Fire facilitates ground layer plant diversity in a Miombo ecosystem.

BACKGROUND AND AIMS: Little is known about the response of ground layer plant communities to fire in Miombo ecosystems, which is a global blind spot of ecological understanding. We aimed: (1) to assess the impact of three experimentally imposed fire treatments on ground layer species composition and compare it with patterns observed for trees; and (2) to analyse the effect of fire treatments on species richness to assess how responses differ among plant functional groups. METHODS: At a 60-year-long fire experiment in Zambia, we quantified the richness and diversity of ground layer plants in terms of taxa and functional groups across three experimental fire treatments of late dry-season fire, early dry-season fire and fire exclusion. Data were collected in five repeat surveys from the onset of the wet season to the early dry season. KEY RESULTS: Of the 140 ground layer species recorded across the three treatments, fire-maintained treatments contributed most of the richness and diversity, with the least number of unique species found in the no-fire treatment. The early-fire treatment was more similar in composition to the no-fire treatment than to the late-fire treatment. C4 grass and geoxyle richness were highest in the late-fire treatment, and there were no shared sedge species between the late-fire and other treatments. At a plot level, the average richness in the late-fire treatment was twice that of the fire exclusion treatment. CONCLUSIONS: Heterogeneity in fire seasonality and intensity supports diversity of a unique flora by providing a diversity of local environments. African ecosystems face rapid expansion of land- and fire-management schemes for carbon offsetting and sequestration. We demonstrate that analyses of the impacts of such schemes predicated on the tree flora alone are highly likely to underestimate impacts on biodiversity. A research priority must be a new understanding of the Miombo ground layer flora integrated into policy and land management.

Functional and ecological diversification of underground organs in Solanum.

The evolution of geophytes in response to different environmental stressors is poorly understood largely due to the great morphological variation in underground plant organs, which includes species with rhizomatous structures or underground storage organs (USOs). Here we compare the evolution and ecological niche patterns of different geophytic organs in Solanum L., classified based on a functional definition and using a clade-based approach with an expert-verified specimen occurrence dataset. Results from PERMANOVA and Phylogenetic ANOVAs indicate that geophytic species occupy drier areas, with rhizomatous species found in the hottest areas whereas species with USOs are restricted to cooler areas in the montane tropics. In addition, rhizomatous species appear to be adapted to fire-driven disturbance, in contrast to species with USOs that appear to be adapted to prolonged climatic disturbance such as unfavorable growing conditions due to drought and cold. We also show that the evolution of rhizome-like structures leads to changes in the relationship between range size and niche breadth. Ancestral state reconstruction shows that in Solanum rhizomatous species are evolutionarily more labile compared to species with USOs. Our results suggest that underground organs enable plants to shift their niches towards distinct extreme environmental conditions and have different evolutionary constraints.

Underground trees inhabit varied environmental extremes across the Afrotropics.

BACKGROUND AND AIMS: Geoxyles, a distinctive feature of Afrotropical savannas and grasslands, survive recurrent disturbances by resprouting subshrub branches from large belowground woody structures. Underground trees are a type of geoxyle that independently evolved within woody genera of at least 40 plant families in Africa. The environmental limits and determinants of underground tree biogeography are poorly understood with the relative influence of frost and fire debated in particular. We aim to quantify variability in the niche of underground tree species relative to their taller, woody tree/shrub congeners. METHODS: Using occurrence records of four Afrotropical genera, Parinari (Chrysobalanaceae), Ozoroa (Anacardiaceae), Syzygium (Myrtaceae) and Lannea (Anacardiaceae), and environmental data of nine climate and disturbance variables, the biogeography and niche of underground trees are compared with their open and closed ecosystem congeners. KEY RESULTS: Along multiple environmental gradients and in a multidimensional environmental space, underground trees inhabit significantly distinct and extreme environments relative to open and closed ecosystem congeners. Niche overlap is low among underground trees and their congeners, and also among underground trees of the four genera. Of the study taxa, Parinari underground trees inhabit hotter, drier and more seasonal environments where herbivory pressure is greatest. Ozoroa underground trees occupy relatively more fire prone environments, while Syzygium underground trees sustain the highest frost frequency and occur in relatively wetter conditions with seasonal waterlogging. Lannea underground trees are associated with the lowest temperatures, highest precipitation, and varying exposure to disturbance. CONCLUSIONS: While underground trees exhibit repeated convergent evolution, distinct environments shape the ecology and biogeography of this iconic plant functional group. The multiplicity of extreme environments related to fire, frost, herbivory and waterlogging that different underground tree taxa occupy, and the distinctiveness of these environments, should be recognised in the management of African grassy ecosystems.

Madagascar's extraordinary biodiversity: Evolution, distribution, and use.

Madagascar's biota is hyperdiverse and includes exceptional levels of endemicity. We review the current state of knowledge on Madagascar's past and current terrestrial and freshwater biodiversity by compiling and presenting comprehensive data on species diversity, endemism, and rates of species description and human uses, in addition to presenting an updated and simplified map of vegetation types. We report a substantial increase of records and species new to science in recent years; however, the diversity and evolution of many groups remain practically unknown (e.g., fungi and most invertebrates). Digitization efforts are increasing the resolution of species richness patterns and we highlight the crucial role of field- and collections-based research for advancing biodiversity knowledge and identifying gaps in our understanding, particularly as species richness corresponds closely to collection effort. Phylogenetic diversity patterns mirror that of species richness and endemism in most of the analyzed groups. We highlight humid forests as centers of diversity and endemism because of their role as refugia and centers of recent and rapid radiations. However, the distinct endemism of other areas, such as the grassland-woodland mosaic of the Central Highlands and the spiny forest of the southwest, is also biologically important despite lower species richness. The documented uses of Malagasy biodiversity are manifold, with much potential for the uncovering of new useful traits for food, medicine, and climate mitigation. The data presented here showcase Madagascar as a unique "living laboratory" for our understanding of evolution and the complex interactions between people and nature. The gathering and analysis of biodiversity data must continue and accelerate if we are to fully understand and safeguard this unique subset of Earth's biodiversity.

Madagascar's extraordinary biodiversity: Threats and opportunities.

Madagascar's unique biota is heavily affected by human activity and is under intense threat. Here, we review the current state of knowledge on the conservation status of Madagascar's terrestrial and freshwater biodiversity by presenting data and analyses on documented and predicted species-level conservation statuses, the most prevalent and relevant threats, ex situ collections and programs, and the coverage and comprehensiveness of protected areas. The existing terrestrial protected area network in Madagascar covers 10.4% of its land area and includes at least part of the range of the majority of described native species of vertebrates with known distributions (97.1% of freshwater fishes, amphibians, reptiles, birds, and mammals combined) and plants (67.7%). The overall figures are higher for threatened species (97.7% of threatened vertebrates and 79.6% of threatened plants occurring within at least one protected area). International Union for Conservation of Nature (IUCN) Red List assessments and Bayesian neural network analyses for plants identify overexploitation of biological resources and unsustainable agriculture as the most prominent threats to biodiversity. We highlight five opportunities for action at multiple levels to ensure that conservation and ecological restoration objectives, programs, and activities take account of complex underlying and interacting factors and produce tangible benefits for the biodiversity and people of Madagascar.

A function-based typology for Earth's ecosystems.

As the United Nations develops a post-2020 global biodiversity framework for the Convention on Biological Diversity, attention is focusing on how new goals and targets for ecosystem conservation might serve its vision of 'living in harmony with nature'1,2. Advancing dual imperatives to conserve biodiversity and sustain ecosystem services requires reliable and resilient generalizations and predictions about ecosystem responses to environmental change and management3. Ecosystems vary in their biota4, service provision5 and relative exposure to risks6, yet there is no globally consistent classification of ecosystems that reflects functional responses to change and management. This hampers progress on developing conservation targets and sustainability goals. Here we present the International Union for Conservation of Nature (IUCN) Global Ecosystem Typology, a conceptually robust, scalable, spatially explicit approach for generalizations and predictions about functions, biota, risks and management remedies across the entire biosphere. The outcome of a major cross-disciplinary collaboration, this novel framework places all of Earth's ecosystems into a unifying theoretical context to guide the transformation of ecosystem policy and management from global to local scales. This new information infrastructure will support knowledge transfer for ecosystem-specific management and restoration, globally standardized ecosystem risk assessments, natural capital accounting and progress on the post-2020 global biodiversity framework.

Grazing lawns and overgrazing in frequently grazed grass communities.

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.

Encroachment diminishes herbaceous plant diversity in grassy ecosystems worldwide.

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.

Madagascar's fire regimes challenge global assumptions about landscape degradation.

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.

Nitrogen concentration and physical properties are key drivers of woody tissue respiration.

BACKGROUND AND AIMS: Despite the critical role of woody tissues in determining net carbon exchange of terrestrial ecosystems, relatively little is known regarding the drivers of sapwood and bark respiration. METHODS: Using one of the most comprehensive wood respiration datasets to date (82 species from Australian rainforest, savanna and temperate forest), we quantified relationships between tissue respiration rates (Rd) measured in vitro (i.e. 'respiration potential') and physical properties of bark and sapwood, and nitrogen concentration (Nmass) of leaves, sapwood and bark. KEY RESULTS: Across all sites, tissue density and thickness explained similar, and in some cases more, variation in bark and sapwood Rd than did Nmass. Higher density bark and sapwood tissues had lower Rd for a given Nmass than lower density tissues. Rd-Nmass slopes were less steep in thicker compared with thinner-barked species and less steep in sapwood than in bark. Including the interactive effects of Nmass, density and thickness significantly increased the explanatory power for bark and sapwood respiration in branches. Among these models, Nmass contributed more to explanatory power in trunks than in branches, and in sapwood than in bark. Our findings were largely consistent across sites, which varied in their climate, soils and dominant vegetation type, suggesting generality in the observed trait relationships. Compared with a global compilation of leaf, stem and root data, Australian species showed generally lower Rd and Nmass, and less steep Rd-Nmass relationships. CONCLUSIONS: To the best of our knowledge, this is the first study to report control of respiration-nitrogen relationships by physical properties of tissues, and one of few to report respiration-nitrogen relationships in bark and sapwood. Together, our findings indicate a potential path towards improving current estimates of autotrophic respiration by integrating variation across distinct plant tissues.

Resprouting grasses are associated with less frequent fire than seeders.

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.

Fire and grazing determined grasslands of central Madagascar represent ancient assemblages.

The ecology of Madagascar's grasslands is under-investigated and the dearth of ecological understanding of how disturbance by fire and grazing shapes these grasslands stems from a perception that disturbance shaped Malagasy grasslands only after human arrival. However, worldwide, fire and grazing shape tropical grasslands over ecological and evolutionary timescales, and it is curious Madagascar should be a global anomaly. We examined the functional and community ecology of Madagascar's grasslands across 71 communities in the Central Highlands. Combining multivariate abundance models of community composition and clustering of grass functional traits, we identified distinct grass assemblages each shaped by fire or grazing. The fire-maintained assemblage is primarily composed of tall caespitose species with narrow leaves and low bulk density. By contrast, the grazer-maintained assemblage is characterized by mat-forming, high bulk density grasses with wide leaves. Within each assemblage, levels of endemism, diversity and grass ages support these as ancient assemblages. Grazer-dependent grasses can only have co-evolved with a now-extinct megafauna. Ironically, the human introduction of cattle probably introduced a megafaunal substitute facilitating modern day persistence of a grazer-maintained grass assemblage in an otherwise defaunated landscape, where these landscapes now support the livelihoods of millions of people.

Lineage-based functional types: characterising functional diversity to enhance the representation of ecological behaviour in Land Surface Models.

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.

Comment on "The global tree restoration potential".

Bastin et al's estimate (Reports, 5 July 2019, p. 76) that tree planting for climate change mitigation could sequester 205 gigatonnes of carbon is approximately five times too large. Their analysis inflated soil organic carbon gains, failed to safeguard against warming from trees at high latitudes and elevations, and considered afforestation of savannas, grasslands, and shrublands to be restoration.

The Trouble with Trees: Afforestation Plans for Africa.

Extensive tree planting is widely promoted for reducing atmospheric CO2. In Africa, 1 million km2, mostly of grassy biomes, have been targeted for 'restoration' by 2030. The target is based on the erroneous assumption that these biomes are deforested and degraded. We discuss the pros and cons of exporting fossil fuel emission problems to Africa.

Alternate Grassy Ecosystem States Are Determined by Palatability-Flammability Trade-Offs.

Fire and mammalian grazers both consume grasses, and feedbacks between grass species, their functional traits, and consumers have profound effects on grassy ecosystem structure worldwide, such that savanna and grassland states determined by fire or grazing can be considered alternate states. These parallel savanna-forest alternate states, which likewise have myriad cascading ecosystem impacts.

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.

Tropical savannas and dry forests.

In the tropics, research, conservation and public attention focus on rain forests, but this neglects that half of the global tropics have a seasonally dry climate. These regions are home to dry forests and savannas (Figures 1 and 2), and are the focus of this Primer. The attention given to rain forests is understandable. Their high species diversity, sheer stature and luxuriance thrill biologists today as much as they did the first explorers in the Age of Discovery. Although dry forest and savanna may make less of a first impression, they support a fascinating diversity of plant strategies to cope with stress and disturbance including fire, drought and herbivory. Savannas played a fundamental role in human evolution, and across Africa and India they support iconic megafauna.