Soil properties constrain forest understory plant distributions along an elevation gradient.

Projections of spatial biodiversity dynamics under climate warming are often based on models including only climate variables, and when non-climatic factors (e.g. soil) are included, data are often at much coarser spatial resolutions than those experienced by plants. Field studies along elevation gradients permit the gathering of detailed soil data, while still covering a wide climatic gradient. Here, an intensive field survey of four spring forest herbs along an elevation gradient showed that soil properties had substantial impacts on the occurrence/abundance of all species, and that soil effects were more pronounced at higher elevations. For Trillium erectum and Claytonia caroliniana, very infrequent occurrences at high elevation were strongly associated with rare microsites with high pH or nutrients. In a seven-year transplant experiment with T. erectum, we found that individuals grew to much smaller sizes at high than low elevation, suggesting that environmental factors rather than dispersal limitation constrain the species' upper range limit, despite substantial warming in recent decades. Our study demonstrates that soil factors interact strongly with climate to determine plant range limits along climatic gradients. Unsuitable soils for plants at high elevations or latitudes may represent an important constraint on future plant migration and biodiversity change. This article is part of the theme issue 'Ecological novelty and planetary stewardship: biodiversity dynamics in a transforming biosphere'.

Soil properties constrain predicted poleward migration of plants under climate change.

Many plant species are predicted to migrate poleward in response to climate change. Species distribution models (SDMs) have been widely used to quantify future suitable habitats, but they often neglect soil properties, despite the importance of soil for plant fitness. As soil properties often change along latitudinal gradients, higher-latitude soils might be more or less suitable than average conditions within the current ranges of species, thereby accelerating or slowing potential poleward migration. In this study, we built three SDMs - one with only climate predictors, one with only soil predictors, and one with both - for each of 1870 plant species in Eastern North America, in order to investigate the relative importance of soil properties in determining plant distributions and poleward shifts under climate change. While climate variables were the most important predictors, soil properties also had a substantial influence on continental-scale plant distributions. Under future climate scenarios, models including soil predicted much smaller northward shifts in distributions than climate-only models (c. 40% reduction). Our findings strongly suggest that high-latitude soils are likely to impede ongoing plant migration, and they highlight the necessity of incorporating soil properties into models and predictions for plant distributions and migration under environmental change.

Mapping canopy traits over Québec using airborne and spaceborne imaging spectroscopy.

The advent of new spaceborne imaging spectrometers offers new opportunities for ecologists to map vegetation traits at global scales. However, to date most imaging spectroscopy studies exploiting satellite spectrometers have been constrained to the landscape scale. In this paper we present a new method to map vegetation traits at the landscape scale and upscale trait maps to the continental level, using historical spaceborne imaging spectroscopy (Hyperion) to derive estimates of leaf mass per area, nitrogen, and carbon concentrations of forests in Québec, Canada. We compare estimates for each species with reference field values and obtain good agreement both at the landscape and continental scales, with patterns consistent with the leaf economic spectrum. By exploiting the Hyperion satellite archive to map these traits and successfully upscale the estimates to the continental scale, we demonstrate the great potential of recent and upcoming spaceborne spectrometers to benefit plant biodiversity monitoring and conservation efforts.

Looking back on biodiversity change: lessons for the road ahead.

Estimating biodiversity change across the planet in the context of widespread human modification is a critical challenge. Here, we review how biodiversity has changed in recent decades across scales and taxonomic groups, focusing on four diversity metrics: species richness, temporal turnover, spatial beta-diversity and abundance. At local scales, change across all metrics includes many examples of both increases and declines and tends to be centred around zero, but with higher prevalence of declining trends in beta-diversity (increasing similarity in composition across space or biotic homogenization) and abundance. The exception to this pattern is temporal turnover, with changes in species composition through time observed in most local assemblages. Less is known about change at regional scales, although several studies suggest that increases in richness are more prevalent than declines. Change at the global scale is the hardest to estimate accurately, but most studies suggest extinction rates are probably outpacing speciation rates, although both are elevated. Recognizing this variability is essential to accurately portray how biodiversity change is unfolding, and highlights how much remains unknown about the magnitude and direction of multiple biodiversity metrics at different scales. Reducing these blind spots is essential to allow appropriate management actions to be deployed. This article is part of the theme issue 'Detecting and attributing the causes of biodiversity change: needs, gaps and solutions'.

Diversity and assembly of lichens and bryophytes on tree trunks along a temperate to boreal elevation gradient.

Based on hypotheses related to environmental filtering vs. stochastic community assembly, we tested taxon-specific predictions regarding the relationships of alpha diversity, beta diversity and species composition of epiphytic macrolichens and bryophytes with elevation and the lateral gradient on trees (the different sides of the tree bole related to aspect and trunk inclination) at Parc national du Mont-Mégantic in Southeastern Québec, Canada. For lichens on firs, increasing elevation was associated with increasing alpha diversity, and a marked shift in community composition, at the scale of whole trees. In contrast, for bryophytes on maples, tree inclination and the lateral gradient had the strongest effects: more inclined trees had greater whole-tree alpha diversity and stronger within-tree contrasts in composition between the upper and lower bole surfaces. For lichens on maples, whole-tree alpha diversity showed a weak, negative relationship with inclination, and beta diversity increased slightly with elevation. Our results are consistent with theories predicting greater alpha diversity in more favorable environments (for lichens: high elevation with high relative air humidity and lower temperatures; for bryophytes: upper surfaces of tree boles with liquid water available), but support was weak for the prediction of greater beta diversity in more favorable environments. Overall, the important predictors of epiphytic cryptogam diversity vary more among the species of tree host (maple vs. fir) than focal taxa (lichens vs. bryophytes), with patterns likely related to different effects of water, temperature, and competition between lichens and bryophytes.

Evolutionary history explains foliar spectral differences between arbuscular and ectomycorrhizal plant species.

Leaf spectra are integrated foliar phenotypes that capture a range of traits and can provide insight into ecological processes. Leaf traits, and therefore leaf spectra, may reflect belowground processes such as mycorrhizal associations. However, evidence for the relationship between leaf traits and mycorrhizal association is mixed, and few studies account for shared evolutionary history. We conduct partial least squares discriminant analysis to assess the ability of spectra to predict mycorrhizal type. We model the evolution of leaf spectra for 92 vascular plant species and use phylogenetic comparative methods to assess differences in spectral properties between arbuscular mycorrhizal and ectomycorrhizal plant species. Partial least squares discriminant analysis classified spectra by mycorrhizal type with 90% (arbuscular) and 85% (ectomycorrhizal) accuracy. Univariate models of principal components identified multiple spectral optima corresponding with mycorrhizal type due to the close relationship between mycorrhizal type and phylogeny. Importantly, we found that spectra of arbuscular mycorrhizal and ectomycorrhizal species do not statistically differ from each other after accounting for phylogeny. While mycorrhizal type can be predicted from spectra, enabling the use of spectra to identify belowground traits using remote sensing, this is due to evolutionary history and not because of fundamental differences in leaf spectra due to mycorrhizal type.

Biodiversity change under adaptive community dynamics.

Compositional change is a ubiquitous response of ecological communities to environmental drivers of global change, but is often regarded as evidence of declining "biotic integrity" relative to historical baselines. Adaptive compositional change, however, is a foundational idea in evolutionary biology, whereby changes in gene frequencies within species boost population-level fitness, allowing populations to persist as the environment changes. Here, we present an analogous idea for ecological communities based on core concepts of fitness and selection. Changes in community composition (i.e., frequencies of genetic differences among species) in response to environmental change should normally increase the average fitnessof community members. We refer to compositional changes that improve the functional match, or "fit," between organisms' traits and their environment as adaptive community dynamics. Environmental change (e.g., land-use change) commonly reduces the fit between antecedent communities and new environments. Subsequent change in community composition in response to environmental changes, however, should normally increase community-level fit, as the success of at least some constituent species increases. We argue that adaptive community dynamics are likely to improve or maintain ecosystem function (e.g., by maintaining productivity). Adaptive community responses may simultaneously produce some changes that are considered societally desirable (e.g., increased carbon storage) and others that are undesirable (e.g., declines of certain species), just as evolutionary responses within species may be deemed desirable (e.g., evolutionary rescue of an endangered species) or undesirable (e.g., enhanced virulence of an agricultural pest). When assessing possible management interventions, it is important to distinguish between drivers of environmental change (e.g., undesired climate warming) and adaptive community responses, which may generate some desirable outcomes. Efforts to facilitate, accept, or resist ecological change require separate consideration of drivers and responses, and may highlight the need to reconsider preferences for historical baseline communities over communities that are better adapted to the new conditions.

Predicting leaf traits across functional groups using reflectance spectroscopy.

Plant ecologists use functional traits to describe how plants respond to and influence their environment. Reflectance spectroscopy can provide rapid, non-destructive estimates of leaf traits, but it remains unclear whether general trait-spectra models can yield accurate estimates across functional groups and ecosystems. We measured leaf spectra and 22 structural and chemical traits for nearly 2000 samples from 103 species. These samples span a large share of known trait variation and represent several functional groups and ecosystems, mainly in eastern Canada. We used partial least-squares regression (PLSR) to build empirical models for estimating traits from spectra. Within the dataset, our PLSR models predicted traits such as leaf mass per area (LMA) and leaf dry matter content (LDMC) with high accuracy (R2 > 0.85; %RMSE < 10). Models for most chemical traits, including pigments, carbon fractions, and major nutrients, showed intermediate accuracy (R2  = 0.55-0.85; %RMSE = 12.7-19.1). Micronutrients such as Cu and Fe showed the poorest accuracy. In validation on external datasets, models for traits such as LMA and LDMC performed relatively well, while carbon fractions showed steep declines in accuracy. We provide models that produce fast, reliable estimates of several functional traits from leaf spectra. Our results reinforce the potential uses of spectroscopy in monitoring plant function around the world.

Different temporal trends in vascular plant and bryophyte communities along elevational gradients over four decades.

Despite many studies showing biodiversity responses to warming, the generality of such responses across taxonomic groups remains unclear. Very few studies have tested for evidence of bryophyte community responses to warming, even though bryophytes are major contributors to diversity and functioning in many ecosystems. Here, we report an empirical study comparing long-term change in bryophyte and vascular plant communities in two sites with contrasting long-term warming trends, using "legacy" botanical records as a baseline for comparison with contemporary resurveys. We hypothesized that ecological changes would be greater in sites with a stronger warming trend and that vascular plant communities, with narrower climatic niches, would be more sensitive than bryophyte communities to climate warming. For each taxonomic group in each site, we quantified the magnitude of changes in species' distributions along the elevation gradient, species richness, and community composition. We found contrasted temporal changes in bryophyte vs. vascular plant communities, which only partially supported the warming hypothesis. In the area with a stronger warming trend, we found a significant increase in local diversity and dissimilarity (ß-diversity) for vascular plants, but not for bryophytes. Presence-absence data did not provide sufficient power to detect elevational shifts in species distributions. The patterns observed for bryophytes are in accordance with recent literature showing that local diversity can remain unchanged despite strong changes in composition. Regardless of whether one taxon is systematically more or less sensitive to environmental change than another, our results suggest that vascular plants cannot be used as a surrogate for bryophytes in terms of predicting the nature and magnitude of responses to warming. Thus, to assess overall biodiversity responses to global change, abundance data from different taxonomic groups and different community properties need to be synthesized.

A millennium of increasing diversity of ecosystems until the mid-20th century.

Land-use change is widely regarded as a simplifying and homogenising force in nature. In contrast, analysing global land-use reconstructions from the 10th to 20th centuries, we found progressive increases in the number, evenness, and diversity of ecosystems (including human-modified land-use types) present across most of the Earth's land surface. Ecosystem diversity increased more rapidly after ~1700 CE, then slowed or slightly declined (depending on the metric) following the mid-20th century acceleration of human impacts. The results also reveal increasing spatial differentiation, rather than homogenisation, in both the presence-absence and area-coverage of different ecosystem types at sub-global scales-at least, prior to the mid-20th century. Nonetheless, geographic homogenization was revealed for a subset of analyses at a global scale, reflecting the now-global presence of certain human-modified ecosystem types. Our results suggest that, while human land-use changes have caused declines in relatively undisturbed or "primary" ecosystem types, they have also driven increases in ecosystem diversity over the last millennium.

Large herbivores trigger spatiotemporal changes in forest plant diversity.

Large herbivores can exert top-down control on terrestrial plant communities, but the magnitude, direction, and scale dependency of their impacts remain equivocal, especially in temperate and boreal forests, where multiple disturbances often interact. Using a unique, long-term, and replicated landscape experiment, we assessed the influence of a high density of white-tailed deer (Odocoileus virginianus) on the spatiotemporal dynamics of diversity, composition, and successional trajectories of understorey plant assemblages in recently logged boreal forests. This experiment provided a rare opportunity to test whether deer herbivory represents a direct filter on plant communities or if it mainly acts to suppress dominant plants, which, in turn, release other plant species from strong negative plant-plant interactions. These two hypotheses make different predictions about changes in community composition and alpha and beta diversity in different vegetation layers and at different spatial scales. Our results showed that deer had strong effects on plant community composition and successional trajectories, but the resulting impacts on plant alpha and beta diversity patterns were markedly scale dependent in both time and space. Responses of tree and non-tree vegetation layers were strongly asymmetric. Deer acted both as a direct filter and as a suppressor of dominant plant species during early forest succession, but the magnitude of both processes was specific to tree and non-tree vegetation layers. Although our data supported the ungulate-driven homogenization hypothesis, compositional shifts and changes of alpha diversity were poor predictors of beta diversity loss. Our findings underscore the importance of long-term studies in revealing nonlinear temporal community trends, and they challenge managers to prioritize particular community properties and scales of interest, given contrasting trends of composition and alpha and beta diversity across spatial scales.

Mycorrhizal dominance reduces local tree species diversity across US forests.

Ectomycorrhizas and arbuscular mycorrhizas, the two most widespread plant-fungal symbioses, are thought to differentially influence tree species diversity, with positive plant-soil feedbacks favouring locally abundant ectomycorrhizal tree species and negative feedbacks promoting species coexistence and diversity in arbuscular mycorrhizal forests. While seedling recruitment studies and cross-biome patterns of plant diversity and mycorrhizal dominance support this hypothesis, it remains to be tested at the forest stand level over continental scales. Here, we analyse approximately 82,000 forest plots across the USA to show that both ectomycorrhizal-dominated and arbuscular mycorrhizal-dominated forests show relatively low tree diversity, while forests with a mixture of mycorrhizal strategies support a higher number of tree species. Our findings suggest that mycorrhizal dominance, rather than mycorrhizal type, shapes tree diversity in forests.

Directional turnover towards larger-ranged plants over time and across habitats.

Species turnover is ubiquitous. However, it remains unknown whether certain types of species are consistently gained or lost across different habitats. Here, we analysed the trajectories of 1827 plant species over time intervals of up to 78 years at 141 sites across mountain summits, forests, and lowland grasslands in Europe. We found, albeit with relatively small effect sizes, displacements of smaller- by larger-ranged species across habitats. Communities shifted in parallel towards more nutrient-demanding species, with species from nutrient-rich habitats having larger ranges. Because these species are typically strong competitors, declines of smaller-ranged species could reflect not only abiotic drivers of global change, but also biotic pressure from increased competition. The ubiquitous component of turnover based on species range size we found here may partially reconcile findings of no net loss in local diversity with global species loss, and link community-scale turnover to macroecological processes such as biotic homogenisation.

Earlier spring reduces potential for gene flow via reduced flowering synchrony across an elevational gradient.

PREMISE: One of the best-documented ecological responses to climate warming involves temporal shifts of phenological events. However, we lack an understanding of how phenological responses to climate change vary among populations of the same species. Such variability has the potential to affect flowering synchrony among populations and hence the potential for gene flow. METHODS: To test whether an earlier start of the growing season affects the potential for gene flow among populations, we quantified the distributions of flowering times of two spring-flowering plants (Trillium erectum and Erythronium americanum) over 6 years along an elevational gradient. We developed a novel model-based metric of potential gene flow between pairs of populations to quantify the potential for pollen-mediated gene flow based on flowering phenology. RESULTS: Earlier onset of spring led to greater separation of peak flowering dates across the elevational gradient for both species investigated, but was only associated with a reduction in potential gene flow in T. erectum, not E. americanum. CONCLUSIONS: Our study suggests that climate change could decrease gene flow via phenological separation among populations along climatic gradients. We also provide a novel method for quantifying potential pollen-mediated gene flow using data on flowering phenology, based on a quantitative, more biologically interpretable model than other available metrics.

Space-for-time inferences about range-edge dynamics of tree species can be influenced by sampling biases.

Differences between the distributions of tree saplings and adults in geographic or niche space have been used to infer climate change effects on tree range dynamics. Previous studies have reported narrower latitudinal or climatic niche ranges of juvenile trees compared to adults, concluding that tree ranges are contracting, contradicting climate-based predictions. However, more comprehensive sampling of adult trees than juvenile trees in most regional forest inventories could potentially bias ontogenetic comparisons. Here we first report spatial simulations showing that reduced sampling intensity can result in underestimates of range and niche limits, but that resampling the same number of individuals of different life stages can eliminate this bias. We then reanalyzed the U.S. Forest Inventory and Analysis data, comparing the range and niche limits between adult trees and saplings of 92 tree species, both using the original data and two resampling procedures. Resampling aimed to reduce sampling biases by controlling for either sampling area or the number of individuals sampled. Overall, these resampling procedures had a major influence on the estimation of range limits, most often by reducing, eliminating, or even reversing the tendency in the original analyses for saplings to have broader distributions than adult trees. These results indicate that previous conclusions that the distributions of juvenile trees were contracting in response to climate change were potentially artifacts of sampling in the underlying data. More generally, sampling effects involved in the estimation of geographic ranges and environmental niche widths need to be taken into account in studies comparing different life stages, and also likely in other types of distribution comparisons.

The geography of biodiversity change in marine and terrestrial assemblages.

Human activities are fundamentally altering biodiversity. Projections of declines at the global scale are contrasted by highly variable trends at local scales, suggesting that biodiversity change may be spatially structured. Here, we examined spatial variation in species richness and composition change using more than 50,000 biodiversity time series from 239 studies and found clear geographic variation in biodiversity change. Rapid compositional change is prevalent, with marine biomes exceeding and terrestrial biomes trailing the overall trend. Assemblage richness is not changing on average, although locations exhibiting increasing and decreasing trends of up to about 20% per year were found in some marine studies. At local scales, widespread compositional reorganization is most often decoupled from richness change, and biodiversity change is strongest and most variable in the oceans.

Understanding Maladaptation by Uniting Ecological and Evolutionary Perspectives.

Evolutionary biologists have long trained their sights on adaptation, focusing on the power of natural selection to produce relative fitness advantages while often ignoring changes in absolute fitness. Ecologists generally have taken a different tack, focusing on changes in abundance and ranges that reflect absolute fitness while often ignoring relative fitness. Uniting these perspectives, we articulate various causes of relative and absolute maladaptation and review numerous examples of their occurrence. This review indicates that maladaptation is reasonably common from both perspectives, yet often in contrasting ways. That is, maladaptation can appear strong from a relative fitness perspective, yet populations can be growing in abundance. Conversely, resident individuals can appear locally adapted (relative to nonresident individuals) yet be declining in abundance. Understanding and interpreting these disconnects between relative and absolute maladaptation, as well as the cases of agreement, is increasingly critical in the face of accelerating human-mediated environmental change. We therefore present a framework for studying maladaptation, focusing in particular on the relationship between absolute and relative fitness, thereby drawing together evolutionary and ecological perspectives. The unification of these ecological and evolutionary perspectives has the potential to bring together previously disjunct research areas while addressing key conceptual issues and specific practical problems.

Global buffering of temperatures under forest canopies.

Macroclimate warming is often assumed to occur within forests despite the potential for tree cover to modify microclimates. Here, using paired measurements, we compared the temperatures under the canopy versus in the open at 98 sites across 5 continents. We show that forests function as a thermal insulator, cooling the understory when ambient temperatures are hot and warming the understory when ambient temperatures are cold. The understory versus open temperature offset is magnified as temperatures become more extreme and is of greater magnitude than the warming of land temperatures over the past century. Tree canopies may thus reduce the severity of warming impacts on forest biodiversity and functioning.

Stronger influence of anthropogenic disturbance than climate change on century-scale compositional changes in northern forests.

Predicting future ecosystem dynamics depends critically on an improved understanding of how disturbances and climate change have driven long-term ecological changes in the past. Here we assembled a dataset of >100,000 tree species lists from the 19th century across a broad region (>130,000km2) in temperate eastern Canada, as well as recent forest inventories, to test the effects of changes in anthropogenic disturbance, temperature and moisture on forest dynamics. We evaluate changes in forest composition using four indices quantifying the affinities of co-occurring tree species with temperature, drought, light and disturbance. Land-use driven shifts favouring more disturbance-adapted tree species are far stronger than any effects ascribable to climate change, although the responses of species to disturbance are correlated with their expected responses to climate change. As such, anthropogenic and natural disturbances are expected to have large direct effects on forests and also indirect effects via altered responses to future climate change.

Four decades of plant community change along a continental gradient of warming.

Many studies of individual sites have revealed biotic changes consistent with climate warming (e.g., upward elevational distribution shifts), but our understanding of the tremendous variation among studies in the magnitude of such biotic changes is minimal. In this study, we resurveyed forest vegetation plots 40 years after the initial surveys in three protected areas along a west-to-east gradient of increasingly steep recent warming trends in eastern Canada (Québec). Consistent with the hypothesis that climate warming has been an important driver of vegetation change, we found an increasing magnitude of changes in species richness and composition from west to east among the three parks. For the two mountainous parks, we found no significant changes in elevational species' distributions in the easternmost park (raw mean = +11.4 m at Forillon Park) where warming has been minimal, and significant upward distribution shifts in the centrally located park (+38.9 m at Mont-Mégantic), where the recent warming trend has been marked. Community Temperature Indices (CTI), reflecting the average affinities of locally co-occurring species to temperature conditions across their geographic ranges ("Species Temperature Indices"), did not change over time as predicted. However, close examination of the underpinnings of CTI values suggested a high sensitivity to uncertainty in individual species' temperature indices, and so a potentially limited responsiveness to warming. Overall, by testing a priori predictions concerning variation among parks in the direction and magnitude of vegetation changes, we have provided stronger evidence for a link between climate warming and biotic responses than otherwise possible and provided a potential explanation for large variation among studies in warming-related biotic changes.