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Darwin’s Naturalization Conundrum holds that both functional similarity and distinctiveness can facilitate biological invasions: invaders similar to natives may succeed through preadaptation to local abiotic conditions, whereas functionally distinct invaders may succeed by reducing competition. Yet the contexts in which either mechanism dominates are unclear. Prior research has primarily attributed variability in native–invasive differentiation to shifts in the balance between biotic and climatic barriers to invasion from local to regional scales. However, similarity and distinctiveness are frequent at both local and regional levels, indicating key drivers of native–invasive differentiation remain overlooked. Crucially, theory and evidence show that as climatic stress increases, competition weakens. This implies that harsh climates should favor invaders functionally similar to natives, whereas mesic climates should favor distinctiveness. However, this hypothesis has not been tested across climate regimes and functional traits. We addressed this gap by combining models of species distributions and flowering phenology for 2,810 species across the United States, estimating phylogenetic distance and phenological differentiation between natives and invasives relative to differentiation among co-occurring natives. In warm, humid regions, invasives flowered earlier, less synchronously, and were more distantly related to natives. In cold or dry regions, they flowered at similar or later times, more synchronously, and were more closely related than natives themselves. Across all climates, invasives consistently exhibited longer flowering durations, with little evidence of greater phenological plasticity. These findings reveal that Darwin’s Conundrum reflects a predictable continuum shaped by environmental context, highlighting climate as a key axis of invasion success. 

Abstract Global warming has caused widespread shifts in plant phenology among species in the temperate zone, but it is unclear how population‐level responses will scale to alter the structure of the flowering season at the community level. This knowledge gap exists largely because—while the climatic sensitivity of first flowering within populations has been studied extensively—little is known about the responsiveness of the duration of a population's flowering period. This limits our ability to anticipate how the entire flowering periods of co‐occurring species may continue to change under warming. Nonetheless, flowering sensitivity to temperature often varies predictably among species between and within communities, which may help forecast temperature‐related changes to a community's flowering season. However, no studies—empirical or theoretical—have assessed how patterns of variation in flowering sensitivity among species could scale to alter community‐level flowering changes under warming. Here, we provide a conceptual overview of how variation in the sensitivity of flowering onset and duration among species can mediate changes to a community's flowering season due to warming trends. Specifically, we focus on the effects of differences in (1) the mean sensitivity of flowering onset and duration among communities and (2) the sensitivity of flowering onsets and durations among species flowering sequentially through the season within a community. We evaluated the manner and degree in which these forms of between‐species variation in sensitivity might affect the structure of the flowering season—both independently and interactively—using simulations, which covered a wide but empirically informed range of parameter values and combinations representing distinct community‐level patterns. Our findings predict that communities across the temperate zone will exhibit varied and often contrasting flowering responses to warming across biomes, underscoring that accounting for the temperature sensitivity of both phenological onset and duration among species is essential for understanding community‐level flowering dynamics in a warming world. 

Summary Phenological response to global climate change can impact ecosystem functions. There are various data sources from which spatiotemporal and taxonomic phenological data may be obtained: mobilized herbaria, community science initiatives, observatory networks, and remote sensing. However, analyses conducted to date have generally relied on single sources of these data. Siloed treatment of data in analyses may be due to the lack of harmonization across different data sources that offer partially nonoverlapping information and are often complementary. Such treatment precludes a deeper understanding of phenological responses at varying macroecological scales. Here, we describe a detailed vision for the harmonization of phenological data, including the direct integration of disparate sources of phenological data using a common schema. Specifically, we highlight existing methods for data harmonization that can be applied to phenological data: data design patterns, metadata standards, and ontologies. We describe how harmonized data from multiple sources can be integrated into analyses using existing methods and discuss the use of automated extraction techniques. Data harmonization is not a new concept in ecology, but the harmonization of phenological data is overdue. We aim to highlight the need for better data harmonization, providing a roadmap for how harmonized phenological data may fill gaps while simultaneously being integrated into analyses. 

Abstract Many plant species alter both the timing and duration of their flowering in response to changing climate and often differ with respect to the magnitudes or directions of their phenological responses to climate changes. These shifts may have cumulative effects on the diversity of species simultaneously in flower throughout a given flowering season, resulting in periods of disproportionately high or low species richness of actively flowering community members relative to historical conditions. Although the potential for such changes to occur is well appreciated, few studies have assessed how climate trends have affected patterns of co‐flowering synchrony due to a scarcity of long‐term datasets documenting flowering duration across species in a community. In this study, we leveraged 1,908,706 plant specimens collected in flower to model the effects of warming throughout the past century on the daily species richness of actively flowering species by developing species‐specific phenoclimate models for 1848 plant species inhabiting 16 well‐documented plant communities across California. These communities are located across a variety of ecoregions, ranging from coastal marshes and grasslands to deserts, chaparral shrublands, and coniferous forests. The recurring patterns in the modeled community‐level flowering displays indicate that recent warming has consistently shortened the period during which many species flower concurrently, and that the bloom season has advanced by nearly 5 days on average. Accordingly, within every flora, recent warming was predicted to increase the daily species richness of flowers early in the local growing season, with corresponding reductions in species richness of flowers later in the growing season. Notably, patterns of change in community‐level bloom displays were driven primarily by differences among species in the timing of flowering onset, as termination dates tended to advance in unison with onset dates, resulting in minor changes to flowering duration among species. 

Forecasting the impacts of changing climate on the phenology of plant populations is essential for anticipating and managing potential ecological disruptions to biotic communities. Herbarium specimens enable assessments of plant phenology across broad spatiotemporal scales. However, specimens are collected opportunistically, and it is unclear whether their collection dates – used as proxies of phenological stages – are closest to the onset, peak, or termination of a phenophase, or whether sampled individuals represent early, average, or late occurrences in their populations. Despite this, no studies have assessed whether these uncertainties limit the utility of herbarium specimens for estimating the onset and termination of a phenophase. Using simulated data mimicking such uncertainties, we evaluated the accuracy with which the onset and termination of population‐level phenological displays (in this case, of flowering) can be predicted from natural‐history collections data (controlling for biases in collector behavior), and how the duration, variability, and responsiveness to climate of the flowering period of a species and temporal collection biases influence model accuracy. Estimates of population‐level onset and termination were highly accurate for a wide range of simulated species' attributes, but accuracy declined among species with longer individual‐level flowering duration and when there were temporal biases in sample collection, as is common among the earliest and latest‐flowering species. The amount of data required to model population‐level phenological displays is not impractical to obtain; model accuracy declined by less than 1 day as sample sizes rose from 300 to 1000 specimens. Our analyses of simulated data indicate that, absent pervasive biases in collection and if the climate conditions that affect phenological timing are correctly identified, specimen data can predict the onset, termination, and duration of a population's flowering period with similar accuracy to estimates of median flowering time that are commonplace in the literature. 

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. 

ABSTRACT Global change is altering the phenology and geographic ranges of flowering species, with potentially profound consequences for the timing and composition of floral resources and the seasonal structure of ecological communities. However, shifts in flowering phenology and species distributions have historically been studied in isolation due to disciplinary silos and limited data, leaving critical gaps in our understanding of their combined effects. To address this, we used millions of herbarium and occurrence records to model phenological and range shifts for 2837 plant species in the United States across historical, recent, and projected climate and land cover conditions, enabling us to scale responses from species to communities, and from local to continental geographies. Our analysis reveals that communities are shifting toward earlier, longer flowering seasons in most biomes, with co‐flowering species richness increasing at the edges of the season and declining at historical peaks—trends projected to intensify under ongoing environmental trends. Although range and phenology shifts operate concurrently, they predominantly affect different aspects of the flowering season: phenological changes primarily alter seasonality—its start, end, and duration—and co‐flowering diversity at the edges of the season, while range shifts more strongly influence co‐flowering species richness during historical seasonal peaks and the identity and degree of flowering synchrony among co‐occurring species pairs. Together, these results demonstrate that shifts in phenology and species ranges act synergistically to restructure the flowering seasons across the conterminous United States, revealing wide variation in the pace and direction of change among biomes.