Butterfly phenology in Mediterranean mountains using space‐for‐time substitution

Inferring species' responses to climate change in the absence of long‐term time series data is a challenge, but can be achieved by substituting space for time. For example, thermal elevational gradients represent suitable proxies to study phenological responses to warming. We used butterfly data from two Mediterranean mountain areas to test whether mean dates of appearance of communities and individual species show a delay with increasing altitude, and an accompanying shortening in the duration of flight periods. We found a 14‐day delay in the mean date of appearance per kilometer increase in altitude for butterfly communities overall, and an average 23‐day shift for 26 selected species, alongside average summer temperature lapse rates of 3°C per km. At higher elevations, there was a shortening of the flight period for the community of 3 days/km, with an 8.8‐day average decline per km for individual species. Rates of phenological delay differed significantly between the two mountain ranges, although this did not seem to result from the respective temperature lapse rates. These results suggest that climate warming could lead to advanced and lengthened flight periods for Mediterranean mountain butterfly communities. However, although multivoltine species showed the expected response of delayed and shortened flight periods at higher elevations, univoltine species showed more pronounced delays in terms of species appearance. Hence, while projections of overall community responses to climate change may benefit from space‐for‐time substitutions, understanding species‐specific responses to local features of habitat and climate may be needed to accurately predict the effects of climate change on phenology.     

Forecasting phenology under global warming

As a consequence of warming temperatures around the world, spring and autumn phenologies have been shifting, with corresponding changes in the length of the growing season. Our understanding of the spatial and interspecific variation of these changes, however, is limited. Not all species are responding similarly, and there is significant spatial variation in responses even within species. This spatial and interspecific variation complicates efforts to predict phenological responses to ongoing climate change, but must be incorporated in order to build reliable forecasts. Here, we use a long-term dataset (1953–2005) of plant phenological events in spring (flowering and leaf out) and autumn (leaf colouring and leaf fall) throughout Japan and South Korea to build forecasts that account for these sources of variability. Specifically, we used hierarchical models to incorporate the spatial variability in phenological responses to temperature to then forecast species'' overall and site-specific responses to global warming. We found that for most species, spring phenology is advancing and autumn phenology is getting later, with the timing of events changing more quickly in autumn compared with the spring. Temporal trends and phenological responses to temperature in East Asia contrasted with results from comparable studies in Europe, where spring events are changing more rapidly than are autumn events. Our results emphasize the need to study multiple species at many sites to understand and forecast regional changes in phenology.     

Early pest development and loss of biological control are associated with urban warming

Climate warming is predicted to cause many changes in ectotherm communities, one of which is phenological mismatch, wherein one species'' development advances relative to an associated species or community. Phenological mismatches already lead to loss of pollination services, and we predict that they also cause loss of biological control. Here, we provide evidence that a pest develops earlier due to urban warming but that phenology of its parasitoid community does not similarly advance. This mismatch is associated with greater egg production that likely leads to more pests on trees.     

Prairie plant phenology driven more by temperature than moisture in climate manipulations across a latitudinal gradient in the Pacific Northwest,USA

Plant phenology will likely shift with climate change, but how temperature and/or moisture regimes will control phenological responses is not well understood. This is particularly true in Mediterranean climate ecosystems where the warmest temperatures and greatest moisture availability are seasonally asynchronous. We examined plant phenological responses at both the population and community levels to four climate treatments (control, warming, drought, and warming plus additional precipitation) embedded within three prairies across a 520 km latitudinal Mediterranean climate gradient within the Pacific Northwest, USA. At the population level, we monitored flowering and abundances in spring 2017 of eight range‐restricted focal species planted both within and north of their current ranges. At the community level, we used normalized difference vegetation index (NDVI) measured from fall 2016 to summer 2018 to estimate peak live biomass, senescence, seasonal patterns, and growing season length. We found that warming exerted a stronger control than our moisture manipulations on phenology at both the population and community levels. Warming advanced flowering regardless of whether a species was within or beyond its current range. Importantly, many of our focal species had low abundances, particularly in the south, suggesting that establishment, in addition to phenological shifts, may be a strong constraint on their future viability. At the community level, warming advanced the date of peak biomass regardless of site or year. The date of senescence advanced regardless of year for the southern and central sites but only in 2018 for the northern site. Growing season length contracted due to warming at the southern and central sites (~3 weeks) but was unaffected at the northern site. Our results emphasize that future temperature changes may exert strong influence on the timing of a variety of plant phenological events, especially those events that occur when temperature is most limiting, even in seasonally water‐limited Mediterranean ecosystems.     

Phenological indices of avian reproduction: cryptic shifts and prediction across large spatial and temporal scales

Climate change‐induced shifts in phenology have important demographic consequences, and are frequently used to assess species' sensitivity to climate change. Therefore, developing accurate phenological predictions is an important step in modeling species' responses to climate change. The ability of such phenological models to predict effects at larger spatial and temporal scales has rarely been assessed. It is also not clear whether the most frequently used phenological index, namely the average date of a phenological event across a population, adequately captures phenological shifts in the distribution of events across the season. We use the long‐tailed tit Aegithalos caudatus (Fig. 1) as a case study to explore these issues. We use an intensive 17‐year local study to model mean breeding date and test the capacity of this local model to predict phenology at larger spatial and temporal scales. We assess whether local models of breeding initiation, termination, and renesting reveal phenological shifts and responses to climate not detected by a standard phenological index, that is, population average lay date. These models take predation timing/intensity into account. The locally‐derived model performs well at predicting phenology at the national scale over several decades, at both high and low temperatures. In the local model, a trend toward warmer Aprils is associated with a significant advance in termination dates, probably in response to phenological shifts in food supply. This results in a 33% reduction in breeding season length over 17 years – a substantial loss of reproductive opportunity that is not detected by the index of population average lay date. We show that standard phenological indices can fail to detect patterns indicative of negative climatic effects, potentially biasing assessments of species' vulnerability to climate change. More positively, we demonstrate the potential of detailed local studies for developing broader‐scale predictive models of future phenological shifts.     

The importance of phylogeny to the study of phenological response to global climate change

Climate change has resulted in major changes in the phenology—i.e. the timing of seasonal activities, such as flowering and bird migration—of some species but not others. These differential responses have been shown to result in ecological mismatches that can have negative fitness consequences. However, the ways in which climate change has shaped changes in biodiversity within and across communities are not well understood. Here, we build on our previous results that established a link between plant species'' phenological response to climate change and a phylogenetic bias in species'' decline in the eastern United States. We extend a similar approach to plant and bird communities in the United States and the UK that further demonstrates that climate change has differentially impacted species based on their phylogenetic relatedness and shared phenological responses. In plants, phenological responses to climate change are often shared among closely related species (i.e. clades), even between geographically disjunct communities. And in some cases, this has resulted in a phylogenetically biased pattern of non-native species success. In birds, the pattern of decline is phylogenetically biased but is not solely explained by phenological response, which suggests that other traits may better explain this pattern. These results illustrate the ways in which phylogenetic thinking can aid in making generalizations of practical importance and enhance efforts to predict species'' responses to future climate change.     

Temperature and snowfall trigger alpine vegetation green‐up on the world's roof

Rapid temperature increase and its impacts on alpine ecosystems in the Qinghai–Tibetan Plateau, the world's highest and largest plateau, are a matter of global concern. Satellite observations have revealed distinctly different trend changes and contradicting temperature responses of vegetation green‐up dates, leading to broad debate about the Plateau's spring phenology and its climatic attribution. Large uncertainties in remote‐sensing estimates of phenology significantly limit efforts to predict the impacts of climate change on vegetation growth and carbon balance in the Qinghai–Tibetan Plateau, which are further exacerbated by a lack of detailed ground observation calibration. Here, we revealed the spatiotemporal variations and climate drivers of ground‐based herbaceous plant green‐up dates using 72 green‐up datasets for 22 herbaceous plant species at 23 phenological stations, and corresponding daily mean air temperature and daily precipitation data from 19 climate stations across eastern and southern parts of the Qinghai–Tibetan Plateau from 1981 to 2011. Results show that neither the continuously advancing trend from 1982 to 2011, nor a turning point in the mid to late 1990s as reported by remote‐sensing studies can be verified by most of the green‐up time series, and no robust evidence for a warmer winter‐induced later green‐up dates can be detected. Thus, chilling requirements may not be an important driver influencing green‐up responses to spring warming. Moreover, temperature‐only control of green‐up dates appears mainly at stations with relatively scarce preseason snowfall and lower elevation, while coupled temperature and precipitation controls of green‐up dates occur mostly at stations with relatively abundant preseason snowfall and higher elevation. The diversified interactions between snowfall and temperature during late winter to early spring likely determine the spatiotemporal variations of green‐up dates. Therefore, prediction of vegetation growth and carbon balance responses to global climate change on the world's roof should integrate both temperature and snowfall variations.     

Disentangling the paradox of insect phenology: are temporal trends reflecting the response to warming?

The strength and direction of phenological responses to changes in climate have been shown to vary significantly both among species and among populations of a species, with the overall patterns not fully resolved. Here, we studied the temporal and spatial variability associated with the response of several insect species to recent global warming. We use hierarchical models within a model comparison framework to analyze phenological data gathered over 40 years by the Japan Meteorological Agency on the emergence dates of 14 insect species at sites across Japan. Contrary to what has been predicted with global warming, temporal trends of annual emergence showed a later emergence day for some species and sites over time, even though temperatures are warming. However, when emergence data were analyzed as a function of temperature and precipitation, the overall response pointed out an earlier emergence day with warmer conditions. The apparent contradiction between the response to temperature and trends over time indicates that other factors, such as declining populations, may be affecting the date phenological events are being recorded. Overall, the responses by insects were weaker than those found for plants in previous work over the same time period in these ecosystems, suggesting the potential for ecological mismatches with deleterious effects for both suites of species. And although temperature may be the major driver of species phenology, we should be cautious when analyzing phenological datasets as many other factors may also be contributing to the variability in phenology.     

Evaluating within‐population variability in behavior and demography for the adaptive potential of a dispersal‐limited species to climate change

Multiple pathways exist for species to respond to changing climates. However, responses of dispersal‐limited species will be more strongly tied to ability to adapt within existing populations as rates of environmental change will likely exceed movement rates. Here, we assess adaptive capacity in Plethodon cinereus, a dispersal‐limited woodland salamander. We quantify plasticity in behavior and variation in demography to observed variation in environmental variables over a 5‐year period. We found strong evidence that temperature and rainfall influence P. cinereus surface presence, indicating changes in climate are likely to affect seasonal activity patterns. We also found that warmer summer temperatures reduced individual growth rates into the autumn, which is likely to have negative demographic consequences. Reduced growth rates may delay reproductive maturity and lead to reductions in size‐specific fecundity, potentially reducing population‐level persistence. To better understand within‐population variability in responses, we examined differences between two common color morphs. Previous evidence suggests that the color polymorphism may be linked to physiological differences in heat and moisture tolerance. We found only moderate support for morph‐specific differences for the relationship between individual growth and temperature. Measuring environmental sensitivity to climatic variability is the first step in predicting species' responses to climate change. Our results suggest phenological shifts and changes in growth rates are likely responses under scenarios where further warming occurs, and we discuss possible adaptive strategies for resulting selective pressures.     

Links between plant species’ spatial and temporal responses to a warming climate

To generate realistic projections of species’ responses to climate change, we need to understand the factors that limit their ability to respond. Although climatic niche conservatism, the maintenance of a species’s climatic niche over time, is a critical assumption in niche-based species distribution models, little is known about how universal it is and how it operates. In particular, few studies have tested the role of climatic niche conservatism via phenological changes in explaining the reported wide variance in the extent of range shifts among species. Using historical records of the phenology and spatial distribution of British plants under a warming climate, we revealed that: (i) perennial species, as well as those with weaker or lagged phenological responses to temperature, experienced a greater increase in temperature during flowering (i.e. failed to maintain climatic niche via phenological changes); (ii) species that failed to maintain climatic niche via phenological changes showed greater northward range shifts; and (iii) there was a complementary relationship between the levels of climatic niche conservatism via phenological changes and range shifts. These results indicate that even species with high climatic niche conservatism might not show range shifts as instead they track warming temperatures during flowering by advancing their phenology.     

Predicting small‐mammal responses to climatic warming: autecology,geographic range,and the Holocene fossil record

Forecasting how species will respond to climatic change requires knowledge of past community dynamics. Here we use time‐series data from the small‐mammal fossil records of two caves in the Great Basin of the American West to evaluate how contrasting and variable local paleoclimates have shaped small‐mammal abundance dynamics over the last ～7500 years of climatic change. We then predict how species and communities will respond to future scenarios of increased warming and aridity coupled with continued spread of an invasive annual grass (Bromus tectorum). We find that most community‐level responses to climatic change occur in the mammalian abundance structure at both sites; the dominance of the community by individuals from species with a southern geographic affinity increases with climatic warming. This suggests that responses occurred in situ rather than by the immigration of new taxa over this time interval. Despite predictability at the community‐scale, species‐level relationships between abundance and climate are variable and are not necessarily explained by a species' geographic affinity. Species present at both sites, however, exhibit remarkably similar responses to climate at each site, indicating that species autecology (specifically dietary functional group) is important in determining response to climatic warming. Regression‐tree analyses show remarkable concordance between the two cave faunas and highlight the importance of a granivorous dietary strategy in this desert ecosystem. Under projections of increased temperature and decreased precipitation over the next 50 years, our results indicate that granivores should thrive as communities become more dominated by individuals with a southern geographic affinity. Granivores, however, are negatively impacted by the invasion of cheatgrass. The last century of anthropogenic impacts has thus placed granivores at a greater risk of extinction than predicted under climate‐only scenarios.     

Contrasting effects of warming and increased snowfall on Arctic tundra plant phenology over the past two decades

Recent changes in climate have led to significant shifts in phenology, with many studies demonstrating advanced phenology in response to warming temperatures. The rate of temperature change is especially high in the Arctic, but this is also where we have relatively little data on phenological changes and the processes driving these changes. In order to understand how Arctic plant species are likely to respond to future changes in climate, we monitored flowering phenology in response to both experimental and ambient warming for four widespread species in two habitat types over 21 years. We additionally used long‐term environmental records to disentangle the effects of temperature increase and changes in snowmelt date on phenological patterns. While flowering occurred earlier in response to experimental warming, plants in unmanipulated plots showed no change or a delay in flowering over the 21‐year period, despite more than 1 °C of ambient warming during that time. This counterintuitive result was likely due to significantly delayed snowmelt over the study period (0.05–0.2 days/yr) due to increased winter snowfall. The timing of snowmelt was a strong driver of flowering phenology for all species – especially for early‐flowering species – while spring temperature was significantly related to flowering time only for later‐flowering species. Despite significantly delayed flowering phenology, the timing of seed maturation showed no significant change over time, suggesting that warmer temperatures may promote more rapid seed development. The results of this study highlight the importance of understanding the specific environmental cues that drive species’ phenological responses as well as the complex interactions between temperature and precipitation when forecasting phenology over the coming decades. As demonstrated here, the effects of altered snowmelt patterns can counter the effects of warmer temperatures, even to the point of generating phenological responses opposite to those predicted by warming alone.     

Warming increases the spread of an invasive thistle

Background

Global warming and shifted precipitation regimes increasingly affect species abundances and distributions worldwide. Despite a large literature on species'' physiological, phenological, growth, and reproductive responses to such climate change, dispersal is rarely examined. Our study aims to test whether the dispersal ability of a non-native, wind-dispersed plant species is affected by climate change, and to quantify the ramifications for future invasion spread rates.

Methodology/Principal Findings

We experimentally increased temperature and precipitation in a two-cohort, factorial field study (n = 80). We found an overwhelming warming effect on plant life history: warming not only improved emergence, survival, and reproduction of the thistle Carduus nutans, but also elevated plant height, which increased seed dispersal distances. Using spatial population models, we demonstrate that these empirical warming effects on demographic vital rates, and dispersal parameters, greatly exacerbate spatial spread. Predicted levels of elevated winter precipitation decreased seed production per capitulum, but this only slightly offset the warming effect on spread. Using a spread rate decomposition technique (c*-LTRE), we also found that plant height-mediated changes in dispersal contribute most to increased spread rate under climate change.

Conclusions/Significance

We found that both dispersal and spread of this wind-dispersed plant species were strongly impacted by climate change. Dispersal responses to climate change can improve, or diminish, a species'' ability to track climate change spatially, and should not be overlooked. Methods that combine both demographic and dispersal responses thus will be an invaluable complement to projections of suitable habitat under climate change.     

A comprehensive overview of the spatial and temporal variability of apple bud dormancy release and blooming phenology in Western Europe

In the current context of global warming, an analysis is required of spatially-extensive and long-term blooming data in fruit trees to make up for insufficient information on regional-scale blooming changes and determinisms that are key to the phenological adaptation of these species. We therefore analysed blooming dates over long periods at climate-contrasted sites in Western Europe, focusing mainly on the Golden Delicious apple that is grown worldwide. On average, blooming advances were more pronounced in northern continental (10 days) than in western oceanic (6–7 days) regions, while the shortest advance was found on the Mediterranean coastline. Temporal trends toward blooming phase shortenings were also observed in continental regions. These regional differences in temporal variability across Western Europe resulted in a decrease in spatial variability, i.e. shorter time intervals between blooming dates in contrasted regions (8–10-day decrease for full bloom between Mediterranean and continental regions). Fitted sequential models were used to reproduce phenological changes. Marked trends toward shorter simulated durations of forcing period (bud growth from dormancy release to blooming) and high positive correlations between these durations and observed blooming dates support the notion that blooming advances and shortenings are mainly due to faster satisfaction of the heating requirement. However, trends toward later dormancy releases were also noted in oceanic and Mediterranean regions. This could tend toward blooming delays and explain the shorter advances in these regions despite similar or greater warming. The regional differences in simulated chilling and forcing periods were consistent with the regional differences in temperature increases.     

Macroevolutionary consequences of profound climate change on niche evolution in marine molluscs over the past three million years

In order to predict the fate of biodiversity in a rapidly changing world, we must first understand how species adapt to new environmental conditions. The long-term evolutionary dynamics of species'' physiological tolerances to differing climatic regimes remain obscure. Here, we unite palaeontological and neontological data to analyse whether species'' environmental tolerances remain stable across 3 Myr of profound climatic changes using 10 phylogenetically, ecologically and developmentally diverse mollusc species from the Atlantic and Gulf Coastal Plains, USA. We additionally investigate whether these species'' upper and lower thermal tolerances are constrained across this interval. We find that these species'' environmental preferences are stable across the duration of their lifetimes, even when faced with significant environmental perturbations. The results suggest that species will respond to current and future warming either by altering distributions to track suitable habitat or, if the pace of change is too rapid, by going extinct. Our findings also support methods that project species'' present-day environmental requirements to future climatic landscapes to assess conservation risks.     

Similarities in butterfly emergence dates among populations suggest local adaptation to climate

Phenology shifts are the most widely cited examples of the biological impact of climate change, yet there are few assessments of potential effects on the fitness of individual organisms or the persistence of populations. Despite extensive evidence of climate‐driven advances in phenological events over recent decades, comparable patterns across species' geographic ranges have seldom been described. Even fewer studies have quantified concurrent spatial gradients and temporal trends between phenology and climate. Here we analyse a large data set (~129 000 phenology measures) over 37 years across the UK to provide the first phylogenetic comparative analysis of the relative roles of plasticity and local adaptation in generating spatial and temporal patterns in butterfly mean flight dates. Although populations of all species exhibit a plastic response to temperature, with adult emergence dates earlier in warmer years by an average of 6.4 days per °C, among‐population differences are significantly lower on average, at 4.3 days per °C. Emergence dates of most species are more synchronised over their geographic range than is predicted by their relationship between mean flight date and temperature over time, suggesting local adaptation. Biological traits of species only weakly explained the variation in differences between space‐temperature and time‐temperature phenological responses, suggesting that multiple mechanisms may operate to maintain local adaptation. As niche models assume constant relationships between occurrence and environmental conditions across a species' entire range, an important implication of the temperature‐mediated local adaptation detected here is that populations of insects are much more sensitive to future climate changes than current projections suggest.     

Long-term temporal changes of plant phenology in the Western Mediterranean

Plants are altering their life cycles in response to current climatic change around the globe. More than 200 000 records for six phenological events (leaf unfolding, flowering, fruit ripening, fruit harvesting, leaf falling and growing season) of 29 perennial species for the period 1943–2003 recorded throughout Spain provide the longest temporal and the broadest spatial assessment of plant phenology changes in the Mediterranean region. The overwhelming majority of the 118 studied phenophases shifted their dates in recent decades. Such changes differed among phenological events. Leaf unfolding, flowering and fruiting are markedly advancing (?0.48, ?0.59 and ?0.32 days yr^?1, respectively), but only since the mid‐1970s. Anemophilous have advanced more days their flowering than entomophilous. However, some species have delayed and others have advanced their leaf falling dates and as a result only a weak shift was observed in this event for the whole of the studied species (+0.12 days yr^?1). The growing season lengthened by 18 days, which implies an increase of 8% in the life of annual leaves. Such an increase was achieved mainly through the advance of leaf unfolding dates in the spring, one of the most productive times of year for vegetation in the Mediterranean. Shifts in the plant calendar were accompanied as well by long‐term changes in the range of onset dates in 39% of studied phenophases. Leaf unfolding, flowering and growing season tended to reduce spatial variability, reflecting a faster and more synchronized onset (or duration) of phenophases across the study area. Changes in spatial variability may aggravate calendar mismatching with other trophic levels resulting from changes in dates. Because temporal responses differed markedly among species, calendar guilds of plants have changed, which suggests alterations of interspecific relationships in plant communities from Mediterranean ecosystems.     

Phenology in a warming world: differences between native and non‐native plant species

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non‐native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non‐native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non‐native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non‐native species, implicating phenology as a potential trait associated with the successful establishment of non‐native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non‐natives are hypothesised to promote invasion success and population persistence, potentially benefiting non‐native over native species under climate change.     

Chilling outweighs photoperiod in preventing precocious spring development

It is well known that increased spring temperatures cause earlier onset dates of leaf unfolding and flowering. However, a temperature increase in winter may be associated with delayed development when species' chilling requirements are not fulfilled. Furthermore, photosensitivity is supposed to interfere with temperature triggers. To date, neither the relative importance nor possible interactions of these three factors have been elucidated. In this study, we present a multispecies climate chamber experiment to test the effects of chilling and photoperiod on the spring phenology of 36 woody species. Several hypotheses regarding their variation with species traits (successional strategy, floristic status, climate of their native range) were tested. Long photoperiods advanced budburst for one‐third of the studied species, but magnitudes of these effects were generally minor. In contrast to prior hypotheses, photosensitive responses were not restricted to climax or oceanic species. Increased chilling length advanced budburst for almost all species; its effect greatly exceeding that of photoperiod. Moreover, we suggest that photosensitivity and chilling effects have to be rigorously disentangled, as the response to photoperiod was restricted to individuals that had not been fully chilled. The results indicate that temperature requirements and successional strategy are linked, with climax species having higher chilling and forcing requirements than pioneer species. Temperature requirements of invasive species closely matched those of native species, suggesting that high phenological concordance is a prerequisite for successful establishment. Lack of chilling not only led to a considerable delay in budburst but also caused substantial changes in the chronological order of species' budburst. The results reveal that increased winter temperatures might impact forest ecosystems more than formerly assumed. Species with lower chilling requirements, such as pioneer or invasive species, might profit from warming winters, if late spring frost events would in parallel occur earlier.     

The roles of shifting and filtering in generating community‐level flowering phenology

Plant phenologies are key components of community assembly and ecosystem function, yet we know little about how phenological patterns differ among ecosystems. Community‐level phenological patterns may be driven by the filtering of species into communities based on their phenology or by intraspecific responses to local conditions that shift when species flower. To understand the relative roles of filtering and shifting on community‐level phenological patterns we compared patterns of first flowering dates (FFD) for herbaceous species at Konza Prairie, KS, USA with those from the colder Fargo, ND, USA area and from Chinnor, England, which has a less continental climate. Comparing patterns of FFD supports that Konza's flowering patterns are potentially influenced both by filtering species that flower early in the growing season and by phenological shifting. Konza species flowering dates were earlier in the spring and later in the fall compared to Fargo, but were not shifted compared to Chinnor, which had a unique suite of early‐flowering species. In all, comparing flowering phenology among three sites reveals that intraspecific responses to climate can generate phenological shifts that compress or stretch community‐level phenological patterns, while novel niches in phenological space can also alter community‐level patterns. Community flowering patterns related to climate suggest that climatic warming has the potential to further distribute flowering of the Konza flora over a longer period, but also could further open it to introductions of non‐native species that have evolved to flower early in the season.