Functional trait trade-offs define plant population stability across different biomes.

Ecological theory posits that temporal stability patterns in plant populations are associated with differences in species' ecological strategies. However, empirical evidence is lacking about which traits, or trade-offs, underlie species stability, especially across different biomes. We compiled a worldwide collection of long-term permanent vegetation records (greater than 7000 plots from 78 datasets) from a large range of habitats which we combined with existing trait databases. We tested whether the observed inter-annual variability in species abundance (coefficient of variation) was related to multiple individual traits. We found that populations with greater leaf dry matter content and seed mass were more stable over time. Despite the variability explained by these traits being low, their effect was consistent across different datasets. Other traits played a significant, albeit weaker, role in species stability, and the inclusion of multi-variate axes or phylogeny did not substantially modify nor improve predictions. These results provide empirical evidence and highlight the relevance of specific ecological trade-offs, i.e. in different resource-use and dispersal strategies, for plant populations stability across multiple biomes. Further research is, however, necessary to integrate and evaluate the role of other specific traits, often not available in databases, and intraspecific trait variability in modulating species stability.

Vulnerability and resistance in the spatial heterogeneity of soil microbial communities under resource additions.

Spatial heterogeneity in composition and function enables ecosystems to supply diverse services. For soil microbes and the ecosystem functions they catalyze, whether such heterogeneity can be maintained in the face of altered resource inputs is uncertain. In a 50-ha northern California grassland with a mosaic of plant communities generated by different soil types, we tested how spatial variability in microbial composition and function changed in response to nutrient and water addition. Fungal composition lost some of its spatial variability in response to nutrient addition, driven by decreases in mutualistic fungi and increases in antagonistic fungi that were strongest on the least fertile soils, where mutualists were initially most frequent and antagonists initially least frequent. Bacterial and archaeal community composition showed little change in their spatial variability with resource addition. Microbial functions related to nitrogen cycling showed increased spatial variability under nutrient, and sometimes water, additions, driven in part by accelerated nitrification on the initially more-fertile soils. Under anthropogenic changes such as eutrophication and altered rainfall, these findings illustrate the potential for significant changes in ecosystem-level spatial heterogeneity of microbial functions and communities.

Synchrony matters more than species richness in plant community stability at a global scale.

The stability of ecological communities is critical for the stable provisioning of ecosystem services, such as food and forage production, carbon sequestration, and soil fertility. Greater biodiversity is expected to enhance stability across years by decreasing synchrony among species, but the drivers of stability in nature remain poorly resolved. Our analysis of time series from 79 datasets across the world showed that stability was associated more strongly with the degree of synchrony among dominant species than with species richness. The relatively weak influence of species richness is consistent with theory predicting that the effect of richness on stability weakens when synchrony is higher than expected under random fluctuations, which was the case in most communities. Land management, nutrient addition, and climate change treatments had relatively weak and varying effects on stability, modifying how species richness, synchrony, and stability interact. Our results demonstrate the prevalence of biotic drivers on ecosystem stability, with the potential for environmental drivers to alter the intricate relationship among richness, synchrony, and stability.

Extreme pre-fire drought decreases shrub regeneration on fertile soils.

Extreme drought and increasing temperatures can decrease the resilience of plant communities to fires. Not only may extremely dry conditions during or after fires lead to higher plant mortality and poorer recruitment, but severe pre-fire droughts may reduce the seed production and belowground vigor that are essential to post-fire plant recovery, and may indirectly facilitate invasion. We studied survival, recruitment, and growth of shrubs and herbs in chaparral (shrubland) communities in Northern California after a 2015 fire that immediately followed California's extreme 3-yr drought. We followed the same protocols used to study similar, adjacent communities after a 1999 fire that did not follow a drought, and we compared the two recovery trajectories. Overall, the 2015 fire was not more severe than the 1999 fire, yet it caused higher mortality and lower growth of resprouting shrubs on fertile (sandstone) soils. In contrast, the 2015 fire did not affect the mortality or growth of resprouting shrubs on infertile (serpentine) soils, the density of shrub seedlings, or the richness or cover of native herbs differently than the 1999 fire. However, the 2015 fire facilitated a massive increase in exotic herbaceous cover, especially on fertile soils, possibly portending the early stages of a type conversion to exotic-dominated grassland. Our findings indicate that the consequences of climate change on fire-dependent communities will include effects of pre-fire as well as post-fire climate, and that resprouting shrubs are particularly likely to be sensitive to pre-fire drought.

Trait-based signatures of cloud base height in a tropical cloud forest.

PREMISE: Clouds have profound consequences for ecosystem structure and function. Yet, the direct monitoring of clouds and their effects on biota is challenging especially in remote and topographically complex tropical cloud forests. We argue that known relationships between climate and the taxonomic and functional composition of plant communities may provide a fingerprint of cloud base height, thus providing a rapid and cost-effective assessment in remote tropical cloud forests. METHODS: To detect cloud base height, we compared species turnover and functional trait values among herbaceous and woody plant communities in an ecosystem dominated by cloud formation. We measured soil and air temperature, soil nutrient concentrations, and extracellular enzyme activity. We hypothesized that woody and herbaceous plants would provide signatures of cloud base height, as evidenced by abrupt shifts in both taxonomic composition and plant function. RESULTS: We demonstrated abrupt changes in taxonomic composition and the community- weighted mean of a key functional trait, specific leaf area, across elevation for both woody and herbaceous species, consistent with our predictions. However, abrupt taxonomic and functional changes occurred 100 m higher in elevation for herbaceous plants compared to woody ones. Soil temperature abruptly decreased where herbaceous taxonomic and functional turnover was high. Other environmental variables including soil biogeochemistry did not explain the abrupt change observed for woody plant communities. CONCLUSIONS: We provide evidence that a trait-based approach can be used to estimate cloud base height. We outline how rises in cloud base height and differential environmental requirements between growth forms can be distinguished using this approach.

Seed banks of native forbs, but not exotic grasses, increase during extreme drought.

Extreme droughts such as the one that affected California in 2012-2015 have been linked to severe ecological consequences in perennial-dominated communities such as forests. In annual communities, drought impacts are difficult to assess because many species persist through facultative multiyear seed dormancy, which leads to the development of seed banks. Impacts of extreme drought on the abundance and composition of the seed banks of whole communities are little known. In 80 heterogeneous grassland plots where cover is dominated by ~15 species of exotic annual grasses and diversity is dominated by ~70 species of native annual forbs, we grew out seeds from soil cores collected early in the California drought (2012) and later in the multiyear drought (2014), and analyzed drought-associated changes in the seed bank. Over the course of the study we identified more than 22,000 seedlings to species. We found that seeds of exotic annual grasses declined sharply in abundance during the drought while seeds of native annual forbs increased, a pattern that resembled but was even stronger than the changes in aboveground cover of these groups. Consistent with the expectation that low specific leaf area (SLA) is an indicator of drought tolerance, we found that the community-weighted mean SLA of annual forbs declined both in the seed bank and in the aboveground community, as low-SLA forbs increased disproportionately. In this system, seed dormancy reinforces the indirect benefits of extreme drought to the native forb community.

Climate-driven diversity change in annual grasslands: Drought plus deluge does not equal normal.

Climate forecasts agree that increased variability and extremes will tend to reduce the availability of water in many terrestrial ecosystems. Increasingly severe droughts may be exacerbated both by warmer temperatures and by the relative unavailability of water that arrives in more sporadic and intense rainfall events. Using long-term data and an experimental water manipulation, we examined the resilience of a heterogeneous annual grassland community to a prolonged series of dry winters that led to a decline in plant species richness (2000-2014), followed by a near-record wet winter (2016-2017), a climatic sequence that broadly resembles the predicted future in its high variability. In our 80, 5-m2 observational plots, species richness did not recover in response to the wet winter, and the positive relationship of richness to annual winter rainfall thus showed a significant weakening trend over the 18-year time period. In experiments on 100, 1-m2 plots, wintertime water supplementation increased and drought shelters decreased the seedling survival and final individual biomass of native annual forbs, the main functional group contributing to the observed long-term decline in richness. Water supplementation also increased the total cover of native annual forbs, but only increased richness within nested subplots to which seeds were also added. We conclude that prolonged dry winters, by increasing seedling mortality and reducing growth of native forbs, may have diminished the seedbank and thus the recovery potential of diversity in this community. However, the wet winter and the watering treatment did cause recovery of the community mean values of a key functional trait (specific leaf area, an indicator of drought intolerance), suggesting that some aggregate community properties may be stabilized by functional redundancy among species.

Resource colimitation governs plant community responses to altered precipitation.

Ecological theory and evidence suggest that plant community biomass and composition may often be jointly controlled by climatic water availability and soil nutrient supply. To the extent that such colimitation operates, alterations in water availability caused by climatic change may have relatively little effect on plant communities on nutrient-poor soils. We tested this prediction with a 5-y rainfall and nutrient manipulation in a semiarid annual grassland system with highly heterogeneous soil nutrient supplies. On nutrient-poor soils, rainfall addition alone had little impact, but rainfall and nutrient addition synergized to cause large increases in biomass, declines in diversity, and near-complete species turnover. Plant species with resource-conservative functional traits (low specific leaf area, short stature) were replaced by species with resource-acquisitive functional traits (high specific leaf area, tall stature). On nutrient-rich soils, in contrast, rainfall addition alone caused substantial increases in biomass, whereas fertilization had little effect. Our results highlight that multiple resource limitation is a critical aspect when predicting the relative vulnerability of natural communities to climatically induced compositional change and diversity loss.

Climate-driven diversity loss in a grassland community.

Local ecological communities represent the scale at which species coexist and share resources, and at which diversity has been experimentally shown to underlie stability, productivity, invasion resistance, and other desirable community properties. Globally, community diversity shows a mixture of increases and decreases over recent decades, and these changes have relatively seldom been linked to climatic trends. In a heterogeneous California grassland, we documented declining plant diversity from 2000 to 2014 at both the local community (5 m(2)) and landscape (27 km(2)) scales, across multiple functional groups and soil environments. Communities became particularly poorer in native annual forbs, which are present as small seedlings in midwinter; within native annual forbs, community composition changed toward lower representation of species with a trait indicating drought intolerance (high specific leaf area). Time series models linked diversity decline to the significant decrease in midwinter precipitation. Livestock grazing history, fire, succession, N deposition, and increases in exotic species could be ruled out as contributing causes. This finding is among the first demonstrations to our knowledge of climate-driven directional loss of species diversity in ecological communities in a natural (nonexperimental) setting. Such diversity losses, which may also foreshadow larger-scale extinctions, may be especially likely in semiarid regions that are undergoing climatic trends toward higher aridity and lower productivity.

Shading and litter mediate the effects of soil fertility on the performance of an understorey herb.

BACKGROUND AND AIMS: Soil fertility and topographic microclimate are common determinants of plant species distributions. However, biotic conditions also vary along these abiotic gradients, and may mediate their effects on plants. In this study, we investigated whether soils and topographic microclimate acted directly on the performance of a focal understorey plant, or indirectly via changing biotic conditions. METHODS: We examined direct and indirect relationships between abiotic variables (soil fertility and topographic microclimate) and biotic factors (overstorey and understorey cover, litter depth and mycorrhizal colonization) and the occurrence, density and flowering of a common understorey herb, Trientalis latifolia, in the Klamath-Siskiyou Mountains, Oregon, USA. RESULTS: We found that the positive effects of soil fertility on Trientalis occurrence were mediated by greater overstorey shading and deeper litter. However, we did not find any effects of topographic microclimate on Trientalis distribution that were mediated by the biotic variables we measured. The predictive success of Trientalis species distribution models with soils and topographic microclimate increased by 12 % with the addition of the biotic variables. CONCLUSIONS: Our results reinforce the idea that species distributions are the outcome of interrelated abiotic gradients and biotic interactions, and suggest that biotic conditions, such as overstorey density, should be included in species distribution models if data are available.

Landscape structure affects specialists but not generalists in naturally fragmented grasslands.

Understanding how biotic communities respond to landscape spatial structure is critically important for conservation management as natural habitats become increasingly fragmented. However, empirical studies of the effects of spatial structure on plant species richness have found inconsistent results, suggesting that more comprehensive approaches are needed. We asked how landscape structure affects total plant species richness and the richness of a guild of specialized plants in a multivariate context. We sampled herbaceous plant communities at 56 dolomite glades (insular, fire-adapted grasslands) across the Missouri Ozarks, USA, and used structural equation modeling (SEM) to analyze the relative importance of landscape structure, soil resource availability, and fire history for plant communities. We found that landscape spatial structure, defined as the area-weighted proximity of glade habitat surrounding study sites (proximity index), had a significant effect on total plant species richness, but only after we controlled for environmental covariates. Richness of specialist species, but not generalists, was positively related to landscape spatial structure. Our results highlight that local environmental filters must be considered to understand the influence of landscape structure on communities and that unique species guilds may respond differently to landscape structure than the community as a whole. These findings suggest that both local environment and landscape context should be considered when developing management strategies for species of conservation concern in fragmented habitats.

Rapid shifts in grassland communities driven by climate change.

Many terrestrial plant communities, especially forests, have been shown to lag in response to rapid climate change. Grassland communities may respond more quickly to novel climates, as they consist mostly of short-lived species, which are directly exposed to macroclimate change. Here we report the rapid response of grassland communities to climate change in the California Floristic Province. We estimated 349 vascular plant species' climatic niches from 829,337 occurrence records, compiled 15 long-term community composition datasets from 12 observational studies and 3 global change experiments, and analysed community compositional shifts in the climate niche space. We show that communities experienced significant shifts towards species associated with warmer and drier locations at rates of 0.0216 ± 0.00592 °C yr-1 (mean ± s.e.) and -3.04 ± 0.742 mm yr-1, and these changes occurred at a pace similar to that of climate warming and drying. These directional shifts were consistent across observations and experiments. Our findings contrast with the lagged responses observed in communities dominated by long-lived plants and suggest greater biodiversity changes than expected in the near future.

Historical and ecological controls on phylogenetic diversity in Californian plant communities.

We addressed the classic question of whether community diversity is determined from the bottom up by the breadth and partitioning of niche space or from the top down by historical and evolutionary forces. Specifically, we contrasted local and regional explanations for the diversity of Californian plant communities using phylogenetic and functional analyses. Our communities were sets of four field plots that sampled alpha (within-plot) and beta (among-plot) sources of variation in diversity. We sampled 93 such communities nested within 78 larger regions for which regional species pools could be independently estimated, spanning the California Floristic Province. We measured phylogenetic and functional diversity within plots and between plots on neighboring soils and slopes. We also measured the phylogenetic diversity of regional species pools and analyzed them in terms of biogeographic groups. We found no evidence linking the phylogenetic diversity of communities to within-plot functional diversity or among-plot beta diversity. Instead, we found that the phylogenetic diversity of communities depends on that of regional species pools. In turn, phylogenetically diverse pools were those with high proportions of species of northern biogeographic affinity, which have relatively mesic distributions and traits. This supports what we call the climatic refuge hypothesis rather than the biogeographic crossroads hypothesis.

Navigating the multiple meanings of ß diversity: a roadmap for the practicing ecologist.

A recent increase in studies of ß diversity has yielded a confusing array of concepts, measures and methods. Here, we provide a roadmap of the most widely used and ecologically relevant approaches for analysis through a series of mission statements. We distinguish two types of ß diversity: directional turnover along a gradient vs. non-directional variation. Different measures emphasize different properties of ecological data. Such properties include the degree of emphasis on presence/absence vs. relative abundance information and the inclusion vs. exclusion of joint absences. Judicious use of multiple measures in concert can uncover the underlying nature of patterns in ß diversity for a given dataset. A case study of Indonesian coral assemblages shows the utility of a multi-faceted approach. We advocate careful consideration of relevant questions, matched by appropriate analyses. The rigorous application of null models will also help to reveal potential processes driving observed patterns in ß diversity.

Is plant community richness regulated over time? Contrasting results from experiments and long-term observations.

There is considerable debate among ecologists as to whether or not communities are saturated. In saturated communities, species richness should remain relatively constant over time, despite compositional turnover, because richness is negatively correlated with colonization and positively correlated with local extinction. Few studies have tested for saturation using temporal observational data as well as diversity-perturbation experiments. We analyzed 10 years of data for plant species richness at 71 sites on contrasting serpentine and non-serpentine soils within Californian (USA) grasslands. We also manipulated local richness and measured its effects on immigration and extinction. Consistent with saturation, we observed that richness was positively correlated with extinction rates and negatively correlated with colonization rates, and randomization tests confirmed that diversity fluctuated less than expected by chance. However, experimental species additions and removals did not affect extinction or colonization, suggesting that richness is not regulated by local species interactions. Instead, we propose three reasons why richness may fluctuate within narrow limits causing the appearance of saturation in temporal observational data sets: negatively autocorrelated patterns of biotic response to yearly conditions, differential affinities of particular species for local conditions, or stochastic abundance-dependent colonization and extinction rates. We illustrate the latter using a metacommunity model.

Can spatial isolation help predict dispersal-limited sites for native species restoration?

When the distribution of species is limited by propagule supply, new populations may be initiated by seed addition, but identifying suitable sites for efficiently targeted seed addition remains a major challenge for restoration. In addition to the biotic or abiotic variables typically used in species distribution models, spatial isolation from conspecifics could help predict the suitability of unoccupied sites. Site suitability might be expected to increase with spatial isolation after other factors are accounted for, since isolation increases the chance that a site is unoccupied only because of propagule limitation. For two native annual forbs in Californian grasslands, we combined experimental seeding and niche modeling to ask whether suitability of unoccupied sites could be predicted by spatial variables (either distances from, or densities of, conspecific populations), either by themselves or in combination with niche models. We also asked whether experimental tests of these predictions held up not only in the short term (one year), but also in the longer term (three years). For Lasthenia californica, seed additions were only successful relatively near existing populations. For Lupinus nanus, seeding success was low and was positively related to the number of conspecifics within 1 km. For both species, a few previously unoccupied sites remained occupied three years after seeding, but this subset was not predictable based on either spatial or niche variables. Seed addition alone may be a limited means of native forb restoration if suitable unoccupied sites are either rare or unpredictable, or if they tend to be close to where the species already occurs.

Niche conservatism as an emerging principle in ecology and conservation biology.

The diversity of life is ultimately generated by evolution, and much attention has focused on the rapid evolution of ecological traits. Yet, the tendency for many ecological traits to instead remain similar over time [niche conservatism (NC)] has many consequences for the fundamental patterns and processes studied in ecology and conservation biology. Here, we describe the mounting evidence for the importance of NC to major topics in ecology (e.g. species richness, ecosystem function) and conservation (e.g. climate change, invasive species). We also review other areas where it may be important but has generally been overlooked, in both ecology (e.g. food webs, disease ecology, mutualistic interactions) and conservation (e.g. habitat modification). We summarize methods for testing for NC, and suggest that a commonly used and advocated method (involving a test for phylogenetic signal) is potentially problematic, and describe alternative approaches. We suggest that considering NC: (1) focuses attention on the within-species processes that cause traits to be conserved over time, (2) emphasizes connections between questions and research areas that are not obviously related (e.g. invasives, global warming, tropical richness), and (3) suggests new areas for research (e.g. why are some clades largely nocturnal? why do related species share diseases?).

Phylogeny, niche conservatism and the latitudinal diversity gradient in mammals.

Biologists have long searched for mechanisms responsible for the increase in species richness with decreasing latitude. The strong correlation between species richness and climate is frequently interpreted as reflecting a causal link via processes linked to energy or evolutionary rates. Here, we investigate how the aggregation of clades, as dictated by phylogeny, can give rise to significant climate-richness gradients without gradients in diversification or environmental carrying capacity. The relationship between climate and species richness varies considerably between clades, regions and time periods in a global-scale phylogenetically informed analysis of all terrestrial mammal species. Many young clades show negative richness-temperature slopes (more species at cooler temperatures), with the ages of these clades coinciding with the expansion of temperate climate zones in the late Eocene. In carnivores, we find steeply positive richness-temperature slopes in clades with restricted distributions and tropical origins (e.g. cat clade), whereas widespread, temperate clades exhibit shallow, negative slopes (e.g. dog-bear clade). We show that the slope of the global climate-richness gradient in mammals is driven by aggregating Chiroptera (bats) with their Eutherian sister group. Our findings indicate that the evolutionary history should be accounted for as part of any search for causal links between environment and species richness.

Invasive species interact with climatic variability to reduce success of natives.

Plants have evolved resource-conservative and resource-acquisitive strategies to deal with variability in rainfall, but interactions with dominant invasive species may undermine these adaptations. To investigate the relative effect of invaders on species with these two strategies, we manipulated rainfall and invasive grass presence and measured demographic rates in three resource-acquisitive and three resource-conservative native annual forbs. We found that invasive grasses were harmful to all of the target species, but especially the resource-acquisitive ones, and that these effects were stronger under experimental drought. Invasive grass presence under drought lowered per capita population growth rates of acquisitive natives through increased mortality and decreased seed set. While invasive grasses also decreased per capita growth rates of resource-conservative natives, they did so by increasing mortality under experimental watering and by limiting the production of seed under experimental drought. Invasive species can thus interact with climatic fluctuations to make bad years worse for resource-acquisitive natives and good years less good for resource-conservative natives, and they may generally tend to undermine the acquisitive strategy more than the conservative one.

Temporal variability and nestedness in California grassland species composition.

Nestedness occurs when species-poor assemblages contain a subset of the species that occur in more species-rich communities and is a commonly observed pattern in spatial data sets. Examination of nested distribution patterns across time rather than space are rarely conducted, even though they may have important implications for species coexistence. Nested temporal assemblages can occur when most species respond similarly to interannual variation in conditions. In contrast, assemblages might be non-nested when different sets of species occur in different years, either because of different resource requirements or as a result of competitive exclusion. Using eight years of plant occurrence data at 71 sites in California grasslands, we found strong signals of temporal nestedness with most species favored by similar conditions. High-quality years enabled the expansion of both grasses and forbs into locales where they were not found during poor-quality years. Native annual forb, exotic annual forb, and exotic annual grass species richness were all greatest in cool, wet years following hot, dry years. Together, these analyses support the hypothesis that, in the absence of community members that specialize on poor-quality years, interannual environmental variation can cause communities to form nested subsets across time much as they do across space.