these ecosystems remain unquantified. The pervasiveness of encroachment is relatively easily observed via remote sensing, but its impacts on plant diversity and richness below the canopy can only be observed via field‐based studies. Via a meta‐analysis of 42 field studies across tropical to temperate grassy ecosystems, we quantified how encroachment altered herbaceous species richness, and the richness of forbs, C3 graminoids and C4 graminoids. Across studies, the natural logarithm of the response ratio (lnRR) of herbaceous species richness ranged from −3.33 to 0.34 with 87% of encroached ecosystems negatively impacted. Assessment of the extent of encroachment, duration of encroachment, mean annual rainfall, latitude, and continent demonstrated that only extent of encroachment had relevance in the data (univariate model including a random effect of study explained 45.4% of variance). The global weighted mean lnRR of species richness decreased from −0.245 at <33% of woody cover increase, to −0.562 at 33%–66%, and to −0.962 at >66%. Continued encroachment results in substantial loss of herbaceous diversity at medium and high extents, with a loss of richness that is not replaced. Although all functional groups are significantly negatively impacted by encroachment, forb richness is relatively more sensitive than graminoid richness, and C4 graminoid richness relatively more than C3 graminoid richness. Although no geographic or climatic correlates had relevance in the data, encroachment as an emergent product of global change coalesces to decrease ground layer light availability, lead to loss of fire and grazers, and alter hydrology and soils. Encroachment is accelerating and grassy ecosystems require urgent attention to determine critical woody cover thresholds that facilitate diverse and resilient grassy ecosystems. [ABSTRACT FROM AUTHOR]

Ziziphus (Rhamnaceae) is a widely distributed genus across the Australasian and African tropics with unusual diversity in habit, and many species of significance to people. Here, we quantify the environmental limits of Ziziphus species and examine inter‐specific relationships among functional traits, environment, biome, and range size. We developed a curated geolocation database for Ziziphus and used it to examine the environmental limits of the genus relative to temperature, rainfall, and seasonality. To assess the relationship between biome and habit, permutational analysis of variance was used, while hierarchical clustering was used to determine whether habit, leaves, and fruit traits were related to biome. For 40 species with adequate geolocation data, range size was calculated to assess its relationship with habit, biomes, and cultivation. Finally, niche identity tests were used to determine niche equivalency among cultivated and non‐cultivated species. Liana species are restricted to closed forests and the geoxylic habit is found only in open grasslands. Further, habit is significantly associated with range size, with trees having on average larger range sizes than shrubs, lianas, and geoxyles, but biome was not correlated with range size. Cultivated species have ranges ~10 times that of non‐cultivated tree species and with significantly different and broader environmental niches. The unusually wide distribution of Ziziphus can be explained by its diversity of habits associated with different biomes spanning continents. This, along with the usage of many Ziziphus species by people for their fruits, expands the range and environmental occupation of the genus. [ABSTRACT FROM AUTHOR]

Frequent grazing can establish high forage value grazing lawns supporting high grazer densities, but can also produce overgrazed grass communities with unpalatable or low grass basal cover, supporting few grazers. Attempts to create grazing lawns via concentrated grazing, with a goal to increase grazer numbers, are thus risky without knowing how environmental conditions influence the likelihood of each outcome. We collected grass species and trait data from 33 frequently grazed grass communities across eastern South Africa (28 sites) and the Serengeti National Park, Tanzania (five sites), covering wide rainfall (336–987 mm year−1) and soil (e.g., 44%–93% sand) gradients. We identified four grass growth forms using hierarchical clustering on principal components analyses of trait data and assessed trait–environment and growth form–environment relationships using fourth corner and principal components analyses. We distinguished two palatable grass growth forms that both attract yet resist grazers and comprise grazing lawns: (1) "lateral attractors" that spread vegetatively via stolons and rhizomes, and (2) "tufted attractors" that form isolated tufts and may have alternate tall growth forms. By contrast, (3) tough, upright, tufted "resisters," and (4) "avoiders" with sparse architectures or that grow appressed to the soil surface, are of little forage value and avoided by grazers. Grazing lawns occurred across a wide range of conditions, typically comprising lateral attractor grasses in drier, sandy environments, and tufted attractor grasses in wetter, low‐sand environments. Resisters occurred on clay‐rich soils in mesic areas, while avoiders were widespread but scarce. While grazing lawns can be established under most conditions, monitoring their composition and cover is important, as the potential for overgrazing seems as widely relevant. Tufted attractor‐dominated lawns appear somewhat more vulnerable to degradation than lateral attractor‐dominated lawns. Increased avoider and resister abundance both reduce forage value, although resisters may provide better soil protection. [ABSTRACT FROM AUTHOR]

The aim of this study was to understand how communities of adult and juvenile (seedlings and saplings) woody plants were impacted by fire and the 2014–2016 El Niño drought in Kruger National Park, South Africa. We used a landscape‐scale fire experiment spanning 2013–2019 in a semi‐arid savanna in the central west of Kruger National Park (mean annual precipitation, 543 mm). Adult and juvenile woody species composition were recorded during and after the drought in 40 plots that experienced a mix of no fire, moderate fire, and frequent fire treatments. Using multivariate modeling, we related community composition in juvenile and adult woody plants to year of sampling and the experimental fire treatments. Post‐drought, there was significant adult woody plant top‐kill, especially in dominant species Dichrostachys cinerea (81% reduction in abundance), Acacia nigrescens (30%), and Combretum apiculatum (19%), but there was no significant change in adult species richness. Two years post‐drought, abundance of all juveniles decreased by 35%, and species richness increased in juveniles in both the frequent fire (7%) and no fire treatments (32%). Counter‐intuitively, the El Niño drought increased species richness of the woody plant community due to the recruitment of new species as juveniles, a potential lasting impact on diversity, and where different fire regimes were associated with differences in community composition. Drought events in semi‐arid savannas could drive temporal dynamics in species richness and composition in previously unrecognized ways. [ABSTRACT FROM AUTHOR]

Species of the Heteropogon‐Themeda clade are ecologically important grasses distributed across the tropics, including widespread species, such as the pantropical Heteropogon contortus and Themeda triandra, and range‐restricted species such as Heteropogon ritchiei and Themeda anathera. Here, we examine habitat preferences of the grassland/savanna and wetland species by describing bioclimatic niche characteristics, characterizing functional traits, and investigating the evolution of functional traits of 31 species in the Heteropogon‐Themeda clade in relation to precipitation and temperature. The climatic limits of the clade are linked to mean annual precipitation and seasonality that also distinguish seven wetland species from 24 grassland/savanna species. Tests of niche equivalency highlighted the unique bioclimatic niche of the wetland species. However, climatic factors do not fully explain species geographic range, and other factors are likely to contribute to their distribution ranges. Trait analyses demonstrated that the wetland and grassland/savanna species were separated by culm height, leaf length, leaf area, awn length, and awn types. Phylogenetic analyses showed that the wetland species had tall stature with long and large leaves and lack of hygroscopic awns, which suggest selective pressures in the shift between savanna/grassland and wetland. The two most widespread species, H. contortus and T. triandra, have significantly different bioclimatic niches, but we also found that climatic niche alone does not explain the current geographic distributions of H. contortus and T. triandra. Our study provides a new understanding of the biogeography and evolutionary history of an ecologically important clade of C4 tropical grasses. [ABSTRACT FROM AUTHOR]

Aim: To understand the geographical distribution of grasses in sub-Saharan Africa with reference to key plant traits thought to affect range size in this family (Poaceae). Specifically, to test hypotheses on the importance of plant height and lifespan in determining range size and invasion potential in the context of their evolutionary history. Location: Sub-Saharan Africa. Taxon: Poaceae. Methods: The range sizes of 757 grass species native to southern Africa were estimated for the sub-Saharan African region from geo-referenced herbarium records using the alpha hull function. Phylogenetic generalised least squares models and linear mixed effects models were fitted to test whether grass range size was related to plant height and lifespan. Tribe-level relationships between range size and plant height were assessed with linear models. For species introduced to other continents, generalised linear mixed effects models were fitted to test whether invasiveness was related to native range size, plant height and lifespan. Differences in native range size among species in four invasion-related categories were assessed with linear mixed effects models. Results: Grass range sizes are larger for taller species and for species with shorter lifespans. The relationship between plant height and range size varies widely among tribes, with some range-restricted tribes having a non-significant effect on plant height. Grasses with larger native range sizes and shorter lifespans are more likely to become invasive after being introduced to other continents. Grass species introduced to other continents have larger native range sizes than those that have not, and native range size increases along the introduced-naturalised-invasive continuum. Main conclusions: The increased dispersal opportunities of annual-biannual grasses appear to have a greater positive effect on range size than do the longer generation times of perennial grasses. Grass height has and continues to be an important driver of grass biogeography, with implications for understanding the spread of certain grass tribes over the Miocene. Factors that promote large native range sizes are also likely to increase the probability of a species becoming invasive. [ABSTRACT FROM AUTHOR]

Summary: Plant populations persist under recurrent fire via resprouting from surviving tissues (resprouters) or seedling recruitment (seeders). Woody species are inherently slow maturing, meaning that seeders are confined to infrequent fire regimes. However, for grasses, which mature faster, the relationships between persistence strategy and fire regime remain unknown.Globally, we analysed associations between fire regimes experienced by hundreds of grass species and their persistence strategy, within a phylogenetic context. We also tested whether persistence strategies are associated with morphological and physiological traits.Resprouters were associated with less frequent fire than seeders. Whilst modal fire frequencies were similar (fire return interval of 4–6 yr), seeders were restricted to regions with more frequent fire than resprouters, suggesting that greater competition with long‐lived resprouters restricts seeder recruitment and survival when fire is rare. Resprouting was associated with lower leaf N, higher C:N ratios and the presence of belowground buds, but was unrelated to photosynthetic pathway.Differences between the life histories of grasses and woody species led to a contrasting prevalence of seeders and resprouters in relation to fire frequency. Rapid sexual maturation in grasses means that seeder distributions, relative to fire regime, are determined by competitive ability and recruitment, rather than time to reproductive maturity. See also the Commentary on this article by Day, 230: 406‐407. [ABSTRACT FROM AUTHOR]

Aim: In savannas, a grass‐dominated ground layer is key to ecosystem function via grass–fire feedbacks that maintain open ecosystems. With woody encroachment, tree density increases, thereby decreasing light in the ground layer and potentially altering ecosystem function. We investigated how light availability can filter individual grass species distributions and whether different functional traits are associated with response to a shade gradient in a landscape experiencing woody encroachment. Location: Savanna–forest mosaic in the Cerrado domain, southeastern Brazil. Methods: Along an encroachment gradient of increasing tree leaf area index (LAI) and shade, we determined how changing light availability alters grass diversity and ground layer structure relative to grass cover and grass functional traits (photosynthetic pathway, underground storage organs, bud protection and traits related to grass shape, size and leaf dimensions). Results: Increasing shade led to a decrease in grass cover and grass species richness, and also compositional and functional changes. We found that where tree LAI reached 1, grass cover was reduced by 50% and species richness by 30%. While C4 grass species abundances decreased with increasing shade, the opposite pattern was true for C3 grasses. There were only small differences in light preferences among C4 subtypes, with phosphoenolpyruvate carboxykinase (PCK) species tolerating slightly more shaded conditions. Persistence of some C4 species under more shaded conditions was possible, likely due to an ability to store starch reserves via underground storage organs. Conclusions: Woody encroachment changes diversity and structure of the grassy layer that is critical to the functioning of savanna ecosystems, highlighting the dependence of the diverse grass layer on open and sunny conditions. Our results suggest a threshold of tree cover close to LAI ≈ 1 as being critical to cerrado grassy layer conservation. [ABSTRACT FROM AUTHOR]

Summary: Process‐based vegetation models attempt to represent the wide range of trait variation in biomes by grouping ecologically similar species into plant functional types (PFTs). This approach has been successful in representing many aspects of plant physiology and biophysics but struggles to capture biogeographic history and ecological dynamics that determine biome boundaries and plant distributions. Grass‐dominated ecosystems are broadly distributed across all vegetated continents and harbour large functional diversity, yet most Land Surface Models (LSMs) summarise grasses into two generic PFTs based primarily on differences between temperate C3 grasses and (sub)tropical C4 grasses. Incorporation of species‐level trait variation is an active area of research to enhance the ecological realism of PFTs, which form the basis for vegetation processes and dynamics in LSMs. Using reported measurements, we developed grass functional trait values (physiological, structural, biochemical, anatomical, phenological, and disturbance‐related) of dominant lineages to improve LSM representations. Our method is fundamentally different from previous efforts, as it uses phylogenetic relatedness to create lineage‐based functional types (LFTs), situated between species‐level trait data and PFT‐level abstractions, thus providing a realistic representation of functional diversity and opening the door to the development of new vegetation models. [ABSTRACT FROM AUTHOR]

Summary: Uapaca bojeri Baill. (Phyllanthaceae), is a tree species endemic to Madagascar, colloquially known as 'tapia', that forms monodominant grassy woodlands in a series of disconnected populations across the Central Highlands. Uapaca bojeri and the ecosystems it forms are culturally, economically and scientifically important to Madagascar. The species, Uapaca bojeri, and the ecosystems it dominates, are in need of new research and conservation efforts to understand how sustainable use of the species can align with its conservation and that of the unique plants and animals special to the tapia woodlands. [ABSTRACT FROM AUTHOR]

Aim: Biomes worldwide are shifting with global change. Biomes whose extents are limited by temperature or precipitation, such as the tundra and savanna, may be particularly strongly affected by climate change. While woody plant encroachment is prevalent across both biomes, its relationship to temperature and precipitation change remains unknown. Here, we quantify the degree to which woody encroachment is related to climate change and identify its main associated drivers. Location: Tundra and savanna biomes. Time period: 1992 ± 20.27–2010 ± 5.62 (mean ± SD). 1876–2016 (range). Major taxa studied: Woody plants (shrubs and trees). Methods: We compiled a dataset comprising 1,089 records from 899 sites of woody plant cover over time and attributed drivers of woody cover change across these two biomes. We calculated cover change in each biome and assessed the degree to which cover change corresponds to concurrent temperature and precipitation changes using multiple climate metrics. Finally, we conducted a quantitative literature review of the relative importance of attributed drivers of woody cover change. Results: Woody encroachment was widespread geographically and across climate gradients. Rates of woody cover change (positive or negative) were 1.8 times lower in the tundra than in the savanna (1.8 vs. 3.2%), while rates of woody cover increase (i.e., encroachment) were c. 1.7 times lower in the tundra compared with the savanna (3.7 vs. 6.3% per decade). In the tundra, magnitudes of woody cover change did not correspond to climate, while in the savanna, greater cover change corresponded with increases in precipitation. We found higher rates of woody cover change in wetter versus drier sites with warming in the tundra biome, and higher rates of woody cover change in drier versus wetter sites with increasing precipitation in the savanna. However, faster rates of woody cover change were not associated with more rapid rates of climate change across sites, except for maximum precipitation in the savanna. Main conclusions: Woody encroachment was positively related to warming in the tundra and increased rainfall in the savanna. However, cover change rates were not predicted by rates of climate change, which can be partially explained by climate interactions in both biomes. Additional likely influences include site‐level factors, time‐lags, plant‐specific responses, and land use and other non‐climate drivers. Our findings highlight the complex nature of climate change impacts in biomes limited by seasonality, which should be accounted for to realistically estimate future responses across open biomes under global change scenarios. [ABSTRACT FROM AUTHOR]

Aim: The sorting of functional traits along environmental gradients is an important driver of community and landscape scale patterns of functional diversity. However, the significance of environmental factors in driving functional gradients within biomes and across continents remains poorly understood. Here, we evaluate the relationship of soil nutrients and climate to leaf traits in grasses (Poaceae) that are hypothesized to reflect different strategies of resource use along gradients of resource availability. Location: Global. Taxon: Poaceae. Methods: We made direct measurements on herbarium specimens to compile a global dataset of functional traits and realized environmental niche for 279 grass species that are common in grassland and savanna biomes. We examined the strength and direction of correlations between pairwise trait combinations and measured the distribution of traits in relation to gradients of soil properties and climate, while accounting for phylogenetic relatedness. Results: Leaf trait variation among species follows two orthogonal axes. One axis represents leaf size and plant height, and we showed positive scaling relationships between these size‐related traits. The other axis corresponds to economic traits associated with resource acquisition and allocation, including leaf tensile strength (LTS), specific leaf area (SLA) and leaf nitrogen content (LNC). Global‐scale variation in LNC was primarily correlated with soil nutrients, while LTS, SLA and size‐related traits showed weak relationships to environment. However, most of the trait variation occurred within different vegetation types, independent of large‐scale environmental gradients. Main conclusions: Our work provides evidence among grasses for relationships at the global scale between leaf economic traits and soil fertility, and for an influence of aridity on traits related to plant size. However, large unexplained variance and strong phylogenetic signal in the model residuals imply that at this scale the evolution of functional traits is driven by factors beyond contemporary environmental or climatic conditions. [ABSTRACT FROM AUTHOR]

Plant growth rates strongly determine ecosystem productivity and are a central element of plant ecological strategies. For laboratory and glasshouse‐grown seedlings, specific leaf area (SLA; ratio of leaf area to mass) is a key driver of interspecific variation in growth rate (GR). Consequently, SLA is often assumed to drive GR variation in field‐grown adult plants. However, there is an increasing evidence that this is not the general case. This suggests that GR – SLA relationships (and perhaps those for other traits) may vary depending on the age or size of the plants being studied. Here we investigated GR – trait relationships and their size dependence among 17 woody species from an open‐canopy, fire‐prone savanna in northern Australia. We tested the predictions that SLA and stem diameter growth rate would be positively correlated in saplings but unrelated in adults while, in both age classes, faster‐GR species would have higher light‐saturated photosynthetic rate (Asat), higher leaf nutrient concentrations, higher branch‐scale biomass allocation to leaf versus stem tissues and lower wood density (WD). SLA showed no relationship to stem diameter GR, even in saplings, and the same was true of leaf N and P concentrations, and WD. However, branch‐scale leaf:stem allocation was strongly related to GR in both age groups, as was Asat. Together, these two traits accounted for up to 80% of interspecific variation in adult GR, and 41% of sapling GR. Asat is rarely measured in field‐based GR studies, and this is the first report of branch‐scale leaf:stem allocation (analogous to a benefit:cost ratio) in relation to plant growth rate. Our results suggest that we may yet find general trait‐drivers of field growth rates, but SLA will not be one. [ABSTRACT FROM AUTHOR]

Summary: 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. [ABSTRACT FROM AUTHOR]

ABSTRACT: Poaceae (the grasses) is arguably the most successful plant family, in terms of its global occurrence in (almost) all ecosystems with angiosperms, its ecological dominance in many ecosystems, and high species richness. We suggest that the success of grasses is best understood in context of their capacity to colonize, persist, and transform environments (the “Viking syndrome”). This results from combining effective long‐distance dispersal, efficacious establishment biology, ecological flexibility, resilience to disturbance and the capacity to modify environments by changing the nature of fire and mammalian herbivory. We identify a diverse set of functional traits linked to dispersal, establishment and competitive abilities. Enhanced long‐distance dispersal is determined by anemochory, epizoochory and endozoochory and is facilitated via the spikelet (and especially the awned lemma) which functions as the dispersal unit. Establishment success could be a consequence of the precocious embryo and large starch reserves, which may underpin the extremely short generation times in grasses. Post‐establishment genetic bottlenecks may be mitigated by wind pollination and the widespread occurrence of polyploidy, in combination with gametic self‐incompatibility. The ecological competitiveness of grasses is corroborated by their dominance across the range of environmental extremes tolerated by angiosperms, facilitated by both C3 and C4 photosynthesis, well‐developed frost tolerance in several clades, and a sympodial growth form that enabled the evolution of both annual and long‐lived life forms. Finally, absence of investment in wood (except in bamboos), and the presence of persistent buds at or below ground level, provides tolerance of repeated defoliation (whether by fire, frost, drought or herbivores). Biotic modification of environments via feedbacks with herbivory or fire reinforce grass dominance leading to open ecosystems. Grasses can be both palatable and productive, fostering high biomass and diversity of mammalian herbivores. Many grasses have a suite of architectural and functional traits that facilitate frequent fire, including a tufted growth form, and tannin‐like substances in leaves which slow decomposition. We mapped these traits over the phylogeny of the Poales, spanning the grasses and their relatives, and demonstrated the accumulation of traits since monocots originated in the mid‐Cretaceous. Although the sympodial growth form is a monocot trait, tillering resulting in the tufted growth form most likely evolved within the grasses. Similarly, although an ovary apparently constructed of a single carpel evolved in the most recent grass ancestor, spikelets and the awned lemma dispersal units evolved within the grasses. Frost tolerance and C4 photosynthesis evolved relatively late (late Palaeogene), and the last significant trait to evolve was probably the production of tannins, associated with pyrophytic savannas. This fits palaeobotanical data, suggesting several phases in the grass success story: from a late Cretaceous origin, to occasional tropical grassland patches in the later Palaeogene, to extensive C3 grassy woodlands in the early–middle Miocene, to the dramatic expansion of the tropical C4 grass savannas and grasslands in the Pliocene, and the C3 steppe grasslands during the Pleistocene glacial periods. Modern grasslands depend heavily on strongly seasonal climates, making them sensitive to climate change. [ABSTRACT FROM AUTHOR]

Tree phenology mediates land-atmosphere mass and energy exchange and is a determinant of ecosystem structure and function. In the dry tropics, including African savannas, many trees grow new leaves during the dry season - weeks or months before the rains typically start. This syndrome of pre-rain green-up has long been recognized at small scales, but the high spatial and interspecific variability in leaf phenology has precluded regional generalizations., We used remote sensing data to show that this precocious phenology is ubiquitous across the woodlands and savannas of southern tropical Africa., In 70% of the study area, green-up preceded rain onset by > 20 d (42% > 40 d). All the main vegetation formations exhibited pre-rain green-up, by as much as 53 ± 18 d (in the wet miombo). Green-up showed low interannual variability (SD between years = 11 d), and high spatial variability (> 100 d)., These results are consistent with a high degree of local phenological adaptation, and an insolation trigger of green-up. Tree-tree competition and niche separation may explain the ubiquity of this precocious phenology. The ubiquity of pre-rain green-up described here challenges existing model representations and suggests resistance (but not necessarily resilience) to the delay in rain onset predicted under climate change. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

1. 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. 2. 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. 3. 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. 4. 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. [ABSTRACT FROM AUTHOR]

Adaptation to changing environments often requires novel traits, but how such traits directly affect the ecological niche remains poorly understood. Multiple plant lineages have evolved C4 photosynthesis, a combination of anatomical and biochemical novelties predicted to increase productivity in warm and arid conditions. Here, we infer the dispersal history across geographical and environmental space in the only known species with both C4 and non-C4 genotypes, the grass Alloteropsis semialata. While non-C4 individuals remained confined to a limited geographic area and restricted ecological conditions, C4 individuals dispersed across three continents and into an expanded range of environments, encompassing the ancestral one. This first intraspecific investigation of C4 evolutionary ecology shows that, in otherwise similar plants, C4 photosynthesis does not shift the ecological niche, but broadens it, allowing dispersal into diverse conditions and over long distances. Over macroevolutionary timescales, this immediate effect can be blurred by subsequent specialisation towards more extreme niches. [ABSTRACT FROM AUTHOR]

Aim We examined differences in the architecture of African and Australian savanna trees. We sought to attribute variation in tree architecture to current environments, wood density and phylogeny, and thereby elucidate the relative importance of biogeographic idiosyncrasies versus current factors in underpinning architectural differences. Location Africa and Australia. Methods We compiled canopy diameters and stem diameters from 4867 trees of 97 species and heights and stem diameters from 10,786 trees of 155 species from a range of African and Australian savanna ecosystems and climates. Using Bayesian methods we first estimated continental-scale savanna tree allometries, ignoring species differences. We then examined continental differences in species-specific allometries accounting for trait covariation using a phylogeny of our study species. Environmental variables and wood density data were included as covariates, allowing us to assess the potential underpinning of regional differences in tree allometries by differences in current environments and traits. Results Substantial allometric differences exist between Australian and African savanna trees. Australian trees are on average 6 m taller at 20 cm diameter, with a 53% smaller canopy area than African trees. However, this extreme continental-scale variation is driven by the architecture of only a few taxa in this study - Vachellia and Senegalia versus Eucalyptus and Corymbia - rather than systematic differences between species, wood density and environment. These same genera often dominate the woody strata of South African and Australian savannas, respectively. Main conclusions Stark differences in the architecture of African and Australian savanna tree taxa are not a product of environmental differences and are not consistent across species. Rather, the most likely explanation is the different evolutionary histories of African and Australian savannas, which share no woody species. We consider that these architectural differences are likely to impact regional patterns of woody biomass accumulation. [ABSTRACT FROM AUTHOR]

Aim Many tropical savannas are undergoing a trend of increasing woody biomass, or 'woody thickening'. Management to reduce fire frequency and intensity in savannas could substantially increase the amount of carbon stored in woody biomass. We addressed two questions: (1) are northern Australian savannas thickening; and (2) to what extent, and by what demographic processes, does fire affect woody biomass accumulation? Location Three large national parks, covering 24,000 km2, in monsoonal northern Australia. Methods We examined changes in woody biomass carbon stocks - inferred from tree basal area and the density of woody understorey plants - over a 10-year period in 136 savanna monitoring plots. We statistically assessed these changes in relation to fire frequency and severity. We used a meta-analysis to identify general trends in woody cover in Australian savannas over the last half-century. Results Woody biomass carbon stocks were relatively stable across the three national parks, but rates of change were statistically indistinguishable from earlier findings of a weak thickening trend. Change was negatively correlated with fire frequency, particularly the frequency of severe fires. High frequencies of severe fires decreased rates of accumulation of biomass by existing trees (through reductions in tree growth and death of individual stems), rather than whole-tree mortality and suppression of recruitment. However, across northern Australia, our meta-analysis identified a general, albeit weak, trend of woody thickening. Main conclusions The drivers of northern Australia's weak thickening trend are uncertain, but likely candidates include increasing atmospheric CO2 concentration and water availability, and pastoral intensification. We demonstrate that changes to fire management have the potential to either increase or decrease rates of woody thickening relative to any underlying trend. Understanding how savanna fires affect woody biomass, and how fire effects are mediated by climate and CO2, are essential research priorities to predict the fate of savannas. [ABSTRACT FROM AUTHOR]

Summary 1. We provide a brief overview of progress in our understanding of introduced plant species. 2. Three main conclusions emerge from our review: (i) Many lines of research, including the search for traits that make species good invaders, or that make ecosystems susceptible to invasion, are yielding idiosyncratic results. To move forward, we advocate a more synthetic approach that incorporates a range of different types of information about the introduced species and the communities and habitats they are invading. (ii) Given the growing evidence for the adaptive capacity of both introduced species and recipient communities, we need to consider the implications of the long-term presence of introduced species in our ecosystems. (iii) Several foundational ideas in invasion biology have become widely accepted without appropriate testing, or despite equivocal evidence from empirical tests. One such idea is the suggestion that disturbance facilitates invasion. 3. We use data from 200 sites around the world to provide a broad test of the hypothesis that invasions are better predicted by a change in disturbance regime than by disturbance per se. Neither disturbance nor change in disturbance regime explained more than 7% of the variation in the % of cover or species richness contributed by introduced species. However, change in disturbance regime was a significantly better predictor than was disturbance per se, explaining approximately twice as much variation as did disturbance. 4. Synthesis. Disturbance is a weak predictor of invasion. To increase predictive power, we need to consider multiple variables (both intrinsic and extrinsic to the site) simultaneously. Variables that describe the changes sites have undergone may be particularly informative. [ABSTRACT FROM AUTHOR]

ABSTRACT Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high-rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree-C4 grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C4 grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life-history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire-resistant tree species in a C4 grass-dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them. [ABSTRACT FROM AUTHOR]

Spatio-temporal variation in tropical savanna tree cover remains poorly understood. We aimed to quantify the drivers of tree cover in tropical mesic savannas in Kakadu National Park by relating changes in tree cover over 40 years to: mean annual rainfall, fire activity, initial tree cover and prior changes in tree cover. Aerial photography, acquired in 1964, 1984 and 2004, was obtained for fifty sites in Kakadu that spanned a rainfall gradient from approximately 1200 to 1600 mm. The remotely sensed estimates of tree cover were validated via field survey. Linear mixed effects modelling and multi-model inference were used to assess the strength and form of the relationships between tree cover and predictor variables. Over the 40 years, tree cover across these savannas increased on average by 4.94 ± 0.88%, but was spatio-temporally variable. Tree cover showed a positive albeit weak trend across the rainfall gradient. The strength of this positive relationship varied over the three measurement times, and this suggests that other factors are important in controlling tree cover. Tree cover was positively related to prior tree cover, and negatively correlated with fire activity. Over 20 years tree cover was more likely to increase if (i) tree cover was initially low or (ii) had decreased in the previous 20-year interval or (iii) there had been fewer fires. Across the examined rainfall gradient, the greater variability in fire activity and inherently higher average tree cover at the wetter latitudes resulted in greater dynamism of tree cover compared with the drier latitudes. This is consistent with savanna tree cover being determined by interactions between mean annual rainfall, tree competition and frequent fire in these tropical mesic savannas. [ABSTRACT FROM AUTHOR]

The article presents information that derived from a systems workshop of the Research Network for Vegetation Function of Darwin and Kakadu National Park in Australia, which took place in September 2008. The workshop examined the anthropogenic alterations of tropical savannas through mechanisms such as land-use change and climate change.

1. How tree cover in tropical savannas changes through time and space is a major unresolved issue in ecology due to the complexity of these systems where fire, grazing, climatic variability and landscape variation interact, determining the spatial patterns of biomass. We aimed to assess the relative importance of fire and grazing in determining multi-decadal and landscape-scale patterns of savanna tree cover and biomass in a mesic savanna. 2. Assessment of digitized aerial photography, from 1964, 1984, 1991 and 2004, was undertaken for 40 sites at the Kapalga fire experimental area (~1470 mm mean annual rainfall) in Kakadu National Park in monsoonal northern Australia. 3. Spatio-temporal changes in tree cover were analysed using linear mixed effects modelling, with multi-model inference in an information theoretic framework. 4. Savanna tree cover fluctuated greatly over the 40-year period, and varied spatially amongst the 40 sites, by 20–30% in a given time period. Tree cover was most likely to decline when initially high, and with a high frequency of fire, especially late dry season fire. The presence of a feral grazer had little effect other than through reducing fire frequency. Our results are consistent with an overall increase in tree cover in these mesic savannas during drier-than-average periods. 5. Ground-based measurements of change in stand basal area under experimental burning regimes agreed with the estimated effects of fire on tree cover at the same site based on our analyses of aerial photography. 6. Synthesis and applications. The dynamism of these mesic savannas is driven by variation in fire frequency and severity, which is influenced by feral grazers, creating a mosaic of tree cover that shifts over time in response to localized variation in disturbance. These findings are relevant to the management of this biome. We show that changing disturbance regimes alter the dynamics of these systems. Fire management incorporating a variety of fire regimes will promote the persistence of a savanna patch mosaic. [ABSTRACT FROM AUTHOR]

Savannas are the major biome of tropical regions, spanning 30% of the Earth's land surface. Tree: grass ratios of savannas are inherently unstable and can be shifted easily by changes in fire, grazing, or climate. We synthesize the history and ecological impacts of the rapid expansion and eradication of an exotic large herbivore, the Asian water buffalo (Buhalus bubalus), on the mesic savannas of Kakadu National Park (KNP), a World Heritage Park located within the Alligator Rivers Region (ARR) of monsoonal north Australia. The study inverts the experience of the Serengeti savannas where grazing herds rapidly declined due to a rinderpest epidemic and then recovered upon disease control. Buffalo entered the ARR by the 1880s, but densities were low until the late 1950s when populations rapidly grew to carrying capacity within a decade. In the 1980s, numbers declined precipitously due to an eradication program. We show evidence that the rapid population expansion and sudden removal of this exotic herbivore created two ecological cascades by altering ground cover abundance and composition, which in turn affected competitive regimes and fuel loads with possible further, long-term effects due to changes in fire regimes. Overall, ecological impacts varied across a north-south gradient in KNP that corresponded to the interacting factors of precipitation, landform, and vegetation type but was also contingent upon the history of buffalo harvest. Floodplains showed the greatest degree of impact during the period of rapid buffalo expansion, but after buffalo removal, they largely reverted to their prior state. Conversely, the woodlands experienced less visible impact during the first cascade. However, in areas of low buffalo harvest and severe impact, there was little recruitment of juvenile trees into the canopy due to the indirect effects of grazing and high frequency of prescribed fires once buffalo were removed. Rain forests were clearly heavily impacted during the first cascade, but the long term consequences of buffalo increase and removal remain unclear. Due to hysteresis effects, the simple removal of an exotic herbivore was not sufficient to return savanna systems to their previous state. [ABSTRACT FROM AUTHOR]

The article presents a research study regarding the limits of savanna on the continents of Africa, Australia, and South America. The effects of rainfall, seasonality, and soil fertility on the growth and distribution of savanna are examined, as well as the interaction of these factors with natural disturbances.