Frequency-dependent tree growth depends on climate.

Climate and competition interact to affect species' performance, such as growth and survival, and help determine species distributions and coexistence. However, it is unclear how climatic conditions modulate frequency-dependent performance, that is, how performance changes as a species becomes locally rare or common. This is critical because declines in performance as a species becomes more common (negative frequency dependence) is a signature of niche differences among species that stabilize coexistence, whereas positive frequency dependence leads to priority effects and hampers species coexistence. Here, we used dendrochronology and hierarchical models to test whether frequency-dependent growth of sugar pine (Pinus lambertiana) depends on climatic conditions. We found that growth rates were strongly dependent on annual precipitation, but no frequency dependence was evident across all years. However, there was a strong interaction between precipitation and frequency dependence, revealing stabilizing niche differences in dry years but positive frequency dependence in wet years. These differences emerged because of precipitation-driven changes in the direction and strength of both con- and heterospecific competition. Overall, these results show how stabilizing and destabilizing effects can be temporally dynamic for long-lived species and interact with climate variation.

Belowground impacts of alpine woody encroachment are determined by plant traits, local climate, and soil conditions.

Global climate and land use change are causing woody plant encroachment in arctic, alpine, and arid/semi-arid ecosystems around the world, yet our understanding of the belowground impacts of this phenomenon is limited. We conducted a globally distributed field study of 13 alpine sites across four continents undergoing woody plant encroachment and sampled soils from both woody encroached and nearby herbaceous plant community types. We found that woody plant encroachment influenced soil microbial richness and community composition across sites based on multiple factors including woody plant traits, site level climate, and abiotic soil conditions. In particular, root symbiont type was a key determinant of belowground effects, as Nitrogen-fixing woody plants had higher soil fungal richness, while Ecto/Ericoid mycorrhizal species had higher soil bacterial richness and symbiont types had distinct soil microbial community composition. Woody plant leaf traits indirectly influenced soil microbes through their impact on soil abiotic conditions, primarily soil pH and C:N ratios. Finally, site-level climate affected the overall magnitude and direction of woody plant influence, as soil fungal and bacterial richness were either higher or lower in woody encroached versus herbaceous soils depending on mean annual temperature and precipitation. All together, these results document global impacts of woody plant encroachment on soil microbial communities, but highlight that multiple biotic and abiotic pathways must be considered to scale up globally from site- and species-level patterns. Considering both the aboveground and belowground effects of woody encroachment will be critical to predict future changes in alpine ecosystem structure and function and subsequent feedbacks to the global climate system.

Changing Climate Drives Divergent and Nonlinear Shifts in Flowering Phenology across Elevations.

Climate change is known to affect regional weather patterns and phenology; however, we lack understanding of how climate drives phenological change across local spatial gradients. This spatial variation is critical for determining whether subpopulations and metacommunities are changing in unison or diverging in phenology. Divergent responses could reduce synchrony both within species (disrupting gene flow among subpopulations) and among species (disrupting interspecific interactions in communities). We also lack understanding of phenological change in environments where life history events are frequently aseasonal, such as the tropical, arid, and semi-arid ecosystems that cover vast areas. Using a 33-year-long dataset spanning a 1,267-m semi-arid elevational gradient in the southwestern United States, we test whether flowering phenology diverged among subpopulations within species and among five communities comprising 590 species. Applying circular statistics to test for changes in year-round flowering, we show flowering has become earlier for all communities except at the highest elevations. However, flowering times shifted at different rates across elevations likely because of elevation-specific changes in temperature and precipitation, indicating diverging phenologies of neighboring communities. Subpopulations of individual species also diverged at mid-elevation but converged in phenology at high elevation. These changes in flowering phenology among communities and subpopulations are undetectable when data are pooled across the gradient. Furthermore, we show that nonlinear changes in flowering times over the 33-year record are obscured by traditional calculations of long-term trends. These findings reveal greater spatiotemporal complexity in phenological responses than previously recognized and indicate climate is driving phenological reshuffling across local spatial gradients.

Disentangling the abundance-impact relationship for invasive species.

To predict the threat of biological invasions to native species, it is critical that we understand how increasing abundance of invasive alien species (IAS) affects native populations and communities. The form of this relationship across taxa and ecosystems is unknown, but is expected to depend strongly on the trophic position of the IAS relative to the native species. Using a global metaanalysis based on 1,258 empirical studies presented in 201 scientific publications, we assessed the shape, direction, and strength of native responses to increasing invader abundance. We also tested how native responses varied with relative trophic position and for responses at the population vs. community levels. As IAS abundance increased, native populations declined nonlinearly by 20%, on average, and community metrics declined linearly by 25%. When at higher trophic levels, invaders tended to cause a strong, nonlinear decline in native populations and communities, with the greatest impacts occurring at low invader abundance. In contrast, invaders at the same trophic level tended to cause a linear decline in native populations and communities, while invaders at lower trophic levels had no consistent impacts. At the community level, increasing invader abundance had significantly larger effects on species evenness and diversity than on species richness. Our results show that native responses to invasion depend critically on invasive species' abundance and trophic position. Further, these general abundance-impact relationships reveal how IAS impacts are likely to develop during the invasion process and when to best manage them.

Shrub range expansion alters diversity and distribution of soil fungal communities across an alpine elevation gradient.

Global climate and land use change are altering plant and soil microbial communities worldwide, particularly in arctic and alpine biomes where warming is accelerated. The widespread expansion of woody shrubs into historically herbaceous alpine plant zones is likely to interact with climate to affect soil microbial community structure and function; however, our understanding of alpine soil ecology remains limited. This study aimed to (i) determine whether the diversity and community composition of soil fungi vary across elevation gradients and to (ii) assess the impact of woody shrub expansion on these patterns. In the White Mountains of California, sagebrush (Artemisia rothrockii) shrubs have been expanding upwards into alpine areas since 1960. In this study, we combined observational field data with a manipulative shrub removal experiment along an elevation transect of alpine shrub expansion. We utilized next-generation sequencing of the ITS1 region for fungi and joint distribution modelling to tease apart effects of the environment and intracommunity interactions on soil fungi. We found that soil fungal diversity declines and community composition changes with increasing elevation. Both abiotic factors (primarily soil moisture and soil organic C) and woody sagebrush range expansion had significant effects on these patterns. However, fungal diversity and relative abundance had high spatial variation, overwhelming the predictive power of vegetation type, elevation and abiotic soil conditions at the landscape scale. Finally, we observed positive and negative associations among fungal taxa which may be important in structuring community responses to global change.

When Climate Reshuffles Competitors: A Call for Experimental Macroecology.

Climate change will likely reshuffle ecological communities, causing novel species interactions that could profoundly influence how populations and communities respond to changing conditions. Nonetheless, predicting the impacts of novel interactions is challenging, partly because many methods of inference are contingent on the current configuration of climatic variables and species distributions. Focusing on competition, we argue that experiments designed to quantify novel interactions in ways that can inform species distribution models are urgently needed, and suggest an empirical agenda to pursue this goal, illustrated using plants. An emerging convergence of ideas from macroecology and demographically focused competition theory offers opportunities to mechanistically incorporate competition into species distribution models, while forging closer ties between experimental ecology and macroecology.

Novel competitors shape species' responses to climate change.

Understanding how species respond to climate change is critical for forecasting the future dynamics and distribution of pests, diseases and biological diversity. Although ecologists have long acknowledged species' direct physiological and demographic responses to climate, more recent work suggests that these direct responses can be overwhelmed by indirect effects mediated via other interacting community members. Theory suggests that some of the most dramatic impacts of community change will probably arise through the assembly of novel species combinations after asynchronous migrations with climate. Empirical tests of this prediction are rare, as existing work focuses on the effects of changing interactions between competitors that co-occur today. To explore how species' responses to climate warming depend on how their competitors migrate to track climate, we transplanted alpine plant species and intact plant communities along a climate gradient in the Swiss Alps. Here we show that when alpine plants were transplanted to warmer climates to simulate a migration failure, their performance was strongly reduced by novel competitors that could migrate upwards from lower elevation; these effects generally exceeded the impact of warming on competition with current competitors. In contrast, when we grew the focal plants under their current climate to simulate climate tracking, a shift in the competitive environment to novel high-elevation competitors had little to no effect. This asymmetry in the importance of changing competitor identity at the leading versus trailing range edges is best explained by the degree of functional similarity between current and novel competitors. We conclude that accounting for novel competitive interactions may be essential to predict species' responses to climate change accurately.

Quantifying invasion resistance: the use of recruitment functions to control for propagule pressure.

Invasive species distributions tend to be biased towards some habitats compared to others due to the combined effects of habitat-specific resistance to invasion and non-uniform propagule pressure. These two factors may also interact, with habitat resistance varying as a function of propagule supply rate. Recruitment experiments, in which the number of individuals recruiting into a population is measured under different propagule supply rates, can help us understand these interactions and quantify habitat resistance to invasion while controlling for variation in propagule supply rate. Here, we constructed recruitment functions for the invasive herb Hieracium lepidulum by sowing seeds at five different densities into six different habitat types in New Zealand's Southern Alps repeated over two successive years, and monitored seedling recruitment and survival over a four year period. We fitted recruitment functions that allowed us to estimate the total number of safe sites available for plants to occupy, which we used as a measure of invasion resistance, and tested several hypotheses concerning how invasion resistance differed among habitats and over time. We found significant differences in levels of H. lepidulum recruitment among habitats, which did not match the species' current distribution in the landscape. Local biotic and abiotic characteristics helped explain some of the between-habitat variation, with vascular plant species richness, vascular plant cover, and light availability, all positively correlated with the number of safe sites for recruitment. Resistance also varied over time however, with cohorts sown in successive years showing different levels of recruitment in some habitats but not others. These results show that recruitment functions can be used to quantify habitat resistance to invasion and to identify potential mechanisms of invasion resistance.

Integrated assessment of biological invasions.

As the main witnesses of the ecological and economic impacts of invasions on ecosystems around the world, ecologists seek to provide the relevant science that informs managers about the potential for invasion of specific organisms in their region(s) of interest. Yet, the assorted literature that could inform such forecasts is rarely integrated to do so, and further, the diverse nature of the data available complicates synthesis and quantitative prediction. Here we present a set of analytical tools for synthesizing different levels of distributional and/or demographic data to produce meaningful assessments of invasion potential that can guide management at multiple phases of ongoing invasions, from dispersal to colonization to proliferation. We illustrate the utility of data-synthesis and data-model assimilation approaches with case studies of three well-known invasive species--a vine, a marine mussel, and a freshwater crayfish--under current and projected future climatic conditions. Results from the integrated assessments reflect the complexity of the invasion process and show that the most relevant climatic variables can have contrasting effects or operate at different intensities across habitat types. As a consequence, for two of the study species climate trends will increase the likelihood of invasion in some habitats and decrease it in others. Our results identified and quantified both bottlenecks and windows of opportunity for invasion, mainly related to the role of human uses of the landscape or to disruption of the flow of resources. The approach we describe has a high potential to enhance model realism, explanatory insight, and predictive capability, generating information that can inform management decisions and optimize phase-specific prevention and control efforts for a wide range of biological invasions.

Beyond seasonal climate: statistical estimation of phenological responses to weather.

Phenological events, such as the timing of flowering or insect emergence, are influenced by a complex combination of climatic and non-climatic factors. Although temperature is generally considered most important, other weather events such as frosts and precipitation events can also influence many species' phenology. Non-climatic variables such as photoperiod and site-specific habitat characteristics can also have important effects on phenology. Forecasting phenological shifts due to climate change requires understanding and quantifying how these multiple factors combine to affect phenology. However, current approaches to analyzing phenological data have a limited ability for quantifying multiple drivers simultaneously. Here, we use a novel statistical approach to estimate the combined effects of multiple variables, including local weather events, on the phenology of several taxa (a tree, an insect, and a fungus). We found that thermal forcing had a significant positive effect on each species, frost events delayed the phenology of the tree and butterfly, and precipitation had a positive effect on fungal fruiting. Using data from sites across latitudinal gradients, we found that these effects are remarkably consistent across sites once latitude and other site effects are accounted for. This consistency suggests an underlying biological response to these variables that is not commonly estimated using data from field observations. This approach's flexibility will be useful for forecasting ongoing phenological responses to changes in climate variability in addition to seasonal trends.

Predicting species-specific responses of fungi to climatic variation using historical records.

Although striking changes have been documented in plant and animal phenology over the past century, less is known about how the fungal kingdom's phenology has been changing. A few recent studies have documented changes in fungal fruiting in Europe in the last few decades, but the geographic and taxonomic extent of these changes, the mechanisms behind these changes, and their relationships to climate are not well understood. Here, we analyzed herbarium data of 274 species of fungi from Michigan to test the hypotheses that fruiting times of fungi depend on annual climate and that responses depend on taxonomic and functional groups. We show that the fungal community overall fruits later in warmer and drier years, which has led to a shift toward later fruiting dates for autumn-fruiting species, consistent with existing evidence. However, we also show that these effects are highly variable among species and are partly explained by basic life-history characteristics. Resulting differences in climate sensitivities are expected to affect community structure as climate changes. This study provides a unique picture of the climate dependence of fungal phenology in North America and an approach for quantifying how individual species and broader fungal communities will respond to ongoing climate change.

Poised to prosper? A cross-system comparison of climate change effects on native and non-native species performance.

Climate change and biological invasions are primary threats to global biodiversity that may interact in the future. To date, the hypothesis that climate change will favour non-native species has been examined exclusively through local comparisons of single or few species. Here, we take a meta-analytical approach to broadly evaluate whether non-native species are poised to respond more positively than native species to future climatic conditions. We compiled a database of studies in aquatic and terrestrial ecosystems that reported performance measures of non-native (157 species) and co-occurring native species (204 species) under different temperature, CO(2) and precipitation conditions. Our analyses revealed that in terrestrial (primarily plant) systems, native and non-native species responded similarly to environmental changes. By contrast, in aquatic (primarily animal) systems, increases in temperature and CO(2) largely inhibited native species. There was a general trend towards stronger responses among non-native species, including enhanced positive responses to more favourable conditions and stronger negative responses to less favourable conditions. As climate change proceeds, aquatic systems may be particularly vulnerable to invasion. Across systems, there could be a higher risk of invasion at sites becoming more climatically hospitable, whereas sites shifting towards harsher conditions may become more resistant to invasions.

Forecasting phenology: from species variability to community patterns.

Shifts in species' phenology in response to climate change have wide-ranging consequences for ecological systems. However, significant variability in species' responses, together with limited data, frustrates efforts to forecast the consequences of ongoing phenological changes. Herein, we use a case study of three North American plant communities to explore the implications of variability across levels of organisation (within and among species, and among communities) for forecasting responses to climate change. We show how despite significant variation among species in sensitivities to climate, comparable patterns emerge at the community level once regional climate drivers are accounted for. However, communities differ with respect to projected patterns of divergence and overlap among their species' phenological distributions in response to climate change. These analyses and a review of hypotheses suggest how explicit consideration of spatial scale and levels of biological organisation may help to understand and forecast phenological responses to climate change.

Disentangling the paradox of insect phenology: are temporal trends reflecting the response to warming?

The strength and direction of phenological responses to changes in climate have been shown to vary significantly both among species and among populations of a species, with the overall patterns not fully resolved. Here, we studied the temporal and spatial variability associated with the response of several insect species to recent global warming. We use hierarchical models within a model comparison framework to analyze phenological data gathered over 40 years by the Japan Meteorological Agency on the emergence dates of 14 insect species at sites across Japan. Contrary to what has been predicted with global warming, temporal trends of annual emergence showed a later emergence day for some species and sites over time, even though temperatures are warming. However, when emergence data were analyzed as a function of temperature and precipitation, the overall response pointed out an earlier emergence day with warmer conditions. The apparent contradiction between the response to temperature and trends over time indicates that other factors, such as declining populations, may be affecting the date phenological events are being recorded. Overall, the responses by insects were weaker than those found for plants in previous work over the same time period in these ecosystems, suggesting the potential for ecological mismatches with deleterious effects for both suites of species. And although temperature may be the major driver of species phenology, we should be cautious when analyzing phenological datasets as many other factors may also be contributing to the variability in phenology.

Negative soil feedbacks accumulate over time for non-native plant species.

The enemy release hypothesis is a common explanation for species invasions, suggesting that introduced species benefit from leaving behind natural enemies in the native range. However, any such advantage may attenuate over time. In this study, we test a prediction of this more dynamic enemy release hypothesis: that non-native plant species that became established longer ago exhibit stronger negative feedbacks with the soil. Consistent with declining enemy release over time, we found increasingly negative soil feedbacks for species established longer ago in New Zealand. Negative soil feedbacks were also stronger for more widespread species, but weaker for more locally abundant species, suggesting that species accumulate negative interactions as they spread and can be locally regulated by these interactions. We also present data to support the common assumption that relatives have similar impacts on and responses to soil communities. Together, these data highlight the dynamic nature of novel interactions arising from species introductions.

Learning from failures: testing broad taxonomic hypotheses about plant naturalization.

Our understanding of broad taxonomic patterns of plant naturalizations is based entirely on observations of successful naturalizations. Omission of the failures, however, can introduce bias by conflating the probabilities of introduction and naturalization. Here, we use two comprehensive datasets of successful and failed plant naturalizations in New Zealand and Australia for a unique, flora-wide comparative test of several major invasion hypotheses. First, we show that some taxa are consistently more successful at naturalizing in these two countries, despite their environmental differences. Broad climatic origins helped to explain some of the differences in success rates in the two countries. We further show that species with native relatives were generally more successful in both countries, contrary to Darwin's naturalization hypothesis, but this effect was inconsistent among families across the two countries. Finally, we show that contrary to studies based on successful naturalizations only, islands need not be inherently more invisible than continents.

Safe sites, seed supply, and the recruitment function in plant populations.

The extent to which plant populations are seed vs. establishment limited can be understood by quantifying the recruitment function, describing the number of seedlings that establish as a function of the number of seeds added. Here, we derive a general equation for the recruitment function based on a mechanistic model describing how the availability of safe sites (sites suitable for germination and establishment) interacts with the number and distribution of seeds added to a plot to determine the number of recruits. The parameters of this recruitment function have a direct biological interpretation that can provide insight into the processes limiting recruitment in plant populations.

Darwin's naturalization conundrum: dissecting taxonomic patterns of species invasions.

Darwin acknowledged contrasting, plausible arguments for how species invasions are influenced by phylogenetic relatedness to the native community. These contrasting arguments persist today without clear resolution. Using data on the naturalization and abundance of exotic plants in the Auckland region, we show how different expectations can be accommodated through attention to scale, assumptions about niche overlap, and stage of invasion. Probability of naturalization was positively related to the number of native species in a genus but negatively related to native congener abundance, suggesting the importance of both niche availability and biotic resistance. Once naturalized, however, exotic abundance was not related to the number of native congeners, but positively related to native congener abundance. Changing the scale of analysis altered this outcome: within habitats exotic abundance was negatively related to native congener abundance, implying that native and exotic species respond similarly to broad scale environmental variation across habitats, with biotic resistance occurring within habitats.

Scales of association: hierarchical linear models and the measurement of ecological systems.

A fundamental challenge to understanding patterns in ecological systems lies in employing methods that can analyse, test and draw inference from measured associations between variables across scales. Hierarchical linear models (HLM) use advanced estimation algorithms to measure regression relationships and variance-covariance parameters in hierarchically structured data. Although hierarchical models have occasionally been used in the analysis of ecological data, their full potential to describe scales of association, diagnose variance explained, and to partition uncertainty has not been employed. In this paper we argue that the use of the HLM framework can enable significantly improved inference about ecological processes across levels of organization. After briefly describing the principals behind HLM, we give two examples that demonstrate a protocol for building hierarchical models and answering questions about the relationships between variables at multiple scales. The first example employs maximum likelihood methods to construct a two-level linear model predicting herbivore damage to a perennial plant at the individual- and patch-scale; the second example uses Bayesian estimation techniques to develop a three-level logistic model of plant flowering probability across individual plants, microsites and populations. HLM model development and diagnostics illustrate the importance of incorporating scale when modelling associations in ecological systems and offer a sophisticated yet accessible method for studies of populations, communities and ecosystems. We suggest that a greater coupling of hierarchical study designs and hierarchical analysis will yield significant insights on how ecological processes operate across scales.