Extreme drought impacts have been underestimated in grasslands and shrublands globally.

Climate change is increasing the frequency and severity of short-term (~1 y) drought events-the most common duration of drought-globally. Yet the impact of this intensification of drought on ecosystem functioning remains poorly resolved. This is due in part to the widely disparate approaches ecologists have employed to study drought, variation in the severity and duration of drought studied, and differences among ecosystems in vegetation, edaphic and climatic attributes that can mediate drought impacts. To overcome these problems and better identify the factors that modulate drought responses, we used a coordinated distributed experiment to quantify the impact of short-term drought on grassland and shrubland ecosystems. With a standardized approach, we imposed ~a single year of drought at 100 sites on six continents. Here we show that loss of a foundational ecosystem function-aboveground net primary production (ANPP)-was 60% greater at sites that experienced statistically extreme drought (1-in-100-y event) vs. those sites where drought was nominal (historically more common) in magnitude (35% vs. 21%, respectively). This reduction in a key carbon cycle process with a single year of extreme drought greatly exceeds previously reported losses for grasslands and shrublands. Our global experiment also revealed high variability in drought response but that relative reductions in ANPP were greater in drier ecosystems and those with fewer plant species. Overall, our results demonstrate with unprecedented rigor that the global impacts of projected increases in drought severity have been significantly underestimated and that drier and less diverse sites are likely to be most vulnerable to extreme drought.

Interactions between tall oatgrass invasion and soil nitrogen cycling.

Increases in nitrogen (N) inputs to the biosphere can exacerbate the introduction and spread of invasive non-native plant species. Often, with elevated soil N levels, invasive plants establish and further enrich soil N pools, changing overall ecosystem function. This study examined the relationship between soil N cycling and an increasingly prevalent, invasive plant species, tall oatgrass (Arrhenatherum elatius subsp. elatius), in foothills ecosystems between the Colorado Rocky Mountains and the Denver-Boulder Metropolitan area-similar to many Western US grasslands and woodlands. It focused on investigating differences in soil N transformations, inorganic N pools, and vegetation characteristics across invaded and uninvaded plots at three sites in two seasons (summer and autumn). There was a statistically significant effect of invasion on rates of net N mineralization, but it was dependent on site and season (p = 0.046). Site had a statistically significant effect on soil moisture and aboveground biomass C:N (p < 0.04). The interactions of invasion x site were statistically significant for ammonium pools (p < 0.03). These findings suggest that A. elatius invasion can be associated with accelerated N cycling, but that the nature of the relationship differs by location and season in the foothills. More broadly, this study contributes to determining how the N cycle is shifting in grassland ecosystems subject to increasing pressures from anthropogenic change.

Ecological similarity is related to phylogenetic distance between species in a cross-niche field transplant experiment.

To determine whether phylogenetic relatedness predicts ecological niche differences and community assembly in the field, we transplanted 16 focal plant species into field niches of species of increasing phylogenetic relatedness, manipulated the presence of plant neighbors, and measured environmental covariates. We found that plant survivorship declined with increasing phylogenetic distance in the presence of neighbors, but with neighbor removal, reached a low point in field niches occupied by species diverged at 63 My, the maximum age of confamilials in our study, and then increased again in the sites of distant relatives. Plant biomass was similarly nonlinear, and niche differences increased with neighbor removal. Competitive response showed a linear decline with relatedness. We compared our experimental results to natural community composition, finding that conspecifics and distant relatives were more likely to co-occur at smaller spatial scales, as predicted by our measures of performance.

Phylogenetic conservatism in plant-soil feedback and its implications for plant abundance.

We examined whether plant-soil feedback and plant-field abundance were phylogenetically conserved. For 57 co-occurring native and exotic plant species from an old field in Canada, we collected a data set on the effects of three soil biota treatments on plant growth: net whole-soil feedback (combined effects of mutualists and antagonists), feedback with arbuscular mycorrhizal fungi (AMF) collected from soils of conspecific plants, and feedback with Glomus etunicatum, a dominant mycorrhizal fungus. We found phylogenetic signal in both net whole-soil feedback and feedback with AMF of conspecifics; conservatism was especially strong among native plants but absent among exotics. The abundance of plants in the field was also conserved, a pattern underlain by shared plant responses to soil biota. We conclude that soil biota influence the abundance of close plant relatives in nature.

The geography and ecology of plant speciation: range overlap and niche divergence in sister species.

A goal of evolutionary biology is to understand the roles of geography and ecology in speciation. The recent shared ancestry of sister species can leave a major imprint on their geographical and ecological attributes, possibly revealing processes involved in speciation. We examined how ecological similarity, range overlap and range asymmetry are related to time since divergence of 71 sister species pairs in the California Floristic Province (CFP). We found that plants exhibit strikingly different age-range correlation patterns from those found for animals; the latter broadly support allopatric speciation as the primary mode of speciation. By contrast, plant sisters in the CFP were sympatric in 80% of cases and range sizes of sisters differed by a mean of 10-fold. Range overlap and range asymmetry were greatest in younger sisters. These results suggest that speciation mechanisms broadly grouped under 'budding' speciation, in which a larger ranged progenitor gives rise to a smaller ranged derivative species, are probably common. The ecological and reproductive similarity of sisters was significantly greater than that of sister-non-sister congeners for every trait assessed. However, shifts in at least one trait were present in 93% of the sister pairs; habitat and soil shifts were especially common. Ecological divergence did not increase with range overlap contrary to expectations under character displacement in sympatry. Our results suggest that vicariant speciation is more ubiquitous in animals than plants, perhaps owing to the sensitivity of plants to fine-scale environmental heterogeneity. Despite high levels of range overlap, ecological shifts in the process of budding speciation may result in low rates of fine-scale spatial co-occurrence. These results have implications for ecological studies of trait evolution and community assembly; despite high levels of sympatry, sister taxa and potentially other close relatives, may be missing from local communities.

The nature of serpentine endemism.

Serpentine soils are a model system for the study of plant adaptation, speciation, and species interactions. Serpentine soil is an edaphically stressful, low productivity soil type that hosts stunted vegetation and a spectacular level of plant endemism. One of the first papers on serpentine plant endemism was by Arthur Kruckeberg, titled "Intraspecific variability in the response of certain native plant species to serpentine soil." Published in the American Journal of Botany in 1951, it has been cited over 100 times. Here, I review the context and content of the paper, as well as its impact. On the basis of the results of reciprocal transplant experiments in the greenhouse, Kruckeberg made three important conclusions on the nature of serpentine plant endemism: (1) Plants are locally adapted to serpentine soils, forming distinct soil ecotypes; (2) soil ecotypes are the first stage in the evolutionary progression toward serpentine endemism; and (3) serpentine endemics are restricted from more fertile nonserpentine soils by competition. Kruckeberg's paper inspired a substantial amount of research, especially in the three areas reviewed here: local adaptation and plant traits, speciation, and the interaction of climate and soil in plant endemism. In documenting soil ecotypes, Kruckeberg identified serpentine soils as a potent selective factor in plant evolution and helped establish serpentine soils as a model system in evolution and ecology.

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.

Plant community data from a statewide survey of paired serpentine and non-serpentine soils in California, USA.

Soils derived from ultramafic parent materials (hereafter serpentine) provide habitat for unique plant communities containing species with adaptations to the low nutrient levels, high magnesium : calcium ratios, and high metal content (Ni, Zn) that characterize serpentine. Plants on serpentine have long been studied in evolution and ecology, and plants adapted to serpentine contribute disproportionately to plant diversity in many parts of the world. In 2000-2003, serpentine plant communities were sampled at 107 locations representing the full range of occurrence of serpentine in California, USA, spanning large gradients in climate. In 2009-2010, plant communities were similarly sampled at 97 locations on nonserpentine soil, near to and paired with 97 of the serpentine sampling locations. (Some serpentine locations were revisited in 2009-2010 to assess the degree of change since 2000-2003, which was minimal.) At each serpentine or nonserpentine location, a north- and a south-facing 50 × 10 m plot were sampled. This design produced 97 "sites" each consisting of four "plots" (north-south exposure, serpentine-nonserpentine soil). All plots were initially visited three or more times over two years to record plant diversity and cover, and a subset were revisited in 2014 to examine community change after a drought. The original question guiding the study was how plant diversity is shaped by the spatially patchy nature of the serpentine habitat. Subsequently, we investigated how climate drives plant diversity at multiple scales (within locations, between locations on the same and different soil types, and across entire regions) and at different levels of organization (taxonomic, functional, and phylogenetic). There are no copyright restrictions and users should cite this data paper in publications that result from use of the data.

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.