Optimal phenology of annual plants under grazing pressure

Plants show phenological responses to herbivory. Some enclosure experiments have demonstrated that the onset of the peak flowering season is dependent on grazing pressure. We constructed a mathematical model using Pontryargin's maximum principle to investigate changes in flowering time by examining shifts in resource allocation from vegetative to reproductive plant components. We represented a primary production of a plant individual by two types of function of vegetative part size, a linear function and a convex non-linear function. The results of a linear production model indicate that optimal phenology follows a schedule that switches from the production of vegetative parts to that of reproductive parts at a given time ('bang-bang' control). However, in a non-linear model, a singular control, wherein the plant invests in both productive and reproductive parts, may be included between obligate production and reproduction periods. We assumed that the peak of the flowering season occurs immediately following the exclusive investment in reproduction. In a linear production model, differential herbivory rates on the vegetative and reproductive parts of a plant resulted in shifts in the peak flowering time. A higher herbivory rate on the vegetative components advanced the peak, whereas it was delayed when grazing pressure focused on reproductive components of the plant. In the non-linear production model, increased grazing pressure tended to postpone the flowering peak. These results corresponded well with results of enclosure experiments, thus suggesting adaptive control of flowering time in plants.     

Nonlinear flowering responses to climate: are species approaching their limits of phenological change?

Many alpine and subalpine plant species exhibit phenological advancements in association with earlier snowmelt. While the phenology of some plant species does not advance beyond a threshold snowmelt date, the prevalence of such threshold phenological responses within plant communities is largely unknown. We therefore examined the shape of flowering phenology responses (linear versus nonlinear) to climate using two long-term datasets from plant communities in snow-dominated environments: Gothic, CO, USA (1974–2011) and Zackenberg, Greenland (1996–2011). For a total of 64 species, we determined whether a linear or nonlinear regression model best explained interannual variation in flowering phenology in response to increasing temperatures and advancing snowmelt dates. The most common nonlinear trend was for species to flower earlier as snowmelt advanced, with either no change or a slower rate of change when snowmelt was early (average 20% of cases). By contrast, some species advanced their flowering at a faster rate over the warmest temperatures relative to cooler temperatures (average 5% of cases). Thus, some species seem to be approaching their limits of phenological change in response to snowmelt but not temperature. Such phenological thresholds could either be a result of minimum springtime photoperiod cues for flowering or a slower rate of adaptive change in flowering time relative to changing climatic conditions.     

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.     

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.     

Marked reduction in demographic rates and reduced fitness advantage for early breeding is not linked to reduced thermal matching of breeding time

Warmer springs may cause animals to become mistimed if advances of spring timing, including available resources and of timing of breeding occur at different speed. We used thermal sums (cumulative sum of degree days) during spring to describe the thermal progression (timing) of spring and investigate its relationship to breeding phenology and demography of a long‐distant migrant bird, the northern wheatear (Oenanthe oenanthe L.). We first compare 20‐year trends in spring timing, breeding time, selection for breeding time, and annual demographic rates. We then explicitly test whether annual variation in selection for breeding time and demographic rates associates with the degree of phenological matching between breeding time and thermal progression of spring. Both thermal progression of spring and breeding time of wheatears advanced in time during the study period. But despite breeding on average 7 days earlier with respect to date, wheatears bred about 4 days later with respect to thermal spring progression. Over the same time period, selection for breeding time changed from distinct within‐season advantage of breeding early to no or very weak advantage. Furthermore, demographic rates (nest success, fledgling production, recruitment, adult survival) and nestling weight declined markedly by 16%–79%. Those temporal trends suggest that a reduced degree of phenological matching may affect within‐season fitness advantage of early breeding and population demographic rates. In contrast, when we investigate links based on annual variation, we find no significant relationship between either demographic rates or fitness advantage of early breeding with annual variation in the degree of phenological matching. Our results show that corresponding temporal trends in phenological matching, selection for breeding time and demographic rates are inconclusive evidence for demographic effects of changed phenological matching. Instead, we suggest that the trends in selection for breeding time and demographic rates are due to a general deterioration of the breeding environment.     

物候变化对北美温带落叶阔叶林生态系统生产力的影响

数据源、时间范围、空间尺度等的差异导致许多物候变化对陆地生态系统碳收支影响的研究缺少可比性。该文基于4级碳通量填充数据, 采用相对阈值方法提取了两个北美典型温带阔叶林站Harvard Forest (HF)和University of Michigan Biological Station (UMBS)共20年的物候参数(返青期、枯黄期和生长季长度), 并研究了物候变化对生态系统生产力的影响。结果表明: 1)生长季长度的延长对年累积总初级生产力(GPP)有显著贡献, 但由于呼吸作用(RE)的干扰, 生长季长度变化对年净生态系统生产力(NEP)的影响并不显著; 2)返青期的提前对上半年生态系统总初级生产力的贡献最为显著, 二者的相关系数分别为0.76 (HF)和0.93 (UMBS); 3)枯黄期的延迟对生产力的影响并不显著; 4)随着春季返青期的提前或秋季枯黄期的延迟, 上、下半年GPP和RE的累积量虽均有增加趋势, 但由于各自增加的幅度不确定, 导致年NEP与二者的响应关系复杂。     

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.     

Traits and climate are associated with first flowering day in herbaceous species along elevational gradients

Phenological responses to changing temperatures are known as “fingerprints of climate change,” yet these reactions are highly species specific. To assess whether different plant characteristics are related to these species‐specific responses in flowering phenology, we observed the first flowering day (FFD) of ten herbaceous species along two elevational gradients, representing temperature gradients. On the same populations, we measured traits being associated with (1) plant performance (specific leaf area), (2) leaf biochemistry (leaf C, N, P, K, and Mg content), and (3) water‐use efficiency (stomatal pore area index and stable carbon isotopes concentration). We found that as elevation increased, FFD was delayed for all species with a highly species‐specific rate. Populations at higher elevations needed less temperature accumulation to start flowering than populations of the same species at lower elevations. Surprisingly, traits explained a higher proportion of variance in the phenological data than elevation. Earlier flowering was associated with higher water‐use efficiency, higher leaf C, and lower leaf P content. In addition to that, the intensity of shifts in FFD was related to leaf N and K. These results propose that traits have a high potential in explaining phenological variations, which even surpassed the effect of temperature changes in our study. Therefore, they have a high potential to be included in future analyses studying the effects of climate change and will help to improve predictions of vegetation changes.     

Artificial selection shifts flowering phenology and other correlated traits in an autotetraploid herb

There is mounting evidence that plants are responding to anthropogenic climate change with shifts in flowering phenologies. We conducted a three-generation artificial selection experiment on flowering time in Campanulastrum americanum, an autotetraploid herb, to determine the potential for adaptive evolution of this trait as well as possible costs associated with enhanced or delayed flowering. Divergent selection for earlier and later flowering resulted in a 25-day difference in flowering time. Experiment-wide heritability was 0.31 and 0.23 for the initiation of flowering in early and late lines, respectively. Selection for earlier flowering resulted in significant correlated responses in other traits including smaller size, fewer branches, smaller floral displays, longer fruit maturation times, fewer seeds per fruit and slower seed germination. Results suggest that although flowering time shows the potential to adapt to a changing climate, phenological shifts may be associated with reduced plant fitness possibly hindering evolutionary change.     

Selection and plasticity both account for interannual variation in life‐history phenology in an annual prairie legume

As the environment changes, so too must plant communities and populations if they are to persist. Life‐history transitions and their timing are often the traits that are most responsive to changing environmental conditions. To compare the contributions of plasticity and natural selective response to variation in germination and flowering phenology, we performed a quantitative genetic study of phenotypic selection on Chamaecrista fasciculata (Fabaceae) across two consecutive years in a restored tallgrass prairie. The earliest dates of germination and flowering were recorded for two parental cohorts and one progeny cohort in an experimental garden. Environmental differences between years were the largest contributors to phenological variation in this population. In addition, there was substantial heritability for flowering time and statistically significant selection for advancement of flowering. Comparison between a progeny cohort and its preselection parental cohort indicated a change in mean flowering time consistent with the direction of selection. Selection on germination time was weaker than that on flowering time, while environmental effects on germination time were stronger. The response to selection on flowering time was detectable when accounting for the effect of the environment on phenotypic differences, highlighting the importance of controlling for year‐to‐year environmental variation in quantitative genetic studies.     

Temperature alone does not explain phenological variation of diverse temperate plants under experimental warming

Anthropogenic climate change has altered temperate forest phenology, but how these trends will play out in the future is controversial. We measured the effect of experimental warming of 0.6–5.0 °C on the phenology of a diverse suite of 11 plant species in the deciduous forest understory (Duke Forest, North Carolina, USA) in a relatively warm year (2011) and a colder year (2013). Our primary goal was to dissect how temperature affects timing of spring budburst, flowering, and autumn leaf coloring for functional groups with different growth habits, phenological niches, and xylem anatomy. Warming advanced budburst of six deciduous woody species by 5–15 days and delayed leaf coloring by 18–21 days, resulting in an extension of the growing season by as much as 20–29 days. Spring temperature accumulation was strongly correlated with budburst date, but temperature alone cannot explain the diverse budburst responses observed among plant functional types. Ring‐porous trees showed a consistent temperature response pattern across years, suggesting these species are sensitive to photoperiod. Conversely, diffuse‐porous species responded differently between years, suggesting winter chilling may be more important in regulating budburst. Budburst of the ring‐porous Quercus alba responded nonlinearly to warming, suggesting evolutionary constraints may limit changes in phenology, and therefore productivity, in the future. Warming caused a divergence in flowering times among species in the forest community, resulting in a longer flowering season by 10‐16 days. Temperature was a good predictor of flowering for only four of the seven species studied here. Observations of interannual temperature variability overpredicted flowering responses in spring‐blooming species, relative to our warming experiment, and did not consistently predict even the direction of flowering shifts. Experiments that push temperatures beyond historic variation are indispensable for improving predictions of future changes in phenology.     

Beyond gradual warming: extreme weather events alter flower phenology of European grassland and heath species

Shifts in the phenology of plant and animal species or in the migratory arrival of birds are seen as ‘fingerprints’ of global warming. However, even if such responses have been documented in large continent‐wide datasets of the northern hemisphere, all studies to date correlate the phenological pattern of various taxa with gradual climatic trends. Here, we report a previously unobserved phenomenon: severe drought and heavy rain events caused phenological shifts in plants of the same magnitude as one decade of gradual warming. We present data from two vegetation periods in an experimental setting containing the first evidence of shifted phenological response of 10 grassland and heath species to simulated 100‐year extreme weather events in Central Europe. Averaged over all species, 32 days of drought significantly advanced the mid‐flowering date by 4 days. The flowering length was significantly extended by 4 days. Heavy rainfall (170 mm over 14 days) had no significant effect on the mid‐flowering date. However, heavy rainfall reduced the flowering length by several days. Observed shifts were species‐specific, (e.g. drought advanced the mid‐flowering date for Holcus lanatus by 1.5 days and delayed the mid‐flowering date for Calluna vulgaris by 5.7 days, heavy rain advanced mid‐flowering date of Lotus corniculatus by 26.6 days and shortened the flowering length of the same species by 36.9 days). Interestingly, the phenological response of individual species was modified by community composition. For example, the mid‐flowering date of C. vulgaris was delayed after drought by 9.3 days in communities composed of grasses and dwarf shrubs compared with communities composed of dwarf shrubs only. This indicates that responses to extreme events are context specific. Additionally, the phenological response of experimental communities to extreme weather events can be modified by the functional diversity of a stand. Future studies on phenological response patterns related to climate change would profit from explicitly addressing the role of extreme weather events.     

Phenology and phenotypic natural selection on the flowering time of a deceit-pollinated tropical orchid, Myrmecophila christinae

BACKGROUND AND AIMS: Flowering phenology is described and the effect of flowering time on pollination success is evaluated in the deceit-pollinated tropical orchid, Myrmecophila christinae. It was expected that, due to this species' deceit pollination strategy and low observed pollinator visit rate, there would be a higher probability of natural selection events favouring individuals flowering away from the population flowering peak. METHODS: The study covers two consecutive years and four populations of M. christinae located along the north coast of the Yucatán Peninsula. For phenological and pollination success data, a total of 110 individuals were monitored weekly in 1998, and 83 individuals in 1999, during all the flowering and fruiting season. KEY RESULTS: The results showed significant differences in the probability of donating and receiving pollen throughout the flowering season. The probability of receiving or donating pollen increased the further an individual flowering was from the flowering peak. Regression analysis showed directional and disruptive phenotypic natural selection gradients, suggesting the presence of selection events unfavourable to flowering during flowering peak, for both male success (pollen removal) and female success (fruit production). However, the intensity and significance of the natural selection events varied between populations from year to year. The variation between seasons and populations was apparently due to variations in the density of reproductive individuals in each population and each season. CONCLUSIONS: As in other deceit-pollinated orchids, natural selection in M. christinae favours individuals flowering early or late in relation to population peak flowering. However, results also suggested a fluctuating regime of selective events act on flowering time of M. christinae.     

Seasonal Patterns of Flowering and Fruiting in a Dry Tropical Forest in Jamaica1

Tropical dry forests occupy more area and are more endangered than rainforests, yet their regeneration ecology has received less study and is consequently poorly understood. We recorded the flowering and fruiting phenology of a tropical dry forest in Jamaica over a period of 26 mo within ten 15 × 15‐m plots. Community‐wide recruitment reached a maximum in the wet season, whereas no recruitment occurred during the dry season. We observed a unimodal peak in rainfall and fruit production, and the periodicity and intensity of seed production were significantly correlated with rainfall seasonality (the optimal time for germination). Flowering at the community and system levels lagged behind a significant increase and subsequent decrease in rainfall by 7 and 3 mo, respectively, indicating that the dominant factor controlling flowering periodicity is the passage of the major (4‐mo long) rainy season and changes in soil moisture conditions. Fruiting lagged behind flowering by 2 mo and a significant increase in fruiting occurred 2 mo prior to a significant increase in rainfall. At the population level, a correspondence analysis identified a major dichotomy in the patterns of flowering and fruiting between species and indicated two broad species groups based on their time of peak fruiting and the number of times they were in fruit. These were either individuals which were usually in peak fruit 1–2 mo prior to the start of the major rainy season or those that were in fruit more or less continuously throughout the year with no peak fruiting time. This study supports the view that seasonal variation in rainfall and hence soil water availability constitutes both the proximate and the ultimate cause of flowering periodicity in tropical dry forests.     

Impact of climate change on plant phenology in Mediterranean ecosystems

Plant phenology is strongly controlled by climate and has consequently become one of the most reliable bioindicators of ongoing climate change. We used a dataset of more than 200 000 records for six phenological events of 29 perennial plant species monitored from 1943 to 2003 for a comprehensive assessment of plant phenological responses to climate change in the Mediterranean region. Temperature, precipitation and North Atlantic Oscillation (NAO) were studied together during a complete annual cycle before phenological events to determine their relative importance and potential seasonal carry‐over effects. Warm and dry springs under a positive phase of NAO advance flowering, leaf unfolding and fruiting dates and lengthen the growing season. Spatial variability of dates (range among sites) was also reduced during warm and dry years, especially for spring events. Climate during previous weeks to phenophases occurrence had the greatest impact on plants, although all events were also affected by climate conditions several months before. Immediate along with delayed climate effects suggest dual triggers in plant phenology. Climatic models accounted for more than 80% of variability in flowering and leaf unfolding dates, and in length of the growing season, but for lower proportions in fruiting and leaf falling. Most part of year‐to‐year changes in dates was accounted for temperature, while precipitation and NAO accounted for <10% of dates' variability. In the case of flowering, insect‐pollinated species were better modelled by climate than wind‐pollinated species. Differences in temporal responses of plant phenology to recent climate change are due to differences in the sensitivity to climate among events and species. Spring events are changing more than autumn events as they are more sensitive to climate and are also undergoing the greatest alterations of climate relative to other seasons. In conclusion, climate change has shifted plant phenology in the Mediterranean region.     

Maintenance of temporal synchrony between syrphid flies and floral resources despite differential phenological responses to climate

Variation in species’ responses to abiotic phenological cues under climate change may cause changes in temporal overlap among interacting taxa, with potential demographic consequences. Here, we examine associations between the abiotic environment and plant–pollinator phenological synchrony using a long‐term syrphid fly–flowering phenology dataset (1992–2011). Degree‐days above freezing, precipitation, and timing of snow melt were investigated as predictors of phenology. Syrphids generally emerge after flowering onset and end their activity before the end of flowering. Neither flowering nor syrphid phenology has changed significantly over our 20‐year record, consistent with a lack of directional change in climate variables over the same time frame. Instead we document interannual variability in the abiotic environment and phenology. Timing of snow melt was the best predictor of flowering onset and syrphid emergence. Snow melt and degree‐days were the best predictors of the end of flowering, whereas degree‐days and precipitation best predicted the end of the syrphid period. Flowering advanced at a faster rate than syrphids in response to both advancing snow melt and increasing temperature. Different rates of phenological advancements resulted in more days of temporal overlap between the flower–syrphid community in years of early snow melt because of extended activity periods. Phenological synchrony at the community level is therefore likely to be maintained for some time, even under advancing snow melt conditions that are evident over longer term records at our site. These results show that interacting taxa may respond to different phenological cues and to the same cues at different rates but still maintain phenological synchrony over a range of abiotic conditions. However, our results also indicate that some individual plant species may overlap with the syrphid community for fewer days under continued climate change. This highlights the role of interannual variation in these flower–syrphid interactions and shows that species‐level responses can differ from community‐level responses in nonintuitive ways.     

Spatial and habitat variation in aphid,butterfly, moth and bird phenologies over the last half century

Global warming has advanced the timing of biological events, potentially leading to disruption across trophic levels. The potential importance of phenological change as a driver of population trends has been suggested. To fully understand the possible impacts, there is a need to quantify the scale of these changes spatially and according to habitat type. We studied the relationship between phenological trends, space and habitat type between 1965 and 2012 using an extensive UK dataset comprising 269 aphid, bird, butterfly and moth species. We modelled phenologies using generalized additive mixed models that included covariates for geographical (latitude, longitude, altitude), temporal (year, season) and habitat terms (woodland, scrub, grassland). Model selection showed that a baseline model with geographical and temporal components explained the variation in phenologies better than either a model in which space and time interacted or a habitat model without spatial terms. This baseline model showed strongly that phenologies shifted progressively earlier over time, that increasing altitude produced later phenologies and that a strong spatial component determined phenological timings, particularly latitude. The seasonal timing of a phenological event, in terms of whether it fell in the first or second half of the year, did not result in substantially different trends for butterflies. For moths, early season phenologies advanced more rapidly than those recorded later. Whilst temporal trends across all habitats resulted in earlier phenologies over time, agricultural habitats produced significantly later phenologies than most other habitats studied, probably because of nonclimatic drivers. A model with a significant habitat‐time interaction was the best‐fitting model for birds, moths and butterflies, emphasizing that the rates of phenological advance also differ among habitats for these groups. Our results suggest the presence of strong spatial gradients in mean seasonal timing and nonlinear trends towards earlier seasonal timing that varies in form and rate among habitat types.     

Increased variance in temperature and lag effects alter phenological responses to rapid warming in a subarctic plant community

Summer temperature on the Cape Churchill Peninsula (Manitoba, Canada) has increased rapidly over the past 75 years, and flowering phenology of the plant community is advanced in years with warmer temperatures (higher cumulative growing degree days). Despite this, there has been no overall shift in flowering phenology over this period. However, climate change has also resulted in increased interannual variation in temperature; if relationships between phenology and temperature are not linear, an increase in temperature variance may interact with an increase in the mean to alter how community phenology changes over time. In our system, the relationship between phenology and temperature was log‐linear, resulting in a steeper slope at the cold end of the temperature spectrum than at the warm end. Because below‐average temperatures had a greater impact on phenology than above‐average temperatures, the long‐term advance in phenology was reduced. In addition, flowering phenology in a given year was delayed if summer temperatures were high the previous year or 2 years earlier (lag effects), further reducing the expected advance over time. Phenology of early‐flowering plants was negatively affected only by temperatures in the previous year, and that of late‐flowering plants primarily by temperatures 2 years earlier. Subarctic plants develop leaf primordia one or more years prior to flowering (preformation); these results suggest that temperature affects the development of flower primordia during this preformation period. Together, increased variance in temperature and lag effects interacted with a changing mean to reduce the expected phenological advance by 94%, a magnitude large enough to account for our inability to detect a significant advance over time. We conclude that changes in temperature variability and lag effects can alter trends in plant responses to a warming climate and that predictions for changes in plant phenology under future warming scenarios should incorporate such effects.     

Selection on flowering time in a life‐cycle context

The main way in which plants can exert control over their local environment is by the timing of different events within their life cycles. Regarding timing of flowering as an integrated part of both the annual cycle and of the whole life cycle, rather than as an isolated event, has important implications for how we assess selection on timing of reproduction and interpret existing phenological patterns in perennial plants. I argue that: 1) we have little unequivocal evidence of pollinator‐mediated selection on flowering time, but perhaps more evidence of antagonist‐mediated selection; 2) much of selection on flowering time might occur before flowers have developed and after reproduction; 3) vital rates of non‐flowering individuals can influence the strength and direction of selection on flowering time, and 4) differences in the direction of selection on flowering date between years might well correspond to consistent selection on the mechanisms determining flowering time. Overall, a life cycle perspective on timing of flowering is likely to facilitate the identification of selective agents and the understanding of the complex mechanisms underlying spatial and temporal variation in selection as well as to enable more accurate predictions of responses to environmental change.     

Is timing of reproduction according to temperature sums an optimal strategy?

Temperature sums are widely used to predict the seasonal timing of yearly recurring biological events, such as flowering, budburst, and hatching. We use a classic energy allocation model for annual plants to compare a strategy for reproductive timing that follows a temperature sum rule (TSR) with a strategy that follows an optimal control rule (OCR) maximizing reproductive output. We show that the OCR corresponds to a certain TSR regardless of how temperature is distributed over the growing season as long as the total temperature sum over the whole growing season is constant between years. We discuss such scenarios, thus outlining under which type of variable growth conditions TSR maximizes reproductive output and should be favored by natural selection. By providing an ultimate explanation for a well‐documented empirical pattern this finding enhances the credibility of temperature sums as predictors of the timing of biological events. However, TSR and OCR respond in opposite directions when the total yearly temperature sum changes between years, representing, for example, variation in the length of the growing season. Our findings have implications for predicting optimal responses of organisms to climatic changes and suggest under which conditions natural selection should favor photoperiod versus temperature control.