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Biodiversity: niches work If organisms are involved in a perpetual struggle for existence, how is it that communities are so diverse? The traditional answer is the ecological 'niche' — even at very small scales, environmental differences are enough to allow different species to coexist. Recently, the 'neutral theory' of biodiversity has suggested that this explanation is too complicated, and species are distributed more by chance effects. Jonathan Levine and Janneke HilleRisLambers test these ideas with an intriguing mix of experiment and theory, showing that diversity declines when niches are removed: in this round, at least, traditional explanations have the edge. If organisms are involved in a perpetual struggle for existence, how is it that communities are so diverse? The traditional answer is the ecological niche but this has recently been challenged by the neutral theory of biodiversity, which explains coexistence with the equivalence of competitors. Here, theory and experimentation are integrated in order to explore this problem; the results show that diversity declines when niches are removed. Ecological communities characteristically contain a wide diversity of species with important functional, economic and aesthetic value. Ecologists have long questioned how this diversity is maintained 1 , 2 , 3 . Classic theory shows that stable coexistence requires competitors to differ in their niches 4 , 5 , 6 ; this has motivated numerous investigations of ecological differences presumed to maintain diversity 3 , 6 , 7 , 8 . That niche differences are key to coexistence, however, has recently been challenged by the neutral theory of biodiversity, which explains coexistence with the equivalence of competitors 9 . The ensuing controversy has motivated calls for a better understanding of the collective importance of niche differences for the diversity observed in ecological communities 10 , 11 . Here we integrate theory and experimentation to show that niche differences collectively stabilize the dynamics of experimental communities of serpentine annual plants. We used field-parameterized population models to develop a null expectation for community dynamics without the stabilizing effects of niche differences. The population growth rates predicted by this null model varied by several orders of magnitude between species, which is sufficient for rapid competitive exclusion. Moreover, after two generations of community change in the field, Shannon diversity was over 50 per cent greater in communities stabilized by niche differences relative to those exhibiting dynamics predicted by the null model. Finally, in an experiment manipulating species’ relative abundances, population growth rates increased when species became rare—the demographic signature of niche differences. Our work thus provides strong evidence that species differences have a critical role in stabilizing species diversity.

Seed availability and suitable microsites for germination are likely to severely constrain the responses of plant species to climate change, especially at and beyond range edges. For example, range shifts may be slow if seed availability is low at range edges due to low parent-tree abundance or reduced fecundity. Even when seeds are available, climatic and biotic factors may further limit the availability of suitable microsites for recruitment. Unfortunately, the importance of seed and microsite limitation during range shifts remains unknown, since few studies have examined both factors simultaneously, particularly across species' ranges. To address this issue, we assessed seed availability and the factors influencing germination for six conifer species across a large environmental gradient encompassing their elevational ranges. Specifically, we assessed (1) how parent-tree abundance influences annual seed availability; (2) how seed limitation varies across species' ranges; (3) how climatic and biotic factors affect germination; and (4) how seed and suitable microsite availability covary annually within and among species. We found that seed availability declined toward species' upper range edges for most species, primarily due to low parent-tree abundance rather than declining fecundity. Range expansions are thus likely to be lagged with respect to climate change, as long generation times preclude rapid increases in tree density. Negative impacts of canopy cover on germination rates suggest range shifts will further be slowed by competition with existing vegetation. Moreover, years of high seed production were generally correlated among species, but not correlated with the availability of suitable microsites, implying that seedling competition and the interaction between seed and microsite limitation will further constrain recruitment. However, the nature of microsite limitation varied strongly between treeline and low-elevation species due to differing responses to snowpack duration and competition, suggesting that treeline species may be quicker to shift their ranges in response to warming than low-elevation species. In all, our results demonstrate that seed and microsite limitation will likely result in lagged responses to climate change but with differences among species leading to complex range shift dynamics.

Using an extensive network of occurrence records for 293 plant species collected over the past 40 years across a climatically diverse geographic section of western North America, we find that plant species distributions were just as likely to shift upwards (i.e., towards higher elevations) as downward (i.e., towards lower elevations)-despite consistent warming across the study area. Although there was no clear directional response to climate warming across the entire study area, there was significant region- to region- variation in responses (i.e. from as many as 73% to as few as 32% of species shifting upward). To understand the factors that might be controlling region-specific distributional shifts of plant species, we explored the relationship between the direction of change in distribution limits and the nature of recent climate change. We found that the direction that distribution limits shifted was explained by an interaction between the rate of change in local summer temperatures and seasonal precipitation. Specifically, species were more likely to shift upward at their upper elevational limit when minimum temperatures increased and snowfall was unchanging or declined at slower rates (<0.5 mm/year). This suggests that both low temperature and water availability limit upward shifts at upper elevation limits. By contrast, species were more likely to shift upwards at their lower elevation limit when maximum temperatures increased, but also shifted upwards under conditions of cooling temperatures when precipitation decreased. This suggests increased water stress may drive upward shifts at lower elevation limits. Our results suggest that species' elevational distribution shifts are not predictable by climate warming alone but depend on the interaction between seasonal temperature and precipitation change.

Succession and community assembly research overlap in many respects, such as through their focus on how ecological processes like dispersal, environmental filters, and biotic interactions influence community structure. Indeed, many recent advances have been made by successional studies that draw on modern analytical techniques introduced by contemporary community assembly studies. However, community assembly studies generally lack a temporal perspective, both on how the forces structuring communities might change over time and on how historical contingency (e.g. priority effects and legacy effects) and complex transitions (e.g. threshold effects) might alter community trajectories. We believe a full understanding of the complex interacting processes that shape community dynamics across large temporal scales can best be achieved by combining concepts, tools, and study systems into an integrated conceptual framework that draws upon both succession and community assembly theory.

Aim The International Tree‐Ring Data Bank (ITRDB) is the most comprehensive database of tree growth. To evaluate its usefulness and improve its accessibility to the broad scientific community, we aimed to: (a) quantify its biases, (b) assess how well it represents global forests, (c) develop tools to identify priority areas to improve its representativity, and d) make available the corrected database. Location Worldwide. Time period Contributed datasets between 1974 and 2017. Major taxa studied Trees. Methods We identified and corrected formatting issues in all individual datasets of the ITRDB. We then calculated the representativity of the ITRDB with respect to species, spatial coverage, climatic regions, elevations, need for data update, climatic limitations on growth, vascular plant diversity, and associated animal diversity. We combined these metrics into a global Priority Sampling Index (PSI) to highlight ways to improve ITRDB representativity. Results Our refined dataset provides access to a network of >52 million growth data points worldwide. We found, however, that the database is dominated by trees from forests with low diversity, in semi‐arid climates, coniferous species, and in western North America. Conifers represented 81% of the ITRDB and even in well‐sampled areas, broadleaves were poorly represented. Our PSI stressed the need to increase the database diversity in terms of broadleaf species and identified poorly represented regions that require scientific attention. Great gains will be made by increasing research and data sharing in African, Asian, and South American forests. Main conclusions The extensive data and coverage of the ITRDB show great promise to address macroecological questions. To achieve this, however, we have to overcome the significant gaps in the representativity of the ITRDB. A strategic and organized group effort is required, and we hope the tools and data provided here can guide the efforts to improve this invaluable database.

Invasive vertebrate predators are directly responsible for the extinction or decline of many vertebrate species, but their indirect impacts often go unmeasured, potentially leading to an underestimation of their full impact. When invasives extirpate functionally important mutualists, dependent species are likely to be affected as well. Here, we show that the invasive brown treesnake, directly responsible for the extirpation of forest birds from the island of Guam, is also indirectly responsible for a severe decline in plant recruitment as a result of disrupting the fruit-frugivore mutualism. To assess the impact of frugivore loss on plants, we compare seed dispersal and recruitment of two fleshy-fruited tree species on Guam and three nearby islands with intact disperser communities. We conservatively estimate that the loss of frugivorous birds caused by the brown treesnake may have caused a 61–92% decline in seedling recruitment. This case study highlights the potential for predator invasions to cause indirect, pervasive and easily overlooked interaction cascades. Invasive brown treesnakes decimated the forest bird community on the island of Guam. Now, Rogers and colleagues document the indirect effects of the snake on trees, linking snake-initiated bird loss to reduced seed dispersal and plant recruitment on Guam compared to nearby uninvaded islands.

Experimental data collected from 40 grasslands on 6 continents show that nutrients and herbivores can serve as counteracting forces to control local plant diversity; nutrient addition reduces local diversity through light limitation, and herbivory rescues diversity at sites where it alleviates light limitation. Shedding light on grazing and biodiversity Human activity has affected grassland biodiversity through the addition of both nutrients and grazing. Theory predicts that these factors could balance each other because they have opposing effects on light limitation, and this international collaboration across 40 experimental sites on six continents — from the 41 Nutrient Network (NutNet) cooperative — puts the theory to the test. The results demonstrate a consistent counteracting effect, with nutrient addition and herbivores jointly controlling plant diversity via light: nutrients reduce ground-level light thereby reducing plant diversity, and herbivores increase plant diversity by reducing competition for light among plants. This work will contribute towards more accurate modelling of the effects of grazing practices and nitrogen deposition on biodiversity in the world's grasslands. In a second paper in this issue of Nature , Yann Hautier et al . studied the influence of eutrophication in the NutNet grassland sites and show that the use of fertilizers is not only a threat to grassland biodiversity but also to the stabilizing effect it has on ecosystem functioning. Human alterations to nutrient cycles 1 , 2 and herbivore communities 3 , 4 , 5 , 6 , 7 are affecting global biodiversity dramatically 2 . Ecological theory predicts these changes should be strongly counteractive: nutrient addition drives plant species loss through intensified competition for light, whereas herbivores prevent competitive exclusion by increasing ground-level light, particularly in productive systems 8 , 9 . Here we use experimental data spanning a globally relevant range of conditions to test the hypothesis that herbaceous plant species losses caused by eutrophication may be offset by increased light availability due to herbivory. This experiment, replicated in 40 grasslands on 6 continents, demonstrates that nutrients and herbivores can serve as counteracting forces to control local plant diversity through light limitation, independent of site productivity, soil nitrogen, herbivore type and climate. Nutrient addition consistently reduced local diversity through light limitation, and herbivory rescued diversity at sites where it alleviated light limitation. Thus, species loss from anthropogenic eutrophication can be ameliorated in grasslands where herbivory increases ground-level light.

Ecosystem productivity commonly increases asymptotically with plant species diversity, and determining the mechanisms responsible for this well-known pattern is essential to predict potential changes in ecosystem productivity with ongoing species loss. Previous studies attributed the asymptotic diversity-–productivity pattern to plant competition and differential resource use (e.g., niche complementarity). Using an analytical model and a series of experiments, we demonstrate theoretically and empirically that host-specific soil microbes can be major determinants of the diversity-–productivity relationship in grasslands. In the presence of soil microbes, plant disease decreased with increasing diversity, and productivity increased nearly 500%%, primarily because of the strong effect of density-dependent disease on productivity at low diversity. Correspondingly, disease was higher in plants grown in conspecific-trained soils than heterospecific-trained soils (demonstrating host-specificity), and productivity increased and host-specific disease decreased with increasing community diversity, suggesting that disease was the primary cause of reduced productivity in species-poor treatments. In sterilized, microbe-free soils, the increase in productivity with increasing plant species number was markedly lower than the increase measured in the presence of soil microbes, suggesting that niche complementarity was a weaker determinant of the diversity-–productivity relationship. Our results demonstrate that soil microbes play an integral role as determinants of the diversity-–productivity relationship.

Shifts in wildflower phenology in response to climate change are well documented in the scientific literature. The majority of studies have revealed phenological shifts using in-situ observations, some aided by citizen science efforts (e.g., National Phenology Network). Such investigations have been instrumental in quantifying phenological shifts but are challenged by the fact that limited resources often make it difficult to gather observations over large spatial scales and long-time frames. However, recent advances in finer scale satellite imagery may provide new opportunities to detect changes in phenology. These approaches have documented plot level changes in vegetation characteristics and leafing phenology, but it remains unclear whether they can also detect flowering in natural environments. Here, we test whether fine-resolution imagery (<10 m) can detect flowering and whether combining multiple sources of imagery improves the detection process. Examining alpine wildflowers at Mt. Rainier National Park (MORA), we found that high-resolution Random Forest (RF) classification from 3-m resolution PlanetScope (from Planet Labs) imagery was able to delineate the flowering season captured by ground-based phenological surveys with an accuracy of 70% (Cohen’s kappa = 0.25). We then combined PlanetScope data with coarser resolution but higher quality imagery from Sentinel and Landsat satellites (10-m Sentinel and 30-m Landsat), resulting in higher accuracy predictions (accuracy = 77%, Cohen’s kappa = 0.39). The model was also able to identify the timing of peak flowering in a particularly warm year (2015), despite being calibrated on normal climate years. Our results suggest PlanetScope imagery holds utility in global change ecology where temporal frequency is important. Additionally, we suggest that combining imagery may provide a new approach to cross-calibrate sensors to account for radiometric irregularity inherent in fine resolution PlanetScope imagery. The development of this approach for wildflower phenology predictions provides new possibilities to monitor climate change effects on flowering communities at broader spatiotemporal scales. In protected and tourist areas where the flowering season draws large numbers of visitors, such as Mt. Rainier National Park, the modeling framework presented here could be a useful tool to manage and prioritize park resources.

How expected increases in climate variability will affect species diversity depends on the role of such variability in regulating the coexistence of competing species. Despite theory linking temporal environmental fluctuations with the maintenance of diversity, the importance of climate variability for stabilizing coexistence remains unknown because of a lack of appropriate long-term observations. Here, we analyze three decades of demographic data from a Kansas prairie to demonstrate that interannual climate variability promotes the coexistence of three common grass species. Specifically, we show that (i) the dynamics of the three species satisfy all requirements of \"storage effect\" theory based on recruitment variability with overlapping generations, (ii) climate variables are correlated with interannual variation in species performance, and (iii) temporal variability increases low-density growth rates, buffering these species against competitive exclusion. Given that environmental fluctuations are ubiquitous in natural systems, our results suggest that coexistence based on the storage effect may be underappreciated and could provide an important alternative to recent neutral theories of diversity. Field evidence for positive effects of variability on coexistence also emphasizes the need to consider changes in both climate means and variances when forecasting the effects of global change on species diversity.