Variable flowering phenology and pollinator use in a community suggest future phenological mismatch

Recent anthropogenic climate change is strongly associated with average shifts toward earlier seasonal timing of activity (phenology) in temperate-zone species. Shifts in phenology have the potential to alter ecological interactions, to the detriment of one or more interacting species. Recent models predict that detrimental phenological mismatch may increasingly occur between plants and their pollinators. One way to test this prediction is to examine data from ecological communities that experience large annual weather fluctuations. Taking this approach, we analyzed interactions over a four-year period among 132 plant species and 665 pollinating insect species within a Mediterranean community. For each plant species we recorded onset and duration of flowering and number of pollinator species. Flowering onset varied among years, and a year of earlier flowering of a species tended to be a year of fewer species pollinating its flowers. This relationship was attributable principally to early-flowering species, suggesting that shifts toward earlier phenology driven by climate change may reduce pollination services due to phenological mismatch. Earlier flowering onset of a species also was associated with prolonged flowering duration, but it is not certain that this will counterbalance any negative effects of lower pollinator species richness on plant reproductive success. Among plants with different life histories, annuals were more severely affected by flowering–pollinator mismatches than perennials. Specialized plant species (those attracting a smaller number of pollinator species) did not experience disproportionate interannual fluctuations in phenology. Thus they do not appear to be faced with disproportionate fluctuations in pollinator species richness, contrary to the expectation that specialists are at greatest risk of losing mutualistic interactions because of climate change.     

Phenotypic plasticity and adaptive evolution contribute to advancing flowering phenology in response to climate change

Anthropogenic climate change has already altered the timing of major life-history transitions, such as the initiation of reproduction. Both phenotypic plasticity and adaptive evolution can underlie rapid phenological shifts in response to climate change, but their relative contributions are poorly understood. Here, we combine a continuous 38 year field survey with quantitative genetic field experiments to assess adaptation in the context of climate change. We focused on Boechera stricta (Brassicaeae), a mustard native to the US Rocky Mountains. Flowering phenology advanced significantly from 1973 to 2011, and was strongly associated with warmer temperatures and earlier snowmelt dates. Strong directional selection favoured earlier flowering in contemporary environments (2010-2011). Climate change could drive this directional selection, and promote even earlier flowering as temperatures continue to increase. Our quantitative genetic analyses predict a response to selection of 0.2 to 0.5 days acceleration in flowering per generation, which could account for more than 20 per cent of the phenological change observed in the long-term dataset. However, the strength of directional selection and the predicted evolutionary response are likely much greater now than even 30 years ago because of rapidly changing climatic conditions. We predict that adaptation will likely be necessary for long-term in situ persistence in the context of climate change.     

Clinal variation for only some phenological traits across a species range

Phenology is the timing of life cycle events of an organism. Alterations in phenology can have profound effects on individual fitness, population growth, and community dynamics. Recent changes in climate have altered the phenology of many organisms, which may result in selection to shift phenological traits. Understanding the relationship between local climates and population differentiation in phenology will allow us to anticipate responses to novel selective environments caused by global climate change. We evaluated population differentiation in the number of days to germination, first flower, and fruit maturation for 33 populations throughout the range of Campanulastrum americanum (American Bellflower). Germination and fruit maturation had geographical clines with earlier timing in populations from northern latitudes. Northern sites were cooler and drier, suggesting potential adaptive differentiation of the shorter life cycle associated with earlier phenology. Similarly, higher elevations were cooler and had earlier fruit maturation. However, seed germination was later in higher elevation populations. Although there was substantial variation in the day to first flower, ranging 40 days between population means, it was idiosyncratic and not related to latitude, suggesting differentiation in response to selective factors distinct from those on germination and fruit maturation. Thus, germination and fruit maturation in C. americanum may shift in response to selection by rising temperatures. However, such changes are not expected for flowering time, a typical indicator of climate change.     

Climatic Variability Leads to Later Seasonal Flowering of Floridian Plants

Understanding species responses to global change will help predict shifts in species distributions as well as aid in conservation. Changes in the timing of seasonal activities of organisms over time may be the most responsive and easily observable indicator of environmental changes associated with global climate change. It is unknown how global climate change will affect species distributions and developmental events in subtropical ecosystems or if climate change will differentially favor nonnative species. Contrary to previously observed trends for earlier flowering onset of plant species with increasing spring temperatures from mid and higher latitudes, we document a trend for delayed seasonal flowering among plants in Florida. Additionally, there were few differences in reproductive responses by native and nonnative species to climatic changes. We argue that plants in Florida have different reproductive cues than those from more northern climates. With global change, minimum temperatures have become more variable within the temperate-subtropical zone that occurs across the peninsula and this variation is strongly associated with delayed flowering among Florida plants. Our data suggest that climate change varies by region and season and is not a simple case of species responding to consistently increasing temperatures across the region. Research on climate change impacts need to be extended outside of the heavily studied higher latitudes to include subtropical and tropical systems in order to properly understand the complexity of regional and seasonal differences of climate change on species responses.     

Climate drives among‐year variation in natural selection on flowering time

To predict long‐term responses to climate change, we need to understand how changes in temperature and precipitation elicit both immediate phenotypic responses and changes in natural selection. We used 22 years of data for the perennial herb Lathyrus vernus to examine how climate influences flowering phenology and phenotypic selection on phenology. Plants flowered earlier in springs with higher temperatures and higher precipitation. Early flowering was associated with a higher fitness in nearly all years, but selection for early flowering was significantly stronger in springs with higher temperatures and lower precipitation. Climate influenced selection through trait distributions, mean fitness and trait?fitness relationships, the latter accounting for most of the among‐year variation in selection. Our results show that climate both induces phenotypic responses and alters natural selection, and that the change in the optimal phenotype might be either weaker, as for spring temperature, or stronger, as for precipitation, than the optimal response.     

Later flowering is associated with a compressed flowering season and reduced reproductive output in an early season floral resource

Climate change‐induced shifts in flowering phenology can expose plants to novel biotic and abiotic environments, potentially leading to decreased temporal overlap with pollinators and exposure to conditions that negatively affect fruit and seed set. We explored the relationship between flowering phenology and reproductive output in the common shrub pointleaf manzanita Arctostaphylos pungens in a lower montane habitat in southeastern Arizona, USA. Contrary to the pattern of progressively earlier flowering observed in many species, long‐term records show that A. pungens flowering onset is shifting later and the flowering season is being compressed. This species can thus provide unusual insight into the effects of altered phenology. To determine the consequences of among‐ and within‐plant variation in flowering time, we documented individual flowering schedules and followed the fates of flowers on over 50 plants throughout two seasons (2012 and 2013). We also measured visitation rates by potential pollinators in 2012, as well as both fruit mass and seeds per fruit of flowers produced at different times. Fruit set was positively related to visitation rate but declined with later dates of flower production in both years. Total fruit production per plant was positively influenced by flowering duration, which declined with later flowering onset, as did fruit mass. Individual flowering schedules were consistent between years, suggesting that plants that begin flowering late have lower reproductive output each year. These patterns suggest that if pointleaf manzanita flowering continues to shift later, its flowering season may continue to become shorter, compressing floral resource availability for pollinators and leading to reduced reproductive output. These results reveal the negative effects of delayed phenology on reproductive output in a long‐lived plant. They highlight the value of using natural variation in flowering time, in combination with long‐term data, to anticipate the consequences of phenological shifts.     

The shape of selection: using alternative fitness functions to test predictions for selection on flowering time

Selection gradient analysis examines the strength and direction of phenotypic selection as well as the curvature of fitness functions, allowing predictions on and insights into the process of evolution in natural populations. However, traditional linear and quadratic selection analyses are not capable of detecting other features of fitness functions, such as asymmetry or thresholds, which may be relevant for understanding key aspects of selection on many traits. In these cases, additional analyses are needed to test specific hypotheses about fitness functions. In this study we used several approaches to analyze selection on a major life-history trait—flowering time—in the annual plant Brassica rapa subjected to experimentally abbreviated and lengthened growing seasons. We used a model that incorporated a tradeoff between the time allocated to growth versus the time allocated to reproduction in order to predict fitness function shape. The model predicted that optimal flowering time shifts to earlier and later dates as the growing season contracts and expands. It also showed the flowering time fitness function to be asymmetrical: reproductive output increases modestly between the earliest and the optimal flowering date, but then falls sharply with later dates, truncating in a ‘tail of zeros’. Our experimental results strongly supported selection for early flowering in short season and selection for late flowering in long season conditions. We also found support for the predicted asymmetry of the flowering time fitness function, including a ‘tail of zeros’ at later flowering dates. The form of the fitness function revealed here has implications for interpreting estimates of selection on flowering time in natural populations and for refining predictions on evolutionary response to climate change. More generally, this study illustrates the value of diverse statistical approaches to understanding mechanisms of natural selection.     

Selection on flowering time in Mediterranean high-mountain plants under global warming

Under climate warming, plants will undergo novel selective pressures to adjust reproductive timing. Adjustment between reproductive phenology and environment is expected to be higher in arctic and alpine habitats because the growing season is considerably short. As early- and late-flowering species reproduce under very different environmental conditions, selective pressures on flowering phenology and potential effects of climate change are likely to differ between them. However, there is no agreement on the magnitude of the benefits and costs of early- vs. late-flowering species under a global warming scenario. In spite of its relevance, phenotypic selection on flowering phenology has rarely been explored in alpine plants and never in Mediterranean high mountain species, where selective pressures are very different due to the summer drought imposed over the short growth season. We hypothesized that late-flowering plants in Mediterranean mountains should present stronger selective pressures towards early onset of reproduction than early-flowering species, because less water is available in the soil as growing season progresses. We performed selection analyses on flowering onset and duration in two high mountain species of contrasting phenology. Since phenotypic selection can be highly context-dependent, we studied several populations of each species for 2 years, covering their local altitudinal ranges and their different microhabitats. Surrogates of biotic selective agents, like fruitset for pollinators and flower and fruit loss for flower and seed predators, were included in the analysis. Differences between the early- and the late-flowering species were less than expected. A consistent negative correlational selection of flowering onset and duration was found affecting plant fitness, i.e., plants that bloomed earlier flowered for longer periods improving plant fitness. Nevertheless, the late-flowering species may experience higher risks under climate warming because in extremely warm and dry years the earlier season does not bring about a longer flowering duration due to summer drought.     

Cascading effects of climate extremes on vertebrate fauna through changes to low‐latitude tree flowering and fruiting phenology

Forest vertebrate fauna provide critical services, such as pollination and seed dispersal, which underpin functional and resilient ecosystems. In turn, many of these fauna are dependent on the flowering phenology of the plant species of such ecosystems. The impact of changes in climate, including climate extremes, on the interaction between these fauna and flora has not been identified or elucidated, yet influences on flowering phenology are already evident. These changes are well documented in the mid to high latitudes. However, there is emerging evidence that the flowering phenology, nectar/pollen production, and fruit production of long‐lived trees in tropical and subtropical forests are also being impacted by changes in the frequency and severity of climate extremes. Here, we examine the implications of these changes for vertebrate fauna dependent on these resources. We review the literature to establish evidence for links between climate extremes and flowering phenology, elucidating the nature of relationships between different vertebrate taxa and flowering regimes. We combine this information with climate change projections to postulate about the likely impacts on nectar, pollen and fruit resource availability and the consequences for dependent vertebrate fauna. The most recent climate projections show that the frequency and intensity of climate extremes will increase during the 21st century. These changes are likely to significantly alter mass flowering and fruiting events in the tropics and subtropics, which are frequently cued by climate extremes, such as intensive rainfall events or rapid temperature shifts. We find that in these systems the abundance and duration of resource availability for vertebrate fauna is likely to fluctuate, and the time intervals between episodes of high resource availability to increase. The combined impact of these changes has the potential to result in cascading effects on ecosystems through changes in pollinator and seed dispersal ecology, and demands a focused research effort.     

Flowering phenology change and climate warming in southwestern Ohio

Global surface temperature has increased markedly over the last 100 years. This increase has a variety of implications for human societies, and for ecological systems. One of the most obvious ways ecosystems are affected by global climate change is through alteration of organisms’ developmental timing (phenology). We used annual botanical surveys that documented the first flowering for an array of species from 1976 to 2003 to examine the potential implications of climate change for plant development. The overall trend for these species was a progressively earlier flowering time. The two earliest flowering taxa (Galanthus and Crocus) also exhibited the strongest shift in first flowering. We detected a significant trend in climate suggesting higher temperatures in winter and spring over the sampling interval and found a significant relationship between warming temperatures and first flowering time for some species. Although 60% of the species in our study flowered earlier over the sampling interval, the remaining species exhibited no statistically detectable change. This variation in response is ostensibly associated with among-species variation in the role of climate cues in plant development. Future work is needed to isolate specific climate cues, and to link plant phenology to the physiological processes that trigger plant development.     

Effects of experimental shifts in flowering phenology on plant-pollinator interactions

Climate change has led to phenological shifts in flowering plants and insect pollinators, causing concern that these shifts will disrupt plant-pollinator mutualisms. We experimentally investigated how shifts in flowering onset affect pollinator visitation for 14 native perennial plant species, six of which have exhibited shifts to earlier flowering over the last 70 years and eight of which have not. We manipulated flowering onset in greenhouses and then observed pollinator visitation in the field. Five of six species with historically advanced flowering received more visits when flowering was experimentally advanced, whereas seven of eight species with historically unchanged flowering received fewer visits when flowering earlier. This pattern suggests that species unconstrained by pollinators have advanced their flowering, whereas species constrained by pollinators have not. In contrast to current concern about phenological mismatches disrupting plant-pollinator mutualisms, mismatches at the onset of flowering are not occurring for most of our study species.     

Re-establishment of clinal variation in flowering time among introduced populations of purple loosestrife (Lythrum salicaria, Lythraceae)

Range expansion during biological invasion requires that invaders adapt to geographical variation in climate, which should yield latitudinal clines in reproductive phenology. We investigated geographic variation in life history among 25 introduced populations of Lythrum salicaria, a widespread European invader of North American wetlands. We detected a strong latitudinal cline in initiation of flowering and size at flowering, which paralleled that reported among native populations. Plants from higher latitudes flowered earlier and at a smaller size than those from lower latitudes, even when raised in a uniform glasshouse. Early flowering was associated with greatly reduced reproductive output, but this was not associated with latitudinal variation in abundance, and probably did not result from a genetic correlation between time to and size at flowering. As introduction to North America c. 200 years ago, L. salicaria has re-established latitudinal clines in life history, probably as an evolutionary response to climatic selection.     

FLOWERING,SEX RATIOS,POLLEN-OVULE RATIOS,FRUIT SET,AND REPRODUCTIVE EFFORT OF A DIOECIOUS TREE,MYRISTICA INSIPIDA (MYRISTICACEAE), IN TWO DIFFERENT RAIN FOREST COMMUNITIES

The flowering of Myristica insipida R. Br. was studied in two rain forest communities in northern Queensland. This dioecious, subcanopy tree had a male-biased sex ratio at both study sites. In the lowland population the male-bias could be attributed to males (trees producing staminate flowers) starting to flower at a smaller average size than females (trees producing pistillate flowers). There were no intersexual differences in spacing or distribution within the study sites. Males trees flowered earlier, flowered longer, and produced over twice as many flowers as females during the study season. Although the onset of flowering was rather variable, 18–22 days following heavy rains, most trees had a synchronous period of maximum flowering. Pollination manipulations determined that there was no fruit development without pollination, and that increasing pollen loads resulted in increased fruit set with diminishing effect. Taking into account the sex-ratios and intersexual differences in flower production, the pollen-ovule ratio was calculated to be 16,000–19,000. Male trees were found to expend more energy on flowering than female trees. Open-pollination resulted in 1.0% of female flowers setting fruit. The much greater cost of fruit production resulted in females expending 421% more energy on reproduction than males. Fruit and seed production were judged to be pollination-limited. Nonetheless, this species exhibited several characteristics that are predicted if dioecy evolved by means of sexual selection.     

Solar radiation and ENSO predict fruiting phenology patterns in a 15‐year record from Kibale National Park,Uganda

Fruiting, flowering, and leaf set patterns influence many aspects of tropical forest communities, but there are few long‐term studies examining potential drivers of these patterns, particularly in Africa. We evaluated a 15‐year dataset of tree phenology in Kibale National Park, Uganda, to identify abiotic predictors of fruit phenological patterns and discuss our findings in light of climate change. We quantified fruiting for 326 trees from 43 species and evaluated these patterns in relation to solar radiance, rainfall, and monthly temperature. We used time‐lagged variables based on seasonality in linear regression models to assess the effect of abiotic variables on the proportion of fruiting trees. Annual fruiting varied over 3.8‐fold, and inter‐annual variation in fruiting is associated with the extent of fruiting in the peak period, not variation in time of fruit set. While temperature and rainfall showed positive effects on fruiting, solar radiance in the two‐year period encompassing a given year and the previous year was the strongest predictor of fruiting. As solar irradiance was the strongest predictor of fruiting, the projected increase in rainfall associated with climate change, and coincident increase in cloud cover suggest that climate change will lead to a decrease in fruiting. ENSO in the prior 24‐month period was also significantly associated with annual ripe fruit production, and ENSO is also affected by climate change. Predicting changes in phenology demands understanding inter‐annual variation in fruit dynamics in light of potential abiotic drivers, patterns that will only emerge with long‐term data.     

Velocity of temperature and flowering time in wheat – assisting breeders to keep pace with climate change

By accelerating crop development, warming climates may result in mismatches between key sensitive growth stages and extreme climate events, with severe consequences for crop yield and food security. Using recent estimates of gene responses to vernalization and photoperiod in wheat, we modelled the flowering times of all ‘potential’ genotypes as influenced by the velocity of climate change across the Australian wheatbelt. In the period 1957–2010, seasonal increases in temperature of 0.012 °C yr^?1 were recorded and changed flowering time of a mid‐season wheat genotype by an average ?0.074 day yr^?1, with flowering ‘velocity’ of up to 0.95 km yr^?1 towards the coastal edges of the wheatbelt; this is an estimate of how quickly the given genotype would have to be ‘moved’ across the landscape to maintain its original flowering time. By 2030, these national changes are projected to accelerate by up to 3‐fold for seasonal temperature and by up to 5‐fold for flowering time between now and 2030, with average national shifts in flowering time of 0.33 and 0.41 day yr^?1 between baseline and the worst climate scenario tested for 2030 and 2050, respectively. Without new flowering alleles in commercial germplasm, the life cycle of wheat crops is predicted to shorten by 2 weeks by 2030 across the wheatbelt for the most pessimistic climate scenario. While current cultivars may be otherwise suitable for future conditions, they will flower earlier due to warmer temperatures. To allow earlier sowing to escape frost, heat and terminal drought, and to maintain current growing period of early‐sown wheat crops in the future, breeders will need to develop and/or introduce new genetic sources for later flowering, more so in the eastern part of the wheatbelt.     

Herbarium records identify sensitivity of flowering phenology of eucalypts to climate: Implications for species response to climate change

Flowering phenology is very sensitive to climate and with increasing global warming the flowering time of plants is shifting to earlier or later dates. Changes in flowering times may affect species reproductive success, associated phenological events, species synchrony, and community composition. Long‐term data on phenological events can provide key insights into the impacts of climate on phenology. For Australia, however, limited data availability restricts our ability to assess the impacts of climate change on plant phenology. To address this limitation other data sources must be explored such as the use of herbarium specimens to conduct studies on flowering phenology. This study uses herbarium specimens for investigating the flowering phenology of five dominant and commercially important Eucalyptus species of south‐eastern Australia and the consequences of climate variability and change on flowering phenology. Relative to precipitation and air humidity, mean temperature of the preceding 3 months was the most influential factor on the flowering time for all species. In response to a temperature increment of 1°C, a shift in the timing of flowering of 14.1–14.9 days was predicted for E. microcarpa and E. tricarpa while delays in flowering of 11.3–15.5 days were found for E. obliqua, E. radiata and E. polyanthemos. Eucalyptus polyanthemos exhibited the greatest sensitivity to climatic variables. The study demonstrates that herbarium data can be used to detect climatic signals on flowering phenology for species with a long flowering duration, such as eucalypts. The robust relationship identified between temperature and flowering phenology indicates that shifts in flowering times will occur under predicted climate change which may affect reproductive success, fitness, plant communities and ecosystems.     

Forecasting fine‐scale changes in the food‐web structure of coastal marine communities under climate change

Climate change is inducing deep modifications in local communities worldwide as a consequence of individualistic species range shifts. Understanding how complex interaction networks will be reorganized under climate change represents a major challenge in the fields of ecology and biogeography. However, forecasting the potential effects of climate change on local communities, and more particularly on food‐web structure, requires the consideration of highly structuring processes, such as trophic interactions. A major breakthrough is therefore expected by combining predictive models integrating habitat selection processes, the physiological limits of marine species and their trophic interactions. In this study, we forecasted the potential impacts of climate change on the local food‐web structure of the highly threatened Gulf of Gabes ecosystem located in the south of the Mediterranean Sea. We coupled the climatic envelope and habitat models to an allometric niche food web model, hence taking into account the different processes acting at regional (climate) and local scales (habitat selection and trophic interactions). Our projections under the A2 climate change scenario showed that future food webs would be composed of smaller species with fewer links, resulting in a decrease of connectance, generality, vulnerability and mean trophic level of communities and an increase of the average path length, which may have large consequences on ecosystem functioning. The unified framework presented here, by connecting food‐web ecology, biogeography and seascape ecology, allows the exploration of spatial aspects of interspecific interactions under climate change and improves our current understanding of climate change impacts on local marine food webs.     

Within‐plant variation in reproductive investment: consequences for selection on flowering time

Variation among the leaves, flowers or fruit produced by a plant is often regarded as a nuisance to the experimenter and an impediment to selection. Here, we suggest that within‐plant variation can drive selection on other plant‐level traits. We examine within‐plant variation in floral sex allocation and in fruit set and predict that such variation generates variation in male success among plants, thereby driving selection on flowering time. We tested this prediction in a simulation model estimating selection on flowering time through male fitness when floral sex allocation and/or fruit set vary directionally among flowers on plants. We parameterized the model through a quantitative literature survey of within‐plant change in sex allocation. As predicted, within‐plant variation in floral sex allocation and in fruit set probability can generate selection on flowering time through male fitness. Declining fruit set from first to last flowers on plants, as occurs in many species, selected for early flowering onset through male fitness. This result was robust to self‐incompatibility and to varying returns on male versus female investment. Selection caused by declining fruit set was strong enough to reverse the selection for late flowering that can be caused by intrafloral protandry. Our model provides testable predictions regarding selection on flowering time through male fitness. The model also establishes the intriguing possibility that within‐plant variation may influence selection on other traits, regardless of whether that variation is under selection itself.     

Spatio‐temporal variation in fitness responses to contrasting environments in Arabidopsis thaliana

The evolutionary response of organisms to global climate change is expected to be strongly conditioned by preexisting standing genetic variation. In addition, natural selection imposed by global climate change on fitness‐related traits can be heterogeneous over time. We estimated selection of life‐history traits of an entire genetic lineage of the plant Arabidopsis thaliana occurring in north‐western Iberian Peninsula that were transplanted over multiple years into two environmentally contrasting field sites in southern Spain, as southern environments are expected to move progressively northwards with climate change in the Iberian Peninsula. The results indicated that natural selection on flowering time prevailed over that on recruitment. Selection favored early flowering in six of eight experiments and late flowering in the other two. Such heterogeneity of selection for flowering time might be a powerful mechanism for maintaining genetic diversity in the long run. We also found that north‐western A. thaliana accessions from warmer environments exhibited higher fitness and higher phenotypic plasticity for flowering time in southern experimental facilities. Overall, our transplant experiments suggested that north‐western Iberian A. thaliana has the means to cope with increasingly warmer environments in the region as predicted by trends in global climate change models.     

A resurrection study reveals limited evolution of phenology in response to recent climate change across the geographic range of the scarlet monkeyflower

Premise of the studyAs global climate change alters drought regimes, rapid evolution of traits that facilitate adaptation to drought can rescue populations in decline. The evolution of phenological advancement can allow plant populations to escape drought, but evolutionary responses in phenology can vary across a species'' range due to differences in drought intensity and standing genetic variation.Methods Mimulus cardinalis, a perennial herb spanning a broad climatic gradient, recently experienced a period of record drought. Here, we used a resurrection study comparing flowering time and stem height at first flower of pre‐drought ancestors and post‐drought descendants from northern‐edge, central, and southern‐edge populations in a common environment to examine the evolution of drought escape across the latitudinal range.Key resultsContrary to the hypothesis of the evolution of advanced phenology in response to recent drought, flowering time did not advance between ancestors and descendants in any population, though storage condition and maternal effects could have impacted these results. Stem height was positively correlated with flowering time, such that plants that flowered earlier were shorter at first flower. This correlation could constrain the evolution of earlier flowering time if selection favors flowering early at a large size.ConclusionsThese findings suggest that rapid evolution of phenology will not rescue these populations from recent climate change. Future work is needed to examine the potential for the evolution of alternative drought strategies and phenotypic plasticity to buffer M. cardinalis populations from changing climate.