Plasticity and not adaptation is the primary source of temperature-mediated variation in flowering phenology in North America.

Phenology varies widely over space and time because of its sensitivity to climate. However, whether phenological variation is primarily generated by rapid organismal responses (plasticity) or local adaptation remains unresolved. Here we used 1,038,027 herbarium specimens representing 1,605 species from the continental United States to measure flowering-time sensitivity to temperature over time (Stime) and space (Sspace). By comparing these estimates, we inferred how adaptation and plasticity historically influenced phenology along temperature gradients and how their contributions vary among species with different phenology and native climates and among ecoregions differing in species composition. Parameters Sspace and Stime were positively correlated (r = 0.87), of similar magnitude and more frequently consistent with plasticity than adaptation. Apparent plasticity and adaptation generated earlier flowering in spring, limited responsiveness in late summer and delayed flowering in autumn in response to temperature increases. Nonetheless, ecoregions differed in the relative contributions of adaptation and plasticity, from consistently greater importance of plasticity (for example, southeastern United States plains) to their nearly equal importance throughout the season (for example, Western Sierra Madre Piedmont). Our results support the hypothesis that plasticity is the primary driver of flowering-time variation along temperature gradients, with local adaptation having a widespread but comparatively limited role.

Relationships of climate, human activity, and fire history to spatiotemporal variation in annual fire probability across California.

In the face of recent wildfires across the Western United States, it is essential that we understand both the dynamics that drive the spatial distribution of wildfire, and the major obstacles to modeling the probability of wildfire over space and time. However, it is well documented that the precise relationships of local vegetation, climate, and ignitions, and how they influence fire dynamics, may vary over space and among local climate, vegetation, and land use regimes. This raises questions not only as to the nature of the potentially nonlinear relationships between local conditions and the fire, but also the possibility that the scale at which such models are developed may be critical to their predictive power and to the apparent relationship of local conditions to wildfire. In this study we demonstrate that both local climate-through limitations posed by fuel dryness (CWD) and availability (AET)-and human activity-through housing density, roads, electrical infrastructure, and agriculture, play important roles in determining the annual probabilities of fire throughout California. We also document the importance of previous burn events as potential barriers to fire in some environments, until enough time has passed for vegetation to regenerate sufficiently to sustain subsequent wildfires. We also demonstrate that long-term and short-term climate variations exhibit different effects on annual fire probability, with short-term climate variations primarily impacting fire probability during periods of extreme climate anomaly. Further, we show that, when using nonlinear modeling techniques, broad-scale fire probability models can outperform localized models at predicting annual fire probability. Finally, this study represents a powerful tool for mapping local fire probability across the state of California under a variety of historical climate regimes, which is essential to avoided emissions modeling, carbon accounting, and hazard severity mapping for the application of fire-resistant building codes across the state of California.

Advancing frost dates have reduced frost risk among most North American angiosperms since 1980.

In recent decades, the final frost dates of winter have advanced throughout North America, and many angiosperm taxa have simultaneously advanced their flowering times as the climate has warmed. Phenological advancement may reduce plant fitness, as flowering prior to the final frost date of the winter/spring transition may damage flower buds or open flowers, limiting fruit and seed production. The risk of floral exposure to frost in the recent past and in the future, however, also depends on whether the last day of winter frost is advancing more rapidly, or less rapidly, than the date of onset of flowering in response to climate warming. This study presents the first continental-scale assessment of recent changes in frost risk to floral tissues, using digital records of 475,694 herbarium specimens representing 1,653 angiosperm species collected across North America from 1920 to 2015. For most species, among sites from which they have been collected, dates of last frost have advanced much more rapidly than flowering dates. As a result, frost risk has declined in 66% of sampled species. Moreover, exotic species consistently exhibit lower frost risk than native species, primarily because the former occupy warmer habitats where the annual frost-free period begins earlier. While reducing the probability of exposure to frost has clear benefits for the survival of flower buds and flowers, such phenological advancement may disrupt other ecological processes across North America, including pollination, herbivory, and disease transmission.

A new phenological metric for use in pheno-climatic models: A case study using herbarium specimens of Streptanthus tortuosus.

PREMISE: Herbarium specimens have been used to detect climate-induced shifts in flowering time by using the day of year of collection (DOY) as a proxy for first or peak flowering date. Variation among herbarium sheets in their phenological status, however, undermines the assumption that DOY accurately represents any particular phenophase. Ignoring this variation can reduce the explanatory power of pheno-climatic models (PCMs) designed to predict the effects of climate on flowering date. METHODS: Here we present a protocol for the phenological scoring of imaged herbarium specimens using an ImageJ plugin, and we introduce a quantitative metric of a specimen's phenological status, the phenological index (PI), which we use in PCMs to control for phenological variation among specimens of Streptanthus tortuosus (Brassicaceeae) when testing for the effects of climate on DOY. We demonstrate that including PI as an independent variable improves model fit. RESULTS: Including PI in PCMs increased the model R 2 relative to PCMs that excluded PI; regression coefficients for climatic parameters, however, remained constant. DISCUSSION: Our protocol provides a simple, quantitative phenological metric for any observed plant. Including PI in PCMs increases R 2 and enables predictions of the DOY of any phenophase under any specified climatic conditions.

Overlooked climate parameters best predict flowering onset: Assessing phenological models using the elastic net.

Determining the manner in which plant species shift their flowering times in response to climatic conditions is essential to understanding and forecasting the impacts of climate change on the world's flora. The limited taxonomic diversity and duration of most phenological datasets, however, have impeded a comprehensive, systematic determination of the best predictors of flowering phenology. Additionally, many studies of the relationship between climate conditions and plant phenology have included only a limited set of climate parameters that are often chosen a priori and may therefore overlook those parameters to which plants are most phenologically sensitive. This study harnesses 894,392 digital herbarium records and 1,959 in situ observations to produce the first assessment of the effects of a large number (25) of climate parameters on the flowering time of a very large number (2,468) of angiosperm taxa throughout North America. In addition, we compare the predictive capacity of phenological models constructed from the collection dates of herbarium specimens vs. repeated in situ observations of individual plants using a regression approach-elastic net regularization-that has not previously been used in phenological modeling, but exhibits several advantages over ordinary least squares and stepwise regression. When herbarium-derived data and in situ phenological observations were used to predict flowering onset, the multivariate models based on each of these data sources had similar predictive capacity (R2  = 0.27). Further, apart from mean maximum temperature (TMAX), the two best predictors of flowering time have not commonly been included in phenological models: the number of frost-free days (NFFD) and the quantity of precipitation as snow (PAS) in the seasons preceding flowering. By vetting these models across an unprecedented number of taxa, this work demonstrates a new approach to phenological modeling.

Long-term herbarium records reveal temperature-dependent changes in flowering phenology in the southeastern USA.

In recent years, a growing body of evidence has emerged indicating that the relationship between flowering phenology and climate may differ throughout various portions of the growing season. These differences have resulted in long-term changes in flowering synchrony that may alter the quantity and diversity of pollinator attention to many species, as well as altering food availability to pollenivorous and nectarivorous animal species. However, long-term multi-season records of past flowering timing have primarily focused on temperate environments. In contrast, changes in flowering phenology within humid subtropical environments such as the southeastern USA remain poorly documented. This research uses herbarium-based methods to examine changes in flowering time across 19,328 samples of spring-, summer-, and autumn-flowering plants in the southeastern USA from the years 1951 to 2009. In this study, species that flower near the onset of the growing season were found to advance under increasing mean March temperatures (-3.391 days/°C, p = 0.022). No long-term advances in early spring flowering or spring temperature were detected during this period, corroborating previous phenological assessments for the southeastern USA. However, late spring through mid-summer flowering exhibited delays in response to higher February temperatures (over 0.1.85 days/°C, p ≤ 0.041 in all cases). Thus, it appears that flowering synchrony may undergo significant restructuring in response to warming spring temperatures, even in humid subtropical environments.

Digital herbarium archives as a spatially extensive, taxonomically discriminate phenological record; a comparison to MODIS satellite imagery.

This study demonstrates that phenological information included in digital herbarium archives can produce annual phenological estimates correlated to satellite-derived green wave phenology at a regional scale (R = 0.183, P = 0.03). Thus, such records may be utilized in a fashion similar to other annual phenological records and, due to their longer duration and ability to discriminate among the various components of the plant community, hold significant potential for use in future research to supplement the deficiencies of other data sources as well as address a wide array of important issues in ecology and bioclimatology that cannot be addressed easily using more traditional methods.