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Whether forests are composed of evergreen or deciduous species largely affects biogeochemical cycles and the functioning, structure and biodiversity of ecosystems. These leaf phenology types may be self-promoted through positive plant-soil feedbacks, which may shift the distribution of forest types towards alternative stable states. However, we still lack empirical evidence of phenological alternative stable states and a spatial understanding of where they might be present. Here, we test the presence of alternative stable states using forest inventory data at the continental (North America and Europe) and global scale. We show that the distribution of forest leaf phenology types is bimodal, and demonstrate the presence of positive feedbacks in recruitment, growth and mortality. Data-driven simulations show that the observed positive feedbacks are sufficient and necessary to produce the alternative stable states, which also lead to hysteresis during ecosystem transition. Spatial random forest models further reveal hotspots of alternative stable states in evergreen-deciduous ecotones and the poleward range limit of forests, which appear largely driven by soil feedbacks. Given the close connection between forest leaf phenology and ecosystem biogeochemical processes, our insights on evergreen vs. deciduous alternative stable states inform our understanding of the distribution of forest biomes, allowing more accurate quantification of carbon turnover and terrestrial climate feedbacks.

The biodiversity-productivity relationship (BPR) is central to our understanding of ecosystem functioning and restoration practices. Quantifying variation in the BPR across environmental gradients is thus critical for a spatially-explicit understanding of this key ecosystem property. Here, by integrating plot-level tree occurrence information from the Global Forest Biodiversity initiative (GFBi), satellite-derived net primary productivity, and environmental covariates, we estimated global variation in the BPR of forests along spatial and environmental gradients. The results show that variation in the forest BPR correlates with temperature and water availability, leading to significant differences in the forest BPR across biomes: the highest positive BPR occurs in arid and boreal forests, the lowest BPR in temperate broadleaved and mixed forests. In addition, forest age played a key role in mediating the BPR, with no BPR found in young forests (

Invasive species utilize a wide array of trait strategies to establish in novel ecosystems. Among these traits is the capacity to produce allelopathic compounds that can directly inhibit neighboring native plants or indirectly suppress native plants via disruption of beneficial belowground microbial mutualisms, or altered soil resources. Despite the well-known prevalence of allelopathy among plant taxa, the pervasiveness of allelopathy among invasive plants is unknown. Here we demonstrate that the majority of the 524 invasive plant species in our database produce allelochemicals with the potential to negatively affect native plant performance. Moreover, allelopathy is widespread across the plant phylogeny, suggesting that allelopathy could have a large impact on native species across the globe. Allelopathic impacts of invasive species are often thought to be present in only a few plant clades (e.g., Brassicaceae). Yet our analysis shows that allelopathy is present in 72% of the 113 plant families surveyed, suggesting that this ubiquitous mechanism of invasion deserves more attention as invasion rates increase across the globe.

The length of the growing season in temperate and boreal forests has a strong effect on the global carbon balance. Yet, our current understanding of the drivers of phenological processes such as autumn leaf senescence in deciduous trees is not sufficient for making reliable estimates of future growing-season lengths under climate change. While temperature has been shown to be an important driver of autumn leaf senescence, recent evidence suggests that the concept of carbon sink limitation might help to reduce unexplained variation in leaf senescence predictions. According to the carbon sink limitation hypothesis, senescence is regulated by the balance of the plant carbon source and the plant carbon sink, so that senescence occurs later when carbon inputs (source) are low and earlier when there is a low carbon demand (sink). In our experiment, we manipulated carbon source–sink dynamics in birch seedlings (Betula pendula L.) to evaluate the evidence for an effect of carbon sink limitation on autumn leaf senescence in a widespread deciduous tree. Specifically, we removed leaves and/or buds from the seedlings and monitored the effects on autumn net photosynthesis and leaf senescence. In agreement with the carbon sink limitation hypothesis, we observed that a decrease in the carbon source through a high degree of leaf removal increased autumn leaf-level photosynthesis by ~14% and postponed senescence by 5.5 ± 2.4 days. Yet, we did not see significant effects of the lower- and medium-degree defoliation treatments. Further, we did not observe an effect of bud removal on either photosynthesis or senescence, which was likely caused by the fact that our bud removal treatment did not considerably change the plant carbon sink. At least partly, our results are in line with the hypothesis of carbon sink limitation as a driver of growing-season length and move the scientific field closer to narrowing a main uncertainty in climate change predictions.

The degree to which elevated CO

The degree to which elevated CO2 concentrations (e[CO2]) increase the amount of carbon (C) assimilated by vegetation plays a key role in climate change. However, due to the short-term nature of CO2 enrichment experiments and the lack of reconciliation between different ecological scales, the effect of e[CO2] on plant biomass stocks remains a major uncertainty in future climate projections. Here, we review the effect of e[CO2] on plant biomass across multiple levels of ecological organization, scaling from physiological responses to changes in population-, community-, ecosystem-, and global-scale dynamics. We find that evidence for a sustained biomass response to e[CO2] varies across ecological scales, leading to diverging conclusions about the responses of individuals, populations, communities, and ecosystems. While the distinct focus of every scale reveals new mechanisms driving biomass accumulation under e[CO2], none of them provides a full picture of all relevant processes. For example, while physiological evidence suggests a possible long-term basis for increased biomass accumulation under e[CO2] through sustained photosynthetic stimulation, population-scale evidence indicates that a possible e[CO2]-induced increase in mortality rates might potentially outweigh the effect of increases in plant growth rates on biomass levels. Evidence at the global scale may indicate that e[CO2] has contributed to increased biomass cover over recent decades, but due to the difficulty to disentangle the effect of e[CO2] from a variety of climatic and land-use-related drivers of plant biomass stocks, it remains unclear whether nutrient limitations or other ecological mechanisms operating at finer scales will dampen the e[CO2] effect over time. By exploring these discrepancies, we identify key research gaps in our understanding of the effect of e[CO2] on plant biomass and highlight the need to integrate knowledge across scales of ecological organization so that large-scale modeling can represent the finer-scale mechanisms needed to constrain our understanding of future terrestrial C storage.

Due to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. When considering trait relationships across the entire plant kingdom, plant trait frameworks typically must omit traits unique to large woody species, thereby limiting our understanding of how these distinct ecological pressures shape trait relationships in trees. Here, by considering 18 functional traits—reflecting leaf economics, wood structure, tree size, reproduction, and below-ground allocation—we quantify the major axes of variation governing trait expression of trees worldwide. We show that trait variation within and across angiosperms and gymnosperms is captured by two independent processes: one reflecting tree size and competition for light, the other reflecting leaf photosynthetic capacity and nutrient economies. By exploring multidimensional relationships across clusters of traits, we further identify a representative set of seven traits which captures the majority of variation in form and function in trees: maximum tree height, stem conduit diameter, specific leaf area, seed mass, bark thickness, root depth, and wood density. Collectively, this work informs future trait-based research into the functional biogeography of trees, and contributes to our fundamental understanding of the ecological and evolutionary controls on forest biodiversity and productivity worldwide.

Biodiversity patterns emerge as a consequence of evolutionary and ecological processes. Their relative importance is frequently tested on model ecosystems such as oceanic islands that vary in both. However, the coarse-scale data typically used in biogeographic studies have limited inferential power to separate the effects of historical biogeographic factors (e.g., island age) from the effects of ecological ones (e.g., island area and habitat heterogeneity). Here, we describe local-scale biodiversity patterns of woody plants using a database of more than 500 forest plots from across the Hawaiian archipelago, where these volcanic islands differ in age by several million years. We show that, after controlling for factors such as island area and heterogeneity, the oldest islands (Kaua’i and O’ahu) have greater native species diversity per unit area than younger islands (Maui and Hawai’i), indicating an important role for macroevolutionary processes in driving not just whole-island differences in species diversity, but also local community assembly. Further, we find that older islands have a greater number of rare species that are more spatially clumped (i.e., higher within-island β-diversity) than younger islands. When we included alien species in our analyses, we found that the signal of macroevolutionary processes via island age was diluted. Our approach allows a more explicit test of the question of how macroevolutionary factors shape not just regional-scale biodiversity, but also local-scale community assembly patterns and processes in a model archipelago ecosystem, and it can be applied to disentangle biodiversity drivers in other systems.

Background: Climates are changing at a rate that exceeds the adaptive capacity of species, especially endangered species. Genetic variation and phenotypic plasticity are important for population persistence, yet few studies have linked traits to seedling performance under drought in endangered species. Aims: We assessed intraspecific variation and trait plasticity under drought, to understand how an endangered species would cope with increasingly severe droughts. Methods: Using greenhouse experiments, we assessed drought tolerance of a critically endangered Hawaiian shrub, Schiedea obovata. Seedlings from five maternal families (three selfed and two interpopulation crosses) were subjected to daily watering or a simulated drought, and we measured growth, traits linked to drought tolerance, and days until death under terminal drought. Results: Drought reduced growth but not days until death. We detected genetic variation in growth, but no trait plasticity except in carbon:nitrogen, which decreased under drought. We did not detect traits that enhanced performance under drought but identified four physiological traits whose effects on growth varied under control and drought. Conclusions: Our results indicate moderate drought tolerance of an endangered shrub, and low trait plasticity. Conservation of endangered species under shifting climates will benefit from studies of stress tolerance, particularly at the vulnerable seedling stage.

The mechanisms causing invasive species impact are rarely empirically tested, limiting our ability to understand and predict subsequent changes in invaded plant communities. Invader disruption of native mutualistic interactions is a mechanism expected to have negative effects on native plant species. Specifically, disruption of native plant‐fungal mutualisms may provide non‐mycorrhizal plant invaders an advantage over mycorrhizal native plants. Invasive Alliaria petiolata (garlic mustard) produces secondary chemicals toxic to soil microorganisms including mycorrhizal fungi, and is known to induce physiological stress and reduce population growth rates of native forest understory plant species. Here, we report on a 11‐yr manipulative field experiment in replicated forest plots testing if the effects of removal of garlic mustard on the plant community support the mutualism disruption hypothesis within the entire understory herbaceous community. We compare community responses for two functional groups: the mycorrhizal vs. the non‐mycorrhizal plant communities. Our results show that garlic mustard weeding alters the community composition, decreases community evenness, and increases the abundance of understory herbs that associate with mycorrhizal fungi. Conversely, garlic mustard has no significant effects on the non‐mycorrhizal plant community. Consistent with the mutualism disruption hypothesis, our results demonstrate that allelochemical producing invaders modify the plant community by disproportionately impacting mycorrhizal plant species. We also demonstrate the importance of incorporating causal mechanisms of biological invasion to elucidate patterns and predict community‐level responses.

Despite widespread evidence that biological invasion influences both the biotic and abiotic soil environments, the extent to which these two pathways underpin the effects of invasion on plant traits and performance remains unknown. Leveraging a long-term (14-year) field experiment, we show that an allelochemical-producing invader affects plants through biotic mechanisms, altering the soil fungal community composition, with no apparent shifts in soil nutrient availability. Changes in belowground fungal communities resulted in high costs of nutrient uptake for native perennials and a shift in plant traits linked to their water and nutrient use efficiencies. Some plants in the invaded community compensate for the disruption of nutritional symbionts and reduced nutrient provisioning by sanctioning more nitrogen to photosynthesis and expending more water, which demonstrates a trade-off in trait investment. For the first time, we show that the disruption of belowground nutritional symbionts can drive plants towards alternative regions of their trait space in order to maintain water and nutrient economics.

Despite widespread evidence that biological invasion influences the biotic and abiotic soil environments, the extent to which each of these pathways underpins the effects of invasion on native plant traits and performance is unknown. Leveraging a long-term (14-yr) manipulative field experiment, we show that an allelochemical-producing invader, Alliaria petiolata, affects native plants through biotic mechanisms, altering the soil fungal community composition, with no apparent shifts in soil nutrient availability. These changes in belowground soil fungal communities resulted in a high cost of resource uptake for native forest perennial herbs and a shift in functional traits linked to their carbon and nutrient economies. Furthermore, we illustrate that some species in the invaded community compensate for high nutrient costs by reducing nutrient uptake and maintaining photosynthesis by expending more water. This demonstrates a trade-off in trait investment that increases nutrient use efficiency as nutrient costs increase. Our results show that invasion-induced disruptions in the soil fungal community belowground can cascade to affect aboveground plant communities via shifts in physiological traits needed to maintain plant water and nutrient economies. These complex above-belowground linkages suggest that plant invasions should be evaluated at the system-level to fully understand and predict their impact on native plants and communities.

The invasive plant, garlic mustard (Alliaria petiolata), has the potential to affect soil microbial communities and ecosystem processes in temperate hardwood forests primarily through the release of allelopathic chemicals into the soil. These forest soils are also often affected (directly and indirectly) by the high abundance of white-tailed deer (Odocoileus virginianus), which can alter plant community composition and productivity. We examined the joint effects of deer and garlic mustard on soil microbial communities, soil nutrients and a native plant species’ vital rates in a temperate forest 8 years after initiation of a paired plot deer exclusion/access study where garlic mustard was either removed from half of each plot or remained at ambient level in the other plot half. We examined soil microbial communities using DNA-based techniques and quantified nutrient availability and physicochemical properties. Deer exclusion affected the community structure of AM fungi, particularly when garlic mustard was present, but had no effect on soil chemistry. Garlic mustard removal plots showed no changes for soil fungi, but displayed higher soil carbon content. Interestingly, we found significant changes to native plant vital rates that mirrored soil responses; the presence of garlic mustard led to higher mortality of large, mature plants and reduced native plant cover and biomass. Our data suggest herbivore-plant-soil feedbacks and synergies can interact to negatively affect the soil ecology of forests. Management activities that reduce deer or invasive plant abundance may positively affect soil microbial communities and chemistry in temperate forests.

Environmental conditions impose restrictions and costs on reproduction. Multiple reproductive options exist when increased reproductive costs drive plant populations toward alternative reproductive strategies. Using 4 years of demographic data across a deer impact gradient, where deer alter the abiotic environment, we parameterize a size-dependent integral projection model for a sexually labile and unpalatable forest perennial to investigate the demographic processes driving differentiation in the operational sex ratio (OSR) of local populations. In addition to a relative increase in asexual reproduction, our results illustrate that nontrophic indirect effects by overabundant deer on this perennial result in delayed female sex expression to unsustainably large plant sizes and lead to more pronounced plant shrinkage following female sex expression, effectively increasing the cost of reproduction. Among plants of reproductive age, increased deer impact decreases the size-dependent probability of flowering and reduces reproductive consistency over time. This pattern in sex expression skews populations toward female-biased OSRs at low deer impact sites and male-biased OSRs at intermediate and high deer impact sites. While this shift toward a male-biased OSR may ameliorate pollen limitation, it also decreases the effective population size when coupled with increased asexual reproduction. The divergence of reproductive strategies and reduced lifetime fitness in response to indirect deer impacts illustrate the persistent long-term effects of overabundant herbivores on unpalatable understory perennials.

Summary Understanding of the role of environmental change in the decline of endangered species is critical to designing scale-appropriate restoration plans. For locally endemic rare plants on the brink of extinction, frugivory can drastically reduce local recruitment by dispersing seeds away from geographically isolated populations. Dispersal of seeds away from isolated populations can ultimately lead to population decline. For localized endemic plants, fine-scale changes in microhabitat can further limit population persistence. Evaluating the individual and combined impact of frugivores and microhabitat heterogeneity on the short-term (i.e. transient) and long-term (i.e. asymptotic) dynamics of plants will provide insight into the drivers of species rarity. In this study, we used four years of demographic data to develop matrix projection models for a long-lived shrub, Cyrtandra dentata (H. St. John & Storey) (Gesneriaceae), which is endemic to the island of O'ahu in Hawai'i. Furthermore, we evaluated the individual and combined influence of a non-native frugivorous bird, Leiothrix lutea, and microhabitat heterogeneity on the short-term and long-term C. dentata population dynamics. Frugivory by L. lutea decreased the short-term and long-term population growth rates. However, under the current level of frugivory at the field site the C. dentata population was projected to persist over time. Conversely, the removal of optimum microhabitat for seedling establishment (i.e. rocky gulch walls and boulders in the gulch bottom) reduced the short-term and long-term population growth rates from growing to declining. Survival of mature C. dentata plants had the greatest influence on long-term population dynamics, followed by the growth of seedlings and immature plants. The importance of mature plant survival was even greater when we simulated the combined effect of frugivory and the loss of optimal microhabitat, relative to population dynamics based on field conditions. In the short-term (10 years), however, earlier life stages had the greatest influence on population growth rate. Synthesis and applications. This study emphasizes how important it is to decouple rare plant management strategies in the short versus long-term in order to prioritize restoration actions, particularly when faced with multiple stressors not all of which can be feasibly managed. From an applied conservation perspective, our findings also illustrate that the life stage that, if improved by management, would have the greatest influence on population dynamics is dependent on the timeframe of interest and initial conditions of the population. This article is protected by copyright. All rights reserved.

Rare species across taxonomic groups and biomes commonly suffer from multiple threats and require intensive restoration, including population reintroduction and threat control. Following reintroduction, it is necessary to identify what level of threat control is needed for species to persist over time. Population reintroduction and threat control are time intensive and costly. Thus, it is pragmatic to develop economically efficient restoration strategies. We combined transfer function analysis and economic cost analysis to evaluate the effects of biologically meaningful increases in demographic processes on the persistence of a reintroduced population of a Hawaii endemic long-lived shrub, Delissea waianaeensis. We show that an increase in fertility by 0.419 following the suppression of non-native rodents or an increase by 0.098 in seedling growth following the suppression of invasive molluscs would stabilize the population (i.e., λ = 1). Though a greater increase in fertility than seedling growth was needed for the reintroduced population to persist over time, increasing fertility by suppressing rodents was the most cost effective restoration strategy. Our study emphasizes the importance of considering the effects of large increases in plant vital rates in population projections and incorporating the economic cost of management actions in demographic models when developing restoration plans for endangered species.

Climate projections forecast more extreme interannual climate variability over time, with an increase in the severity and duration of extreme drought and rainfall events. Based on bioclimatic envelope models, it is projected that changing precipitation patterns will drastically alter the spatial distributions and density of plants and be a primary driver of biodiversity loss. However, many other underlying mechanisms can impact plant vital rates (i.e., survival, growth, and reproduction) and population dynamics. In this study, we developed a size‐dependent integral projection model (IPM) to evaluate how interannual precipitation and mollusk herbivory influence the dynamics of a Hawaii endemic short‐lived shrub, Schiedea obovata (Caryophyllaceae). Assessing how wet season precipitation effects population dynamics it critical, as it is the timeframe when most of the foliar growth occurs, plants flower and fruit, and seedlings establish. Temporal variation in wet season precipitation had a greater effect than mollusk herbivory on S. obovata population growth rate λ, and the impact of interannual precipitation on vital rates shifted across plant ontogeny. Furthermore, wet season precipitation influenced multiple vital rates in contrasting ways and the effect of precipitation on the survival of larger vegetative and reproductively mature individuals contributed the most to variation in the population growth rate. Among all combination of wet season precipitation and herbivory intensities, the only scenario that led to a growing population was when high wet precipitation was associated with low herbivory. Our study highlights the importance of evaluating how abiotic factors and plant–consumer interactions influence an organism across its life cycle to fully understand the underpinning mechanisms that structure its spatial and temporal distribution and abundance. Our results also illustrate that for short‐lived species, like S. obovata, seedling herbivory can have less of an effect on the dynamics of plant populations than decreased interannual precipitation.

Large-scale rodent control can help to manage endangered species that are vulnerable to invasive rodent consumption. A 26 ha rodent snap-trap grid was installed in montane forest on Oahu Island, Hawaii, in order to protect endangered snails and plants. To assess the effectiveness of this trapping operation in reducing fruit consumption and seed predation of the endangered Hawaiian lobeliad, Cyanea superba subsp. superba, pre- and post-dispersal C. superba fruit consumption were monitored for 36 plants at the site with rodent control (Kahanahaiki) and 42 plants at an adjacent site without rodent control (Pahole). Over 47 % of all monitored fruit were eaten on the plants at Pahole compared to 4 % at Kahanahaiki. Images captured using motion-sensing cameras suggest that black rats (Rattus rattus) were the only pre-dispersal fruit consumers. To quantify post-dispersal fruit consumption, and to identify the culprit frugivore(s), mature fruit were placed in tracking tunnels positioned on the forest floor and checked daily. At Pahole, all of the fruit were consumed by rats compared to 29 % at Kahanahaiki. Lastly, to determine if rodents from the sites were predators or dispersers of C. superba seed, fruit were fed to captive black rats and house mice (Mus musculus). Black rats consumed entire fruit, killing all the seed, while mice did little damage to the fruit and seed. Therefore, large-scale rat trapping can directly benefit the reproduction of C. superba subsp. superba. Controlling black rats at restoration sites appears integral to the successful restoration of this endangered plant species.