Echoes of ancient introgression punctuate stable genomic lineages in the evolution of figs.

Studies investigating the evolution of flowering plants have long focused on isolating mechanisms such as pollinator specificity. Some recent studies have proposed a role for introgressive hybridization between species, recognizing that isolating processes such as pollinator specialization may not be complete barriers to hybridization. Occasional hybridization may therefore lead to distinct yet reproductively connected lineages. We investigate the balance between introgression and reproductive isolation in a diverse clade using a densely sampled phylogenomic study of fig trees (Ficus, Moraceae). Codiversification with specialized pollinating wasps (Agaonidae) is recognized as a major engine of fig diversity, leading to about 850 species. Nevertheless, some studies have focused on the importance of hybridization in Ficus, highlighting the consequences of pollinator sharing. Here, we employ dense taxon sampling (520 species) throughout Moraceae and 1,751 loci to investigate phylogenetic relationships and the prevalence of introgression among species throughout the history of Ficus. We present a well-resolved phylogenomic backbone for Ficus, providing a solid foundation for an updated classification. Our results paint a picture of phylogenetically stable evolution within lineages punctuated by occasional local introgression events likely mediated by local pollinator sharing, illustrated by clear cases of cytoplasmic introgression that have been nearly drowned out of the nuclear genome through subsequent lineage fidelity. The phylogenetic history of figs thus highlights that while hybridization is an important process in plant evolution, the mere ability of species to hybridize locally does not necessarily translate into ongoing introgression between distant lineages, particularly in the presence of obligate plant-pollinator relationships.

Root-associated fungal communities are influenced more by soils than by plant-host root traits in a Chinese tropical forest.

Forest fungal communities are shaped by the interactions between host tree root systems and the associated soil conditions. We investigated how the soil environment, root morphological traits, and root chemistry influence root-inhabiting fungal communities in three tropical forest sites of varying successional status in Xishuangbanna, China. For 150 trees of 66 species, we measured root morphology and tissue chemistry. Tree species identity was confirmed by sequencing rbcL, and root-associated fungal (RAF) communities were determined using high-throughput ITS2 sequencing. Using distance-based redundancy analysis and hierarchical variation partitioning, we quantified the relative importance of two soil variables (site average total phosphorus and available phosphorus), four root traits (dry matter content, tissue density, specific tip abundance, and forks), and three root tissue elemental concentrations (nitrogen, calcium, and manganese) on RAF community dissimilarity. The root and soil environment collectively explained 23% of RAF compositional variation. Soil phosphorus explained 76% of that variation. Twenty fungal taxa differentiated RAF communities among the three sites. Soil phosphorus most strongly affects RAF assemblages in this tropical forest. Variation in root calcium and manganese concentrations and root morphology among tree hosts, principally an architectural trade-off between dense, highly branched vs less-dense, herringbone-type root systems, are important secondary determinants.

Plant functional traits have globally consistent effects on competition.

Phenotypic traits and their associated trade-offs have been shown to have globally consistent effects on individual plant physiological functions, but how these effects scale up to influence competition, a key driver of community assembly in terrestrial vegetation, has remained unclear. Here we use growth data from more than 3 million trees in over 140,000 plots across the world to show how three key functional traits--wood density, specific leaf area and maximum height--consistently influence competitive interactions. Fast maximum growth of a species was correlated negatively with its wood density in all biomes, and positively with its specific leaf area in most biomes. Low wood density was also correlated with a low ability to tolerate competition and a low competitive effect on neighbours, while high specific leaf area was correlated with a low competitive effect. Thus, traits generate trade-offs between performance with competition versus performance without competition, a fundamental ingredient in the classical hypothesis that the coexistence of plant species is enabled via differentiation in their successional strategies. Competition within species was stronger than between species, but an increase in trait dissimilarity between species had little influence in weakening competition. No benefit of dissimilarity was detected for specific leaf area or wood density, and only a weak benefit for maximum height. Our trait-based approach to modelling competition makes generalization possible across the forest ecosystems of the world and their highly diverse species composition.

The global spectrum of plant form and function.

Earth is home to a remarkable diversity of plant forms and life histories, yet comparatively few essential trait combinations have proved evolutionarily viable in today's terrestrial biosphere. By analysing worldwide variation in six major traits critical to growth, survival and reproduction within the largest sample of vascular plant species ever compiled, we found that occupancy of six-dimensional trait space is strongly concentrated, indicating coordination and trade-offs. Three-quarters of trait variation is captured in a two-dimensional global spectrum of plant form and function. One major dimension within this plane reflects the size of whole plants and their parts; the other represents the leaf economics spectrum, which balances leaf construction costs against growth potential. The global plant trait spectrum provides a backdrop for elucidating constraints on evolution, for functionally qualifying species and ecosystems, and for improving models that predict future vegetation based on continuous variation in plant form and function.

Root anatomy helps to reconcile observed root trait syndromes in tropical tree species.

PREMISE: Studying the organization of functional traits in plant leaves and stems has revealed notable patterns linking function and form; however, evidence of similarly robust organization in root tissues remains controversial. We posit that anatomical traits in roots can provide insight on the overall organization of the root system. We hypothesized that size variation in the tissue outside the stele is related in a nonlinear fashion with functional traits associated with direct resource uptake, including a negative relationship with root architectural traits, and that similar relationships detected in tropical areas also hold true in other biomes. METHODS: We addressed our hypotheses using empirical data from 24 tropical tree species in French Guiana, including anatomical measurements in first order roots and functional trait description for the entire fine root system. In addition, we compiled a global meta-analysis of root traits for 500+ forest species across tropical, subtropical, and temperate forests. RESULTS: Our results supported the expected nonlinear relationships between cortical size and morphological traits and a negative linear trend with architectural traits. We confirmed a global negative relationship among specific root length (SRL), diameter, and tissue density, suggesting similar anatomical constraints in root systems across woody plants. However, the importance of factors varies across biomes, possibly related to the unequal phylogenetic representation across latitudes. CONCLUSIONS: Our findings imply that the rhizocentric hypothesis can be a valuable approach to understand fine root trait syndromes and the evolution of absorptive roots in vascular plants.

The physiological acclimation and growth response of Populus trichocarpa to warming.

Plant metabolic acclimation to thermal stress remains underrepresented in current global climate models. Gaps exist in our understanding of how metabolic processes (i.e., photosynthesis, respiration) acclimate over time and how aboveground versus belowground acclimation differs. We measured the thermal acclimation of Populus trichocarpa, comparing aboveground versus belowground physiology over time. Ninety genetically identical ramets were propagated in mesocosms that separated root and microbial components. After establishment at 25°C for 6 weeks, 60 clones were warmed +4 or +8°C and monitored for 10 weeks, measuring photosynthesis (A), leaf respiration (R), soil respiration (Rs ), root plus soil respiration (Rs+r ), and root respiration (Rr ). We observed thermal acclimation in both A and R, with rates initially increasing, then declining as the thermal photosynthetic optimum (Topt ) and the temperature-sensitivity (Q10 ) of respiration adjusted to warmer conditions. Photosynthetic acclimation was constructive, based on an increase in both Topt and peak A. Belowground, Rs+r decreased linearly with warming, while Rs rates declined abruptly, then remained constant with additional warming. Plant biomass was greatest at +4°C, with 30% allocated belowground. Rates of mass-based Rr were similar among treatments; however, root nitrogen declined at +8°C leading to less mass nitrogen-based Rr in that treatment. The Q10 -temperature relationship of Rr was affected by warming, leading to differing values among treatments. Aboveground acclimation exceeded belowground acclimation, and plant nitrogen-use mediated the acclimatory response. Results suggest that moderate climate warming (+4°C) may lead to acclimation and increased plant biomass production but increases in production could be limited with severe warming (+8°C).

Surprising low diversity of the plant pathogen Phytophthora in Amazonian forests.

The genus Phytophthora represents a group of plant pathogens with broad global distribution. The majority of them cause the collar and root-rot of diverse plant species. Little is known about Phytophthora communities in forest ecosystems, especially in the Neotropical forests where natural enemies could maintain the huge plant diversity via negative density dependence. We characterized the diversity of soil-borne Phytophthora communities in the North French Guiana rainforest and investigated how they are structured by host identity and environmental factors. In this little-explored habitat, 250 soil cores were sampled from 10 plots hosting 10 different plant families across three forest environments (Terra Firme, Seasonally Flooded and White Sand). Phytophthora diversity was studied using a baiting approach and metabarcoding (High-Throughput Sequencing) on environmental DNA extracted from both soil samples and baiting-leaves. These three approaches revealed very similar communities, characterized by an unexpected low diversity of Phytophthora species, with the dominance of two cryptic species close to Phytophthora heveae. As expected, the Phytophthora community composition of the French Guiana rainforest was significantly impacted by the host plant family and environment. However, these plant pathogen communities are very small and are dominated by generalist species, questioning their potential roles as drivers of plant diversity in these Amazonian forests.

Disentangling the effects of environment and ontogeny on tree functional dimensions for congeneric species in tropical forests.

Soil water and nutrient availability are key drivers of tree species distribution and forest ecosystem functioning, with strong species differences in water and nutrient use. Despite growing evidence for intraspecific trait differences, it remains unclear under which circumstances the effects of environmental gradients trump those of ontogeny and taxonomy on important functional dimensions related to resource use, particularly in tropical forests. Here, we explore how physiological, chemical, and morphological traits related to resource use vary between life stages in four species within the genus Micropholis that is widespread in lowland Amazonia. Specifically, we evaluate how environment, developmental stage, and taxonomy contribute to single-trait variation and multidimensional functional strategies. We find that environment, developmental stage, and taxonomy differentially contribute to functional dimensions. Habitats and seasons shape physiological and chemical traits related to water and nutrient use, whereas developmental stage and taxonomic identity impact morphological traits -especially those related to the leaf economics spectrum. Our findings suggest that combining environment, ontogeny, and taxonomy allows for a better understanding of important functional dimensions in tropical trees and highlights the need for integrating tree physiological and chemical traits with classically used morphological traits to improve predictions of tropical forests' responses to environmental change.

Independent evolutionary changes in fine-root traits among main clades during the diversification of seed plants.

Changes in fine-root morphology are typically associated with transitions from the ancestral arbuscular mycorrhizal (AM) to the alternative ectomycorrhizal (ECM) or nonmycorrhizal (NM) associations. However, the modifications in root morphology may also coincide with new modifications in leaf hydraulics and growth habit during angiosperm diversification. These hypotheses have not been evaluated concurrently, and this limits our understanding of the causes of fine-root evolution. To explore the evolution of fine-root systems, we assembled a 600+ species database to reconstruct historical changes in seed plants over time. We utilise ancestral reconstruction approaches together with phylogenetically informed comparative analyses to test whether changes in fine-root traits were most strongly associated with mycorrhizal affiliation, leaf hydraulics or growth form. Our findings showed significant shifts in root diameter, specific root length and root tissue density as angiosperms diversified, largely independent from leaf changes or mycorrhizal affiliation. Growth form was the only factor associated with fine-root traits in statistical models including mycorrhizal association and leaf venation, suggesting substantial modifications in fine-root morphology during transitions from woody to nonwoody habits. Divergences in fine-root systems were crucial in the evolution of seed plant lineages, with important implications for ecological processes in terrestrial ecosystems.

Compositional response of Amazon forests to climate change.

Most of the planet's diversity is concentrated in the tropics, which includes many regions undergoing rapid climate change. Yet, while climate-induced biodiversity changes are widely documented elsewhere, few studies have addressed this issue for lowland tropical ecosystems. Here we investigate whether the floristic and functional composition of intact lowland Amazonian forests have been changing by evaluating records from 106 long-term inventory plots spanning 30 years. We analyse three traits that have been hypothesized to respond to different environmental drivers (increase in moisture stress and atmospheric CO2 concentrations): maximum tree size, biogeographic water-deficit affiliation and wood density. Tree communities have become increasingly dominated by large-statured taxa, but to date there has been no detectable change in mean wood density or water deficit affiliation at the community level, despite most forest plots having experienced an intensification of the dry season. However, among newly recruited trees, dry-affiliated genera have become more abundant, while the mortality of wet-affiliated genera has increased in those plots where the dry season has intensified most. Thus, a slow shift to a more dry-affiliated Amazonia is underway, with changes in compositional dynamics (recruits and mortality) consistent with climate-change drivers, but yet to significantly impact whole-community composition. The Amazon observational record suggests that the increase in atmospheric CO2 is driving a shift within tree communities to large-statured species and that climate changes to date will impact forest composition, but long generation times of tropical trees mean that biodiversity change is lagging behind climate change.

Habitats shape taxonomic and functional composition of Neotropical ant assemblages.

Determining assembly rules of co-occurring species persists as a fundamental goal in community ecology. At local scales, the relative importance of environmental filtering vs. competitive exclusion remains a subject of debate. In this study, we assessed the relative importance of habitat filtering and competition in structuring understory ant communities in tropical forests of French Guiana. Leaf-litter ants were collected using pitfall and Winkler traps across swamp, slope and plateau forests near Saül, French Guiana. We used a combination of univariate and multivariate analyses to evaluate trait response of ants to habitat characteristics. Null model analyses were used to investigate the effects of habitat filtering and competitive interactions on community assembly at the scale of assemblages and sampling points, respectively. Swamp forests presented a much lower taxonomic and functional richness compared to slope and plateau forests. Furthermore, marked differences in taxonomic and functional composition were observed between swamp forests and slope or plateau forests. We found weak evidence for competitive exclusion based on null models. Nevertheless, the contrasting trait composition observed between habitats revealed differences in the ecological attributes of the species in the different forest habitats. Our analyses suggest that competitive interactions may not play an important role in structuring leaf-litter ant assemblages locally. Rather, habitats are responsible for driving both taxonomic and functional composition of ant communities.

Precipitation mediates sap flux sensitivity to evaporative demand in the neotropics.

Transpiration in humid tropical forests modulates the global water cycle and is a key driver of climate regulation. Yet, our understanding of how tropical trees regulate sap flux in response to climate variability remains elusive. With a progressively warming climate, atmospheric evaporative demand [i.e., vapor pressure deficit (VPD)] will be increasingly important for plant functioning, becoming the major control of plant water use in the twenty-first century. Using measurements in 34 tree species at seven sites across a precipitation gradient in the neotropics, we determined how the maximum sap flux velocity (vmax) and the VPD threshold at which vmax is reached (VPDmax) vary with precipitation regime [mean annual precipitation (MAP); seasonal drought intensity (PDRY)] and two functional traits related to foliar and wood economics spectra [leaf mass per area (LMA); wood specific gravity (WSG)]. We show that, even though vmax is highly variable within sites, it follows a negative trend in response to increasing MAP and PDRY across sites. LMA and WSG exerted little effect on vmax and VPDmax, suggesting that these widely used functional traits provide limited explanatory power of dynamic plant responses to environmental variation within hyper-diverse forests. This study demonstrates that long-term precipitation plays an important role in the sap flux response of humid tropical forests to VPD. Our findings suggest that under higher evaporative demand, trees growing in wetter environments in humid tropical regions may be subjected to reduced water exchange with the atmosphere relative to trees growing in drier climates.

Coordination and trade-offs among hydraulic safety, efficiency and drought avoidance traits in Amazonian rainforest canopy tree species.

Predicting responses of tropical forests to climate change-type drought is challenging because of high species diversity. Detailed characterization of tropical tree hydraulic physiology is necessary to evaluate community drought vulnerability and improve model parameterization. Here, we measured xylem hydraulic conductivity (hydraulic efficiency), xylem vulnerability curves (hydraulic safety), sapwood pressure-volume curves (drought avoidance) and wood density on emergent branches of 14 common species of Eastern Amazonian canopy trees in Paracou, French Guiana across species with the densest and lightest wood in the plot. Our objectives were to evaluate relationships among hydraulic traits to identify strategies and test the ability of easy-to-measure traits as proxies for hard-to-measure hydraulic traits. Xylem efficiency was related to capacitance, sapwood water content and turgor loss point, and other drought avoidance traits, but not to xylem safety (P50 ). Wood density was correlated (r = -0.57 to -0.97) with sapwood pressure-volume traits, forming an axis of hydraulic strategy variation. In contrast to drier sites where hydraulic safety plays a greater role, tropical trees in this humid tropical site varied along an axis with low wood density, high xylem efficiency and high capacitance at one end of the spectrum, and high wood density and low turgor loss point at the other.

Understanding the recruitment response of juvenile Neotropical trees to logging intensity using functional traits.

Selective logging remains a widespread practice in tropical forests, yet the long-term effects of timber harvest on juvenile tree (i.e., sapling) recruitment across the hundreds of species occurring in most tropical forests remain difficult to predict. This uncertainty could potentially exacerbate threats to some of the thousands of timber-valuable tree species in the Amazon. Our objective was to determine to what extent long-term responses of tree species regeneration in logged forests can be explained by their functional traits. We integrate functional trait data for 13 leaf, stem, and seed traits from 25 canopy tree species with a range of life histories, such as the pioneer Goupia glabra and the shade-tolerant Iryanthera hostmannii, together with over 30 yr of sapling monitoring in permanent plots spanning a gradient of harvest intensity at the Paracou Forest Disturbance Experiment (PFDE), French Guiana. We anticipated that more intensive logging would increase recruitment of pioneer species with higher specific leaf area, lower wood densities, and smaller seeds, due to the removal of canopy trees. We define a recruitment response metric to compare sapling regeneration to timber harvest intensity across species. Although not statistically significant, sapling recruitment decreased with logging intensity for eight of 23 species and these species tended to have large seeds and dense wood. A generalized linear mixed model fit using specific leaf area, seed mass, and twig density data explained about 45% of the variability in sapling dynamics. Effects of specific leaf area outweighed those of seed mass and wood density in explaining recruitment dynamics of the sapling community in response to increasing logging intensity. The most intense treatment at the PFDE, which includes stand thinning of non-timber-valuable adult trees and poison-girdling for competitive release, showed evidence of shifting community composition in sapling regeneration at the 30-yr mark, toward species with less dense wood, lighter seeds, and higher specific leaf area. Our results indicate that high-intensity logging can have lasting effects on stand regeneration dynamics and that functional traits can help simplify general trends of sapling recruitment for highly diverse logged tropical forests.

Geographical Variation in Community Divergence: Insights from Tropical Forest Monodominance by Ectomycorrhizal Trees.

Convergence occurs in both species traits and community structure, but how convergence at the two scales influences each other remains unclear. To address this question, we focus on tropical forest monodominance, in which a single, often ectomycorrhizal (EM) tree species occasionally dominates forest stands within a landscape otherwise characterized by diverse communities of arbuscular mycorrhizal (AM) trees. Such monodominance is a striking potential example of community divergence resulting in alternative stable states. However, it is observed only in some tropical regions. A diverse suite of AM and EM trees locally codominate forest stands elsewhere. We develop a hypothesis to explain this geographical difference using a simulation model of plant community assembly. Simulation results suggest that in a region with a few EM species (e.g., South America), EM trees experience strong selection for convergent traits that match the abiotic conditions of the environment. Consequently, EM species successfully compete against other species to form monodominant stands via positive plant-soil feedbacks. By contrast, in a region with many EM species (e.g., Southeast Asia), species maintain divergent traits because of complex plant-soil feedbacks, with no species having traits that enable monodominance. An analysis of plant trait data from Borneo and Peruvian Amazon was inconclusive. Overall, this work highlights the utility of geographical comparison in understanding the relationship between trait convergence and community convergence.

There's no place like home: seedling mortality contributes to the habitat specialisation of tree species across Amazonia.

Understanding the mechanisms generating species distributions remains a challenge, especially in hyperdiverse tropical forests. We evaluated the role of rainfall variation, soil gradients and herbivory on seedling mortality, and how variation in seedling performance along these gradients contributes to habitat specialisation. In a 4-year experiment, replicated at the two extremes of the Amazon basin, we reciprocally transplanted 4638 tree seedlings of 41 habitat-specialist species from seven phylogenetic lineages among the three most important forest habitats of lowland Amazonia. Rainfall variation, flooding and soil gradients strongly influenced seedling mortality, whereas herbivory had negligible impact. Seedling mortality varied strongly among habitats, consistent with predictions for habitat specialists in most lineages. This suggests that seedling performance is a primary determinant of the habitat associations of adult trees across Amazonia. It further suggests that tree diversity, currently mostly harboured in terra firme forests, may be strongly impacted by the predicted climate changes in Amazonia.

Rare species contribute disproportionately to the functional structure of species assemblages.

There is broad consensus that the diversity of functional traits within species assemblages drives several ecological processes. It is also widely recognized that rare species are the first to become extinct following human-induced disturbances. Surprisingly, however, the functional importance of rare species is still poorly understood, particularly in tropical species-rich assemblages where the majority of species are rare, and the rate of species extinction can be high. Here, we investigated the consequences of local and regional extinctions on the functional structure of species assemblages. We used three extensive datasets (stream fish from the Brazilian Amazon, rainforest trees from French Guiana, and birds from the Australian Wet Tropics) and built an integrative measure of species rarity versus commonness, combining local abundance, geographical range, and habitat breadth. Using different scenarios of species loss, we found a disproportionate impact of rare species extinction for the three groups, with significant reductions in levels of functional richness, specialization, and originality of assemblages, which may severely undermine the integrity of ecological processes. The whole breadth of functional abilities within species assemblages, which is disproportionately supported by rare species, is certainly critical in maintaining ecosystems particularly under the ongoing rapid environmental transitions.

Evolutionary heritage influences Amazon tree ecology.

Lineages tend to retain ecological characteristics of their ancestors through time. However, for some traits, selection during evolutionary history may have also played a role in determining trait values. To address the relative importance of these processes requires large-scale quantification of traits and evolutionary relationships among species. The Amazonian tree flora comprises a high diversity of angiosperm lineages and species with widely differing life-history characteristics, providing an excellent system to investigate the combined influences of evolutionary heritage and selection in determining trait variation. We used trait data related to the major axes of life-history variation among tropical trees (e.g. growth and mortality rates) from 577 inventory plots in closed-canopy forest, mapped onto a phylogenetic hypothesis spanning more than 300 genera including all major angiosperm clades to test for evolutionary constraints on traits. We found significant phylogenetic signal (PS) for all traits, consistent with evolutionarily related genera having more similar characteristics than expected by chance. Although there is also evidence for repeated evolution of pioneer and shade tolerant life-history strategies within independent lineages, the existence of significant PS allows clearer predictions of the links between evolutionary diversity, ecosystem function and the response of tropical forests to global change.

Variation in stem mortality rates determines patterns of above-ground biomass in Amazonian forests: implications for dynamic global vegetation models.

Understanding the processes that determine above-ground biomass (AGB) in Amazonian forests is important for predicting the sensitivity of these ecosystems to environmental change and for designing and evaluating dynamic global vegetation models (DGVMs). AGB is determined by inputs from woody productivity [woody net primary productivity (NPP)] and the rate at which carbon is lost through tree mortality. Here, we test whether two direct metrics of tree mortality (the absolute rate of woody biomass loss and the rate of stem mortality) and/or woody NPP, control variation in AGB among 167 plots in intact forest across Amazonia. We then compare these relationships and the observed variation in AGB and woody NPP with the predictions of four DGVMs. The observations show that stem mortality rates, rather than absolute rates of woody biomass loss, are the most important predictor of AGB, which is consistent with the importance of stand size structure for determining spatial variation in AGB. The relationship between stem mortality rates and AGB varies among different regions of Amazonia, indicating that variation in wood density and height/diameter relationships also influences AGB. In contrast to previous findings, we find that woody NPP is not correlated with stem mortality rates and is weakly positively correlated with AGB. Across the four models, basin-wide average AGB is similar to the mean of the observations. However, the models consistently overestimate woody NPP and poorly represent the spatial patterns of both AGB and woody NPP estimated using plot data. In marked contrast to the observations, DGVMs typically show strong positive relationships between woody NPP and AGB. Resolving these differences will require incorporating forest size structure, mechanistic models of stem mortality and variation in functional composition in DGVMs.

Rare species support vulnerable functions in high-diversity ecosystems.

Around the world, the human-induced collapses of populations and species have triggered a sixth mass extinction crisis, with rare species often being the first to disappear. Although the role of species diversity in the maintenance of ecosystem processes has been widely investigated, the role of rare species remains controversial. A critical issue is whether common species insure against the loss of functions supported by rare species. This issue is even more critical in species-rich ecosystems where high functional redundancy among species is likely and where it is thus often assumed that ecosystem functioning is buffered against species loss. Here, using extensive datasets of species occurrences and functional traits from three highly diverse ecosystems (846 coral reef fishes, 2,979 alpine plants, and 662 tropical trees), we demonstrate that the most distinct combinations of traits are supported predominantly by rare species both in terms of local abundance and regional occupancy. Moreover, species that have low functional redundancy and are likely to support the most vulnerable functions, with no other species carrying similar combinations of traits, are rarer than expected by chance in all three ecosystems. For instance, 63% and 98% of fish species that are likely to support highly vulnerable functions in coral reef ecosystems are locally and regionally rare, respectively. For alpine plants, 32% and 89% of such species are locally and regionally rare, respectively. Remarkably, 47% of fish species and 55% of tropical tree species that are likely to support highly vulnerable functions have only one individual per sample on average. Our results emphasize the importance of rare species conservation, even in highly diverse ecosystems, which are thought to exhibit high functional redundancy. Rare species offer more than aesthetic, cultural, or taxonomic diversity value; they disproportionately increase the potential breadth of functions provided by ecosystems across spatial scales. As such, they are likely to insure against future uncertainty arising from climate change and the ever-increasing anthropogenic pressures on ecosystems. Our results call for a more detailed understanding of the role of rarity and functional vulnerability in ecosystem functioning.