Susan Harrison’s research while affiliated with University of California, Davis and other places

Anthropogenic biodiversity decline threatens the functioning of ecosystems and the many benefits they provide to humanity¹. As well as causing species losses in directly affected locations, human influence might also reduce biodiversity in relatively unmodified vegetation if far-reaching anthropogenic effects trigger local extinctions and hinder recolonization. Here we show that local plant diversity is globally negatively related to the level of anthropogenic activity in the surrounding region. Impoverishment of natural vegetation was evident only when we considered community completeness: the proportion of all suitable species in the region that are present at a site. To estimate community completeness, we compared the number of recorded species with the dark diversity—ecologically suitable species that are absent from a site but present in the surrounding region². In the sampled regions with a minimal human footprint index, an average of 35% of suitable plant species were present locally, compared with less than 20% in highly affected regions. Besides having the potential to uncover overlooked threats to biodiversity, dark diversity also provides guidance for nature conservation. Species in the dark diversity remain regionally present, and their local populations might be restored through measures that improve connectivity between natural vegetation fragments and reduce threats to population persistence.

Forbs (“wildflowers”) are important contributors to grassland biodiversity but are vulnerable to environmental changes. In a factorial experiment at 94 sites on 6 continents, we test the global generality of several broad predictions: (1) Forb cover and richness decline under nutrient enrichment, particularly nitrogen enrichment. (2) Forb cover and richness increase under herbivory by large mammals. (3) Forb richness and cover are less affected by nutrient enrichment and herbivory in more arid climates, because water limitation reduces the impacts of competition with grasses. (4) Forb families will respond differently to nutrient enrichment and mammalian herbivory due to differences in nutrient requirements. We find strong evidence for the first, partial support for the second, no support for the third, and support for the fourth prediction. Our results underscore that anthropogenic nitrogen addition is a major threat to grassland forbs, but grazing under high herbivore intensity can offset these nutrient effects.

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.

In this review and synthesis, we argue that California is an important test case for the nation and world because terrestrial biodiversity is very high, present and anticipated threats to biodiversity from climate change and other interacting stressors are severe, and innovative approaches to protecting biodiversity in the context of climate change are being developed and tested. We first review salient dimensions of California’s terrestrial physical, biological, and human diversity. Next, we examine four facets of the threat to their sustainability of these dimensions posed by climate change: direct impacts, illustrated by a new analysis of shifting diversity hotspots for plants; interactive effects involving invasive species, land-use change, and other stressors; the impacts of changing fire regimes; and the impacts of land-based renewable energy development. We examine recent policy responses in each of these areas, representing attempts to better protect biodiversity while advancing climate adaptation and mitigation. We conclude that California’s ambitious 30 × 30 Initiative and its efforts to harmonize biodiversity conservation with renewable energy development are important areas of progress. Adapting traditional suppression-oriented fire policies to the reality of new fire regimes is an area in which much progress remains to be made.

Forbs (“wildflowers”) are important contributors to grassland biodiversity and services, but they are vulnerable to environmental changes that affect their coexistence with grasses. In a factorial experiment at 94 sites on 6 continents, we tested the global generality of several broad predictions arising from previous studies: (1) Forb cover and richness decline under nutrient enrichment, particularly nitrogen enrichment, which benefits grasses at the expense of forbs. (2) Forb cover and richness increase under herbivory by large mammals, especially when nutrients are enriched. (3) Forb richness and cover are less affected by nutrient enrichment and herbivory in more arid climates, because water limitation reduces the impacts of competition with grasses. We found strong evidence for the first, partial support for the second, and no support for the third prediction. Forb richness and cover are reduced by nutrient addition, with nitrogen having the greatest effect; forb cover is enhanced by large mammal herbivory, although only under conditions of nutrient enrichment and high herbivore intensity; and forb richness is lower in more arid sites, but is not affected by consistent climate-nutrient or climate-herbivory interactions. We also found that nitrogen enrichment disproportionately affects forbs in certain families (Asteraceae, Fabaceae). Our results underscore that anthropogenic nitrogen addition is a major threat to grassland forbs and the ecosystem services they support, but grazing under high herbivore intensity can offset these nutrient effects.

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.

In 1949–1951, ecologist Robert H. Whittaker sampled plant community composition at 470 sites in the Siskiyou Mountains (Oregon and California; also known as Klamath or Klamath‐Siskiyou Mountains). His primary goal was to develop methods to quantify plant community variation across environmental gradients, following on his seminal work challenging communities as discrete entities. He selected the Siskiyous because of their diverse and endemic‐rich flora, which he attributed to geological complexity and an ancient stable climate. He chose sites to span gradients of topography, elevation, geologic substrate, and distance from the coast. He used the frequencies of indicator species in his data to assign sampling locations to positions on the topographic gradient, nested within the elevational and substrate gradients. He originated in this study the concept of diversity partitioning, in which gamma diversity (species richness of a community) equals alpha diversity (species richness in homogeneous sites) times beta diversity (species turnover among sites along gradients). Diversity partitioning subsequently became highly influential and new developments on it continue. Whittaker published his Siskiyou work covering paleohistory, biogeography, floristics, vegetation, gradient analysis, and diversity partitioning in Ecological Monographs in 1960. Discussed in 2 pages of his 60‐page monograph, diversity partitioning accounts for >95% of its current >4300 citations. In 2006, we retrieved Whittaker's Siskiyou data in hard copy from the Cornell University archives and entered them in a database. We used these data for multiple published analyses, including some based on (re)sampling the approximate locations of a subset of his sites. Because of the continued interest in diversity partitioning and in historic data sets, here we present his data, including 359 sampling locations and their descriptors and, for each sample, a list of species with their estimated percent cover (herbs and shrubs) and numbers by diameter at breast height (DBH) category (trees). Site descriptors include the approximate location (road, trail, or stream), elevation, topographic aspect, geologic substrate (serpentine, gabbro, or diorite), and dominant woody vegetation of each location. For 111 sites, including the small number chosen to represent the distance‐to‐coast gradient, we could not locate his data. There are no copyright restrictions and users of these data should cite this data paper in any publications that result from its use. The authors are available for consultations about and collaborations involving the data.

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, specially across different ecosystems. To address this, we compiled a global collection of long-term permanent vegetation records (>7000 plots from 78 datasets) from a wide range of habitats and combined this with existing trait databases. We tested whether the observed inter-annual variability in species abundance (coefficient of variation) was related to multiple individual traits and multivariate axes of trait variations (PCoA axes). We found that species with greater leaf dry matter content and seed mass were consistently more stable over time (lower variability in species abundance) although other leaf traits played a significant role as well, albeit weaker. Using multivariate axes did not improve predictions by specific traits. Our results confirm existing theory, providing compelling empirical evidence on the importance of specific traits, which point at ecological trade-offs in different resource use and dispersal strategies, on the stability of plant populations worldwide.

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.