Phylogenetic congruence between Neotropical primates and plants is driven by frugivory.

Seed dispersal benefits plants and frugivores, and potentially drives co-evolution, with consequences to diversification evidenced for, e.g., primates. Evidence for macro-coevolutionary patterns in multi-specific, plant-animal mutualisms is scarce, and the mechanisms driving them remain unexplored. We tested for phylogenetic congruences in primate-plant interactions and showed strong co-phylogenetic signals across Neotropical forests, suggesting that both primates and plants share evolutionary history. Phylogenetic congruence between Platyrrhini and Angiosperms was driven by the most generalist primates, modulated by their functional traits, interacting with a wide-range of Angiosperms. Consistently similar eco-evolutionary dynamics seem to be operating irrespective of local assemblages, since co-phylogenetic signal emerged independently across three Neotropical regions. Our analysis supports the idea that macroevolutionary, coevolved patterns among interacting mutualistic partners are driven by super-generalist taxa. Trait convergence among multiple partners within multi-specific assemblages appears as a mechanism favouring these likely coevolved outcomes.

Revisiting florivory: an integrative review and global patterns of a neglected interaction.

Florivory is an ancient interaction which has rarely been quantified due to a lack of standardized protocols, thus impairing biogeographical and phylogenetic comparisons. We created a global, continuously updated, open-access database comprising 180 species and 64 families to compare floral damage between tropical and temperate plants, to examine the effects of plant traits on floral damage, and to explore the eco-evolutionary dynamics of flower-florivore interactions. Flower damage is widespread across angiosperms, but was two-fold higher in tropical vs temperate species, suggesting stronger fitness impacts in the tropics. Flowers were mostly damaged by chewers, but neither flower color nor symmetry explained differences in florivory. Herbivory and florivory levels were positively correlated within species, even though the richness of the florivore community does not affect florivory levels. We show that florivory impacts plant fitness via multiple pathways and that ignoring this interaction makes it more difficult to obtain a broad understanding of the ecology and evolution of angiosperms. Finally, we propose a standardized protocol for florivory measurements, and identify key research avenues that will help fill persistent knowledge gaps. Florivory is expected to be a central research topic in an epoch characterized by widespread decreases in insect populations that comprise both pollinators and florivores.

Towards the flower economics spectrum.

Understanding how floral traits affect reproduction is key for understanding genetic diversity, speciation, and trait evolution in the face of global changes and pollinator decline. However, there has not yet been a unified framework to characterize the major trade-offs and axes of floral trait variation. Here, we propose the development of a floral economics spectrum (FES) that incorporates the multiple pathways by which floral traits can be shaped by multiple agents of selection acting on multiple flower functions. For example, while pollinator-mediated selection has been considered the primary factor affecting flower evolution, selection by nonpollinator agents can reinforce or oppose pollinator selection, and, therefore, affect floral trait variation. In addition to pollinators, the FES should consider nonpollinator biotic agents and floral physiological costs, broadening the drivers of floral traits beyond pollinators. We discuss how coordinated evolution and trade-offs among floral traits and between floral and vegetative traits may influence the distribution of floral traits across biomes and lineages, thereby influencing organismal evolution and community assembly.

Frugivory and seed dispersal in a hyperdiverse plant clade and its role as a keystone resource for the Neotropical fauna.

BACKGROUND AND AIMS: Much of our understanding of the ecology and evolution of seed dispersal in the Neotropics is founded on studies involving the animal-dispersed, hyperdiverse plant clade Miconia (Melastomataceae). Nonetheless, no formal attempt has been made to establish its relevance as a model system or indeed provide evidence of the role of frugivores as Miconia seed dispersers. METHODS: We built three Miconia databases (fruit phenology/diaspore traits, fruit-frugivore interactions and effects on seed germination after gut passage) to determine how Miconia fruiting phenology and fruit traits for >350 species interact with and shape patterns of frugivore selection. In addition, we conducted a meta-analysis evaluating the effects of animal gut passage/seed handling on Miconia germination. KEY RESULTS: Miconia produce numerous small berries that enclose numerous tiny seeds within water- and sugar-rich pulps. In addition, coexisting species provide sequential, year long availability of fruits within communities, with many species producing fruits in periods of resource scarcity. From 2396 pairwise interactions, we identified 646 animal frugivore species in five classes, 22 orders and 60 families, including birds, mammals, reptiles, fish and ants that consume Miconia fruits. Endozoochory is the main dispersal mechanism, but gut passage effects on germination were specific to animal clades; birds, monkeys and ants reduced seed germination percentages, while opossums increased it. CONCLUSIONS: The sequential fruiting phenologies and wide taxonomic and functional diversity of animal vectors associated with Miconia fruits underscore the likely keystone role that this plant clade plays in the Neotropics. By producing fruits morphologically and chemically accessible to a variety of animals, Miconia species ensure short- and long-distance seed dispersal and constitute reliable resources that sustain entire frugivore assemblages.

A research agenda for seed-trait functional ecology.

Trait-based approaches have improved our understanding of plant evolution, community assembly and ecosystem functioning. A major challenge for the upcoming decades is to understand the functions and evolution of early life-history traits, across levels of organization and ecological strategies. Although a variety of seed traits are critical for dispersal, persistence, germination timing and seedling establishment, only seed mass has been considered systematically. Here we suggest broadening the range of morphological, physiological and biochemical seed traits to add new understanding on plant niches, population dynamics and community assembly. The diversity of seed traits and functions provides an important challenge that will require international collaboration in three areas of research. First, we present a conceptual framework for a seed ecological spectrum that builds upon current understanding of plant niches. We then lay the foundation for a seed-trait functional network, the establishment of which will underpin and facilitate trait-based inferences. Finally, we anticipate novel insights and challenges associated with incorporating diverse seed traits into predictive evolutionary ecology, community ecology and applied ecology. If the community invests in standardized seed-trait collection and the implementation of rigorous databases, major strides can be made at this exciting frontier of functional ecology.

Fire and legume germination in a tropical savanna: ecological and historical factors.

BACKGROUND AND AIMS: In many flammable ecosystems, physically dormant seeds show dormancy-break patterns tied to fire, but the link between heat shock and germination in the tropical savannas of Africa and South America remains controversial. Seed heat tolerance is important, preventing seed mortality during fire passage, and is usually predicted by seed traits. This study investigated the role of fire frequency (ecological effects) and seed traits through phylogenetic comparison (historical effects), in determining post-fire germination and seed mortality in legume species of the Cerrado, a tropical savanna-forest mosaic. METHODS: Seeds of 46 legume species were collected from three vegetation types (grassy savannas, woody savannas and forests) with different fire frequencies. Heat shock experiments (100 °C for 1 min; 100 °C for 3 min; 200 °C for 1 min) were then performed, followed by germination and seed viability tests. Principal component analysis, generalized linear mixed models and phylogenetic comparisons were used in data analyses. KEY RESULTS: Heat shocks had little effect on germination, but seed mortality was variable across treatments and species. Seed mortality was lowest under the 100 °C 1 min treatment, and significantly higher under 100 °C 3 min and 200 °C 1 min; larger seed mass decreased seed mortality, especially at 200 °C. Tree species in Detarioideae had the largest seeds and were unaffected by heat. Small-seeded species (mostly shrubs from grassy savannas) were relatively sensitive to the hottest treatment. Nevertheless, the presence of physical dormancy helped to avoid seed mortality in small-seeded species under the hottest treatment. CONCLUSIONS: Physical dormancy-break is not tied to fire in the Cerrado mosaic. Heat tolerance appears in both forest and savanna species and is predicted by seed traits (seed mass and physical dormancy), which might have helped forest lineages to colonize the savannas. The results show seed fire responses are better explained by historical than ecological factors in the Cerrado, contrasting with different fire-prone ecosystems throughout the world.

Timing of seed dispersal and seed dormancy in Brazilian savanna: two solutions to face seasonality.

Background and Aims: The relationship between fruiting phenology and seed dispersal syndrome is widely recognized; however, the interaction of dormancy classes and plant life-history traits in relation to fruiting phenology and seed dispersal is understudied. Here we examined the relationship between fruiting season and seed dormancy and how this relationship is modulated by dormancy classes, dispersal syndromes, seed mass and seed moisture content in a Brazilian savanna (cerrado). Methods: Dormancy classes (non-dormancy and physical, morphological, morphophysiological, physiological and physiophysical dormancy) of 34 cerrado species were experimentally determined. Their seed dispersal syndrome (autochory, anemochory, zoochory), dispersal season (rainy, dry, rainy-to-dry and dry-to-rainy transitions), seed mass and moisture contents, and the estimated germination date were also determined. Log-linear models were used to evaluate how dormancy and dormancy classes are related to dispersal season and syndrome. Key Results: The proportions of dormant and non-dormant species were similar in cerrado. The community-estimated germination date was seasonal, occurring at the onset of rainy season. Overall, anemochorous non-dormant species released seeds during the dry-to-rainy transition; autochorous physically dormant species dispersed seeds during the dry season and rainy-to-dry transition; zoochorous species dispersed non-dormant seeds during the dry and rainy seasons, while species with morphological, morphophysiological or physiological dormancy dispersed seeds in the transitional seasons. Seed mass differed among dispersal seasons and dormancy classes, but seed moisture content did not vary with dispersal syndrome, season or dormancy class. Conclusions: The beginning of the rainy season was the most favourable period for seed germination in cerrado, and the germination phenology was controlled by both the timing of seed dispersal and seed dormancy. Dormancy class was influenced by dispersal syndrome and season. Moreover, dormancy avoided seed germination during the rainy-to-dry transition, independently of dispersal syndrome. The variability of dormancy classes with dispersal syndrome allowed animal-dispersed species to fruit all year round, but seeds germinated only during the rainy season. Conversely, seasonally restricted wind-dispersal species dispersed and germinated their non-dormant seeds only in the rainy season.

Phylogeny strongly drives seed dormancy and quality in a climatically buffered hotspot for plant endemism.

BACKGROUND AND AIMS: Models of costs and benefits of dormancy (D) predict that the evolutionarily stable strategy in long-term stable environments is for non-dormancy (ND), but this prediction remains to be tested empirically. We reviewed seed traits of species in the climatically buffered, geologically stable and nutrient-impoverished campo rupestre grasslands in Brazil to test the hypothesis that ND is favoured over D. We examined the relative importance of life-history traits and phylogeny in driving the evolution of D and assessed seed viability at the community level. METHODS: Germination and viability data were retrieved from 67 publications and ND/D was determined for 168 species in 25 angiosperm families. We also obtained the percentage of embryoless, viable and dormant seeds for 74 species. Frequencies of species with dormant and non-dormant seeds were compared with global databases of dormancy distribution. KEY RESULTS: The majority of campo rupestre taxa (62·5 %) had non-dormant seeds, and the ND/D ratio was the highest for any vegetation type on Earth. Dormancy was unrelated to other species life-history traits, suggesting that contemporary factors are poor predictors of D. We found a significant phylogenetic structure in the dormancy categorical trait. Dormancy diversity was highly skewed towards the root of the phylogenetic tree and there was a strong phylogenetic signal in the data, suggesting a major role of phylogeny in determining the evolution of D versus ND and seed viability. Quantitative analysis of the data revealed that at least half of the seeds produced by 46 % of the surveyed populations were embryoless and/or otherwise non-viable. CONCLUSIONS: Our results support the view that long-term climatic and geological stability favour ND. Seed viability data show that campo rupestre species have a markedly low investment in regeneration from seeds, highlighting the need for specific in situ and ex situ conservation strategies to avoid loss of biodiversity.

Intraspecific variation in fruit-frugivore interactions: effects of fruiting neighborhood and consequences for seed dispersal.

The extent of specialization/generalization continuum in fruit-frugivore interactions at the individual level remains poorly explored. Here, we investigated the interactions between the Neotropical treelet Miconia irwinii (Melastomataceae) and its avian seed dispersers in Brazilian campo rupestre. We built an individual-based network to derive plant degree of interaction specialization regarding disperser species. Then, we explored how intraspecific variation in interaction niche breadth relates to fruit availability on individual plants in varying densities of fruiting conspecific neighbors, and how these factors affect the quantity of viable seeds dispersed. We predicted broader interaction niche breadths for individuals with larger fruit crops in denser fruiting neighborhoods. The downscaled network included nine bird species and 15 plants, which varied nearly five-fold in their degree of interaction specialization. We found positive effects of crop size on visitation and fruit removal rates, but not on degree of interaction specialization. Conversely, we found that an increase in the density of conspecific fruiting neighbors both increased visitation rate and reduced plant degree of interaction specialization. We suggest that tracking fruit-rich patches by avian frugivore species is the main driver of density-dependent intraspecific variation in plants' interaction niche breadth. Our study shed some light on the overlooked fitness consequences of intraspecific variation in interaction niches by showing that individuals along the specialization/generalization continuum may have their seed dispersed with similar effectiveness. Our study exemplifies how individual-based networks linking plants to frugivore species that differ in their seed dispersal effectiveness can advance our understanding of intraspecific variation in the outcomes of fruit-frugivore interactions.

How far do Neotropical primates disperse seeds?

Seed dispersal distance (SDD) is a vital component of vertebrate-mediated seed dispersal process: the average distance at which seeds are deposited away from the parent plant represents the starting template of plant regeneration. We present a simple model to explain and predict observed measures of average dispersal distance and we hypothesize that it is a consequence of how long seeds are retained in the disperser's gut, how rapidly the disperser moves per unit time and how twisted the animal travel path is relative to the straight-line distance moved away from the seed source. We retrieved data on dispersal distances from 26 published studies including nine primate species dispersing up to 112 plant species per study. We used gut transit time (TT) as a proxy for residence time inside the gut, the disperser's travel path per hour as proxy for movement rate, and the daily path length relative to the home range area as a correlate of path twisting (PT). We illustrate this model with comparative data on Neotropical primates. These three variables explained 90% of the variation in the average SDD. Path analysis indicates that additional variables exerted only indirect effects. Our model can be applied to primate populations for which detailed seed dispersal data are missing, and help evaluate conservation priorities for primate species according to the potential service they provide in terms of forest regeneration.

Assessing bias and knowledge gaps on seed ecology research: implications for conservation agenda and policy.

Sampling biases permeate ecological research and result in knowledge gaps that have vital consequences for conservation planning. The consequences of knowledge gaps on species identity and distribution (the Wallacean and Linnean shortfalls, respectively) have become apparent recently, but we know little about the extent that research biases and knowledge gaps on traits that influence species' niches (the Hutchinsonian shortfall) affect conservation policy. To examine whether knowledge of species' traits based on seed ecology is geographically, phylogenetically, and ecologically biased, we retrieved research data on seed germination, seed dormancy, seed dispersal, seed banks, seed predation, and seed removal from a database of 847 papers, 1648 species, and 5322 cases. Brazil was selected as a model system for megadiverse, undersampled countries. Kernel density maps showed that research was geographically biased towards highly populated sites, with vast areas remaining historically unexplored. We also show that research was clustered into protected areas. We detected a significant positive phylogenetic bias at genus-level, indicating research concentration in few genera and lower relative bias rates for many herbaceous genera. Unexpectedly, information on seed banking was available for only 74 (3.4%) of threatened species, which suggests that information deficits are highest for species with critical needs for ex situ conservation strategies. Tree, fleshy-fruited, and biotic dispersal species were disproportionately overstudied. Our data indicate that information deficits on seed ecology preclude our ability to effectively restore ecosystems and to safeguard endangered species. We call for a systematic improvement of environmental agenda in which policy makers and scientists target sites, clades, and functional groups historically neglected. Lessons from developed countries and collaborative efforts will be important for megadiverse, underdeveloped countries to achieve the targets of international agreements that depend on seed ecology knowledge aiming to secure biological diversity and ecosystem services.

MelastomaTRAITs 1.0: A database of functional traits in Melastomataceae, a large pantropical angiosperm family.

The recent availability of open-access repositories of functional traits has revolutionized trait-based approaches in ecology and evolution. Nevertheless, the underrepresentation of tropical regions and lineages remains a pervasive bias in plant functional trait databases, which constrains large-scale assessments of plant ecology, evolution, and biogeography. Here, we present MelastomaTRAITs 1.0, a comprehensive and updatable database of functional traits for the pantropical Melastomataceae, the ninth-largest angiosperm family with 177 genera and more than 5800 species. Melastomataceae encompass species with a wide diversity of growth forms (herbs, shrubs, trees, epiphytes, and woody climbers), habitats (including tropical forests, savannas, grasslands, and wetlands from sea level to montane areas above the treeline), ecological strategies (from pioneer, edge-adapted and invasive species to shade-tolerant understory species), geographic distribution (from microendemic to continental-wide distribution), reproductive, pollination, and seed dispersal systems. MelastomaTRAITs builds on 581 references, such as taxonomic monographs, ecological research, and unpublished data, and includes four whole-plant traits, six leaf traits, 11 flower traits, 18 fruit traits, and 27 seed traits for 2520 species distributed in 144 genera across all 21 tribes. Most data come from the Neotropics where the family is most species-rich. Miconieae (the largest tribe) contains the highest number of trait records (49.6%) and species (41.1%) records. The trait types with the most information in the database were whole-plant traits, flowers, and leaf traits. With the breadth of functional traits recorded, our database helps to fill a gap in information for tropical plants and will significantly improve our capacity for large-scale trait-based syntheses across levels of organization, plant-animal interactions, regeneration ecology, and thereby support conservation and restoration programs. There are no copyright restrictions on the dataset; please cite this data paper when reusing the data.

How much leaf area do insects eat? A data set of insect herbivory sampled globally with a standardized protocol.

Herbivory is ubiquitous. Despite being a potential driver of plant distribution and performance, herbivory remains largely undocumented. Some early attempts have been made to review, globally, how much leaf area is removed through insect feeding. Kozlov et al., in one of the most comprehensive reviews regarding global patterns of herbivory, have compiled published studies regarding foliar removal and sampled data on global herbivory levels using a standardized protocol. However, in the review by Kozlov et al., only 15 sampling sites, comprising 33 plant species, were evaluated in tropical areas around the globe. In Brazil, which ranks first in terms of plant biodiversity, with a total of 46,097 species, almost half (43%) being endemic, a single data point was sampled, covering only two plant species. In an attempt to increase knowledge regarding herbivory in tropical plant species and to provide the raw data needed to test general hypotheses related to plant-herbivore interactions across large spatial scales, we proposed a joint, collaborative network to evaluate tropical herbivory. This network allowed us to update and expand the data on insect herbivory in tropical and temperate plant species. Our data set, collected with a standardized protocol, covers 45 sampling sites from nine countries and includes leaf herbivory measurements of 57,239 leaves from 209 species of vascular plants belonging to 65 families from tropical and temperate regions. They expand previous data sets by including a total of 32 sampling sites from tropical areas around the globe, comprising 152 species, 146 of them being sampled in Brazil. For temperate areas, it includes 13 sampling sites, comprising 59 species. Thus, when compared to the most recent comprehensive review of insect herbivory (Kozlov et al.), our data set has increased the base of available data for the tropical plants more than 460% (from 33 to 152 species) and the Brazilian sampling was increased 7,300% (from 2 to 146 species). Data on precise levels of herbivory are presented for more than 57,000 leaves worldwide. There are no copyright restrictions. Please cite this paper when using the current data in publications; the authors request to be informed how the data is used in the publications.

Seed Functional Traits Provide Support for Ecological Restoration and ex situ Conservation in the Threatened Amazon Ironstone Outcrop Flora.

Cangas (ironstone outcrops) host a specialized flora, characterized by high degree of edaphic endemism and an apparent lack of natural history knowledge of its flora. Due to intense pressure from iron ore mining this ecosystem is under threat and in need of restoration. We studied seed functional traits that are relevant for restoration, translocation and ex situ conservation in 48 species from cangas in eastern Amazon. Were determined the thermal niche breadth, classified seed dormancy and determined methods to overcome it, determined the effect of seed storage on germination, tested the association between germination traits and functional groups, and tested whether seed traits are phylogenetically conserved. We found a broad interspecific variation in most seed traits, except for seed water content. Large interspecific variation in the temperature niche breadth was found among the studied species, but only four species, showed optimum germination at high temperatures of 35-40°C, despite high temperatures under natural conditions. Only 35% of the studied species produced dormant seeds. Mechanical scarification was effective in overcoming physical dormancy and application of gibberellic acid was effective in overcoming physiological dormancy in five species. For the 29 species that seeds were stored for 24 months, 76% showed decreases in the germination percentage. The weak association between germination traits and life-history traits indicate that no particular plant functional type requires specific methods for seed-based translocations. Exceptions were the lianas which showed relatively larger seeds compared to the other growth-forms. Dormancy was the only trait strongly related to phylogeny, suggesting that phylogenetic relatedness may not be a good predictor of regeneration from seeds in cangas. Our study provides support to better manage seed sourcing, use, storage and enhancement techniques with expected reduced costs and increased seedling establishment success.

Linking Plant Functional Ecology to Island Biogeography.

The study of insular systems has a long history in ecology and biogeography. Island plants often differ remarkably from their noninsular counterparts, constituting excellent models for exploring eco-evolutionary processes. Trait-based approaches can help to answer important questions in island biogeography, yet plant trait patterns on islands remain understudied. We discuss three key hypotheses linking functional ecology to island biogeography: (i) plants in insular systems are characterized by distinct functional trait syndromes (compared with noninsular environments); (ii) these syndromes differ between true islands and terrestrial habitat islands; and (iii) island characteristics influence trait syndromes in a predictable manner. We are convinced that implementing trait-based comparative approaches would considerably further our understanding of plant ecology and evolution in insular systems.

Ants on plants: a meta-analysis of the role of ants as plant biotic defenses.

We reviewed the evidence on the role of ants as plant biotic defenses, by conducting meta-analyses for the effects of experimental removal of ants on plant herbivory and fitness with data pooled from 81 studies. Effects reviewed were plant herbivory, herbivore abundance, hemipteran abundance, predator abundance, plant biomass and reproduction in studies where ants were experimentally removed (n = 273 independent comparisons). Ant removal exhibited strong effects on herbivory rates, as plants without ants suffered almost twice as much damage and exhibited 50% more herbivores than plants with ants. Ants also influenced several parameters of plant fitness, as plants without ants suffered a reduction in biomass (-23.7%), leaf production (-51.8%), and reproduction (-24.3%). Effects were much stronger in tropical regions compared to temperate ones. Tropical plants suffered almost threefold higher herbivore damage than plants from temperate regions and exhibited three times more herbivores. Ant removal in tropical plants resulted in a decrease in plant fitness of about 59%, whereas in temperate plants this reduction was not statistically significant. Ant removal effects were also more important in obligate ant-plants (=myrmecophytes) compared to plants exhibiting facultative relationships with hemiptera or those plants with extrafloral nectaries and food bodies. When only tropical plants were considered and the strength of the association between ants and plants taken into account, plants with obligate association with ants exhibited almost four times higher herbivory compared to plants with facultative associations with ants, but similar reductions in plant reproduction. The removal of a single ant species increased plant herbivory by almost three times compared to the removal of several ant species. Altogether, these results suggest that ants do act as plant biotic defenses, but the effects of their presence are more pronounced in tropical systems, especially in myrmecophytic plants.

Water-use strategies in flowers from a neotropical savanna under contrasting environmental conditions during flowering.

Flowers require high amounts of water, which ultimately may compromise pollinator attractiveness under water limitation. Water-use and -conservation strategies in leaves from hot and dry ecosystems are well documented, yet little is known about mechanisms of water allocation in flowers, particularly in tropical savanna ecosystems. We evaluated traits related to corolla water status in two Kielmeyera species that differ in flowering phenology and flower size: larger-flowered K. regalis blooms during the rainy summer and smaller-flowered K. coriacea blooms during the dry winter. To test the hypothesis that water demand in corollas increases with increasing vapor pressure deficit (VPD), we analyzed interspecific differences in corolla stomatal conductance and density, water content, and fresh and dry mass per unit area. We also performed hand-pollination and pollinator-exclusion experiments to determine variation in floral longevity. Corolla transpiration rates were higher in K. coriacea (157 vs 95 g·H2O·m-2·h-1 for K. coriacea and K. regalis, respectively), and increased with VPD in both species. Stomatal density was 25-fold higher in K. coriacea, and corolla fresh and dry mass per unit of area were 47% and 21% higher, respectively, in K. coriacea, due to thick pectin-rich cell walls. The high pectin content increases water content in corollas of K. coriacea. Regardless of pollination, flowers lasted one day in K. coriacea and three in K. regalis. Our study suggests structure-function relationships of floral traits with flowering season, and that K. coriacea displays small and short-lived corollas with high water content to buffer the high evaporative demand during the dry period.

Resilience and restoration of tropical and subtropical grasslands, savannas, and grassy woodlands.

Despite growing recognition of the conservation values of grassy biomes, our understanding of how to maintain and restore biodiverse tropical grasslands (including savannas and open-canopy grassy woodlands) remains limited. To incorporate grasslands into large-scale restoration efforts, we synthesised existing ecological knowledge of tropical grassland resilience and approaches to plant community restoration. Tropical grassland plant communities are resilient to, and often dependent on, the endogenous disturbances with which they evolved - frequent fires and native megafaunal herbivory. In stark contrast, tropical grasslands are extremely vulnerable to human-caused exogenous disturbances, particularly those that alter soils and destroy belowground biomass (e.g. tillage agriculture, surface mining); tropical grassland restoration after severe soil disturbances is expensive and rarely achieves management targets. Where grasslands have been degraded by altered disturbance regimes (e.g. fire exclusion), exotic plant invasions, or afforestation, restoration efforts can recreate vegetation structure (i.e. historical tree density and herbaceous ground cover), but species-diverse plant communities, including endemic species, are slow to recover. Complicating plant-community restoration efforts, many tropical grassland species, particularly those that invest in underground storage organs, are difficult to propagate and re-establish. To guide restoration decisions, we draw on the old-growth grassland concept, the novel ecosystem concept, and theory regarding tree cover along resource gradients in savannas to propose a conceptual framework that classifies tropical grasslands into three broad ecosystem states. These states are: (1) old-growth grasslands (i.e. ancient, biodiverse grassy ecosystems), where management should focus on the maintenance of disturbance regimes; (2) hybrid grasslands, where restoration should emphasise a return towards the old-growth state; and (3) novel ecosystems, where the magnitude of environmental change (i.e. a shift to an alternative ecosystem state) or the socioecological context preclude a return to historical conditions.