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.     

Factors affecting fruit and seed production of Paeonia ostii “Feng Dan”, an economically important oil tree

Many factors may affect reproduction of animal-pollinated species. In this study, the effects of pollen limitation, attractive traits (flower number, plant height and flower width) and flowering phenological traits (flowering onset, duration and synchrony) on female reproduction, as well as the patterns of variation in fruit and seed production within plants, were investigated in Paeonia ostii “Feng Dan” over two flowering seasons (2018 and 2019). Fruit set was very high (90%), and pollen supplementation did not increase fruit and seed production in either year, indicating no pollen limitation. Fruit set, ovule number per fruit and mean individual seed weight per fruit were not affected by any of the six attractive and phenological traits in either year, whereas seed number per fruit was related to the three attractive traits in one or both years. Seed number per plant was positively affected by the three attractive traits and best explained by flower number in both years, but the effect of each of the three phenological traits on seed number per plant differed between years. Within plants, the fruit set, ovule number, seed set and seed number per fruit declined from early- to late-opening flowers, presumably because of resource preemption, but the mean individual seed weight did not vary across the flowering sequence. Our study shows that attractive traits of Paeonia ostii “Feng Dan” are more important than flowering phenological traits in the prediction of total seed production per plant.     

Cattle change plant reproductive phenology,promoting community changes in a post-fire Nothofagus forest in northern Patagonia,Argentina

Aims Variations in rates and length of flowering and fruiting not only affect the reproduction of a given plant species but also the behavior and reproduction of associated taxa. Flowering and fruiting variations may be influenced by herbivory, especially by large mammals. The aim of this study was to determine the effects of cattle browsing on the reproductive phenology of understory species in a subalpine post-fire Nothofagus forest in Patagonia.Methods The effects of herbivory on plant reproductive phenology were studied in a set of experimental exclosures (fenced plots) installed since 2001 in a post-fire N. pumilio forest, located in Nahuel Huapi National Park (NHNP), Argentina. We monitored the beginning and duration of each reproductive phenological stage: floral bud, open flower, immature fruit and mature fruit. We also counted the number of flowers, fruits, seeds and viable seeds of the dominant plants to assess whether browsing modifies temporal patterns of the flowering and fruiting periods.Important findings Cattle reduced the total number of species flowering and fruiting and changed the reproductive phenology of some species. We found that palatable species seem to be negatively affected by browsing in terms of reduced fitness due to changes in flowering and fruiting periods. In contrast, cattle benefitted the reproduction of non-palatable species and could promote the invasion of shade-intolerant exotic forbs such as Cirsium vulgare. The effects of livestock reported in this study are important to understanding how browsing could alter native species establishment and possibly alter successional trajectories during recolonization after fire.     

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.     

Reproductive phenology in three Genisteae (Fabaceae) shrub species of the W Mediterranean Region

We studied the flower duration, the phenology of flowering and fruiting, and flower, fruit, and seed production in three Genisteae shrub species of the W Mediterranean Region: Cytisus multiflorus, C. striatus , and Retama sphaerocarpa . Flower duration was negatively correlated with temperature, and in the case of C. striatus it was also influenced by pollination. In Cytisus multiflorus , which flowers during winter in the population studied, two floral morphs were recognized differentiated by flower size, phenological pattern, and production of the reproductive organs: morph LF (large flowers) and morph SF (small flowers). The former is earlier in phenology but its reproductive success is less than the second since it not only produces significantly fewer flowers/plant, but also a smaller crop of fruit and seeds. The flowering of the LF morph lasted some three months and of the SF morph 11 weeks. The winter-spring flowering C. striatus , with a four month duration of flowering time, is sympatric with C. multiflorus and their flowering periods overlap, but the former is more successful reproductively. Retama sphaerocarpa is clearly spring flowering, with an extremely short duration of flowering (c. 6.5 weeks), but, unlike the other two species which disperse their seeds in the same season in which they are produced, it staggers the dispersal of its diaspores over more than one year. Despite the great flower per plant production, especially in Retama sphaerocarpa , and of the number of fruit initiated, the final crop of fruit and seed/plant is low in the three species (C. multiflorus : fruit 8.40% and seeds 1.96%; C. striatus : fruit 13.09% and seeds 4.12%; Retama sphaerocarpa : fruit 5.65% and seeds 1.33%).     

The phenology of growth and reproduction in plants

The study of phenological aspects of plants involves the observation, recording and interpretation of the timing of their life history events. This review considers the phenology of leafing, flowering and fruit production in a range of species and communities. The selective forces (both abiotic and biotic) that influence the timing of these events are discussed. Within the limits imposed by phylogenetic constraints, the phenological patterns (timing, frequency, duration, degree of synchrony, etc.) of each phase are probably the result of a compromise between a variety of selective pressures, such as seasonal climatic changes, resource availability, and the presence of pollinators, predators and seed dispersers. Many studies on flowering times stress the role of interactions between plant species which share pollinators or predators. The timing of fruiting plays a key role in controlling the abundance and variety of obligate frugivores in many tropical communities. The importance of long-term recording is stressed, particularly in species which fruit irregularly. An understanding of the phenology of plants is crucial to the understanding of community function and diversity.     

Phylogenetic conservatism and climate factors shape flowering phenology in alpine meadows

The study of phylogenetic conservatism in alpine plant phenology is critical for predicting climate change impacts; currently we have a poor understanding of how phylogeny and climate factors interactively influence plant phenology. Therefore, we explored the influence of phylogeny and climate factors on flowering phenology in alpine meadows. For two different types of alpine plant communities, we recorded phenological data, including flowering peak, first flower budding, first flowering, first fruiting and the flowering end for 62 species over the course of 5 years (2008–2012). From sequences in two plastid regions, we constructed phylogenetic trees. We used Blomberg’s K and Pagel’s lambda to assess the phylogenetic signal in phenological traits and species’ phenological responses to climate factors. We found a significant phylogenetic signal in the date of all reproductive phenological events and in species’ phenological responses to weekly day length and temperature. The number of species in flower was strongly associated with the weekly day lengths and followed by the weekly temperature prior to phenological activity. Based on phylogenetic eigenvector regression (PVR) analysis, we found a highly shared influence of phylogeny and climate factors on alpine species flowering phenology. Our results suggest the phylogenetic conservatism in both flowering and fruiting phenology may depend on the similarity of responses to external environmental cues among close relatives.     

Reproductive output in Mediterranean shrubs under climate change experimentally induced by drought and warming

The effects of climate change on plant reproductive performance affects the sequence of different plant reproductive stages from flowering to seed production and viability, as well as the network of relationships between them. These effects are expected to respond to different components of climate change, such as temperature and water availability, and may be sensitive to differences in species phenology.We used long-term experimental drought and warming treatments to study the effect of climate change on flower production, fruit and seed-set, seed size and seed germination rate (proportion of germinating seeds) in three Mediterranean shrubs coexisting in a coastal shrubland.Larger plants produced significantly more flowers in all three species, and higher fruit-set in Dorycnium pentaphyllum. Flower production was reduced in drought and warming treatments in the spring-flowering species D. pentaphyllum and Helianthemum syriacum, but not in the autumn–winter species Erica multiflora, which increased flowering in the warming treatment. However, the drought treatment eventually resulted in a decreased seed-set in E. multiflora. Structural equation modelling revealed strong correlations between the sequential reproductive stages. Specifically, flower density in inflorescences determined seed-set in H. syriacum, and seed size and germination rate in E. multiflora. Nevertheless, the relevance of relationships between reproductive traits changed between climatic treatments: in D. pentaphyllum a direct relationship between plant size and seed size only arised in the drought treatment, while in H. syriacum climate treatments resulted in a stronger relationship between the number of flowers and seed-set.This experimental study shows the ability of changing climatic variables to determine the reproductive sequential process of woody species. We show that several parameters of the reproductive performance of some Mediterranean species are affected by drought and warming treatments simulating climate change, highlighting the importance of changes in both water availability and temperature, and the sequential relationship between reproductive stages. Phenological patterns also contribute to species’ differential responses to climatic change, due to the relationship of these patterns with resource availability, environmental conditions and plant–pollinator interactions.     

A Range-Expanding Shrub Species Alters Plant Phenological Response to Experimental Warming

Shifts in plant species phenology (the timing of life-history events such as flowering) have been observed worldwide in concert with rising global temperatures. While most species display earlier phenology with warming, there is large variation among, and even within, species in phenological sensitivity to rising temperatures. Other indirect effects of climate change, such as shifting species composition and altered species interactions, may also be contributing to shifting plant phenology. Here, we describe how experimental warming and the presence of a range-expanding species, sagebrush (Artemisia rothrockii), interact to influence the flowering phenology (day of first and peak flowering) and production (number of flowers) of an alpine cushion plant, Trifolium andersonii, in California’s White Mountains. Both first flowering and peak flowering were strongly accelerated by warming, but not when sagebrush was present. Warming significantly increased flower production of T. andersonii, but less so in the presence of sagebrush. A shading treatment delayed phenology and lowered flower production, suggesting that shading may be the mechanism by which sagebrush presence delayed flowering of the understory species. This study demonstrates that species interactions can modify phenological responses to climate change, and suggests that indirect effects of rising temperatures arising from shifting species ranges and altered species interactions may even exceed the direct effects of rising temperatures on phenology.     

Vulnerability and determinants of reproductive success in the narrow endemic Antirrhinum microphyllum (Scrophulariaceae)

The breeding system and flowering phenology of the narrow endemic Antirrhinum microphyllum (Scrophulariaceae) were studied in order to assess the main factors affecting female reproductive success and to identify existing or potential threats to the viability of its populations. Hand-pollination experiments showed that A. microphyllum is an allogamous self-incompatible species. In both populations studied, the flowering season was 4 mo long and mean flowering duration per plant was about 1 mo. Peak flower production took place between mid-April and mid-May and overlapped with the period of activity of the main pollinator, Rhodanthidium sticticum (Megachilidae). Estimated mean number of seeds produced per plant was 9391, showing that population viability is not presently limited by seed output. The study of the direct and indirect effects of plant size, phenological traits (first flowering date, flowering duration, and flowering synchrony), and distance to neighbor plants on reproductive success was performed using structural equation modeling (SEM). In both populations, number of flowers and plant size were the main factors that determined the total number of fruits produced by a plant. First flowering date and flowering synchrony also affected fruit production. Multisample comparison of path coefficients for the two populations rejected the possibility that reproductive patterns could be described by one single model.     

Plant reproductive phenology over four years including an episode of general flowering in a lowland dipterocarp forest,Sarawak, Malaysia

The first systematic observation of a general flowering, a phenomenon unique to lowland mixed-dipterocarp forests in Southeast Asia, is presented. During general flowering, which occurs at irregular intervals of 3–10 yr, nearly all dipterocarp species together with species of other families come heavily into flower. We monitored reproductive phenology of 576 individual plants representing 305 species in 56 families in Sarawak, Malaysia. Observations continued for 53 mo from August 1992 and covered one episode of a general flowering cycle. Among 527 effective reproductive events during 43 mo, 57% were concentrated in the general flowering period (GFP) of 10 mo in 1996. We classified 257 species into flowering types based on timing and frequency of flowering. The most abundant type was “general flowering” (35%), which flowered only during GFP. The others were “supra-annual” (19%), “annual” (13%), and “sub-annual” (5%) types. General flowering type and temporal aggregation in reproductive events were commonly found among species in various categories of taxonomic groups, life forms, pollination systems, and fruit types. Possible causes for general flowering, such as promotion of pollination brought about by interspecific synchronization and paucity of climatic cues suitable for flowering trigger, are proposed, in addition to the predator satiation hypothesis of Janzen (1974) .     

Global warming reduces plant reproductive output for temperate multi-inflorescence species on the Tibetan plateau

? Temperature is projected to increase more during the winter than during the summer in cold regions. The effects of winter warming on reproductive effort have not been examined for temperate plant species. ? Here, we report the results of experimentally induced seasonal winter warming (0.4 and 2.4°C increases in growing and nongrowing seasons, respectively, using warmed and ambient open-top chambers in a Tibetan Plateau alpine meadow) for nine indeterminate-growing species producing multiple (single-flowered or multi-flowered) inflorescences and three determinate-growing species producing single inflorescences after a 3-yr period of warming. ? Warming reduced significantly flower number and seed production per plant for all nine multi-inflorescence species, but not for the three single-inflorescence species. Warming had an insignificant effect on the fruit to flower number ratio, seed size and seed number per fruit among species. The reduction in seed production was largely attributable to the decline in flower number per plant. The flowering onset time was unaffected for nine of the 12 species. Therefore, the decline in flower production and seed production in response to winter warming probably reflects a physiological response (e.g. metabolic changes associated with flower production). ? Collectively, the data indicate that global warming may reduce flower and seed production for temperate herbaceous species and will probably have a differential effect on single- vs multi-inflorescence species.     

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.     

Phenological responses to nitrogen and water addition are linked to plant growth patterns in a desert herbaceous community

Increases in nitrogen (N) deposition and variation in precipitation have been occurring in temperate deserts; however, little information is available regarding plant phenological responses to environmental cues and their relationships with plant growth pattern in desert ecosystems. In this study, plant phenology and growth of six annuals in response to N and water addition were monitored throughout two consecutive growing seasons in 2011 and 2012 in a temperate desert in northwestern China. The effects of N and water addition on reproductive phenology differed among plant species. N and water addition consistently advanced the flowering onset time and fruiting time of four spring ephemerals; however, their effects on two spring‐summer annuals were inconsistent, with advances being noted in one species and delays in another. N and water addition alone increased plant height, relative growth rate, leaf number, flower number, and individual biomass, while their combinative effects on plant growth and reproductive phenology were dependent on species. Multiple regression analysis showed that flowering onset time was negatively correlated with relative growth rate of two species, and negatively correlated with maximum plant height of the other four species. Our study demonstrates that phenological responses to increasing precipitation and N deposition varied in annuals with different life histories, whereby the effects of climate change on plant growth rate were related to reproductive phenology. Desert annuals that were able to accelerate growth rate under increasing soil resource availability tended to advance their flowering onset time to escape drought later in the growing season. This study promotes our understanding of the responses of temperate desert annuals to increasing precipitation and N deposition in this desert.     

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.     

Flower morphology, phenology and visitor patterns in an alpine community on Mt Olympos, Greece

Alpine vegetation, restricted to the top of high mountains, is among the vegetation types most endangered by global warming, currently predicted to raise temperatures from 1.1 to 6.4 °C, by the end of the century. Nevertheless, background information allowing evaluation of impacts is rather scarce for some geographic zones. Our study of an alpine community on the Plateau of Muses (2600–2750 m a.s.l.) of Mt Olympos, the highest mountain of Greece, conducted in 1993–1994, can provide such background information for the Mediterranean region. We studied features relating to phenology of flowering, floral morphology, distribution and abundance, and flower visitors of plant species that exhibit a biotic pollination syndrome. We identified dominant patterns and we further (i) explored the relative contribution of the plant features and abiotic factors studied in explaining the activity patterns of flower visitors, (ii) examined if flower and visitor traits of the alpine community match each other according to the classical pollination syndromes, and (iii) investigated whether the responses of individual plant species to the yearly climatic variability result into phenological patterns that characterize the whole community. The common strategy of the alpine community was for early flowering and long flower life span; consistently early flowering species were twice as many as late flowering ones, whereas floral longevity (estimated for 36 species) averaged 5.2 days. Duration of flowering (estimated for 57 species) averaged 18.2 days. Climatic variability affected onset of flowering; all late flowering species delayed their flowering during the year characterized by a humid and cold summer. Duration of flowering and floral longevity did not change in a consistent way. Hymenoptera (Aculeates) were the dominant flower visitors. They accounted for 43.3% of the visits recorded, with bumblebees making a little less than half. Diptera followed making 37.5% of the visits (most made by syrphid flies). There was a mismatch between flower-morphology and flower visitor traits; the alpine community had predominantly non-specialized, pale-colour flowers, which are traits assumed to correspond to Diptera dominance and absence of social bees. Visitation was influenced by flower abundance and duration of flowering; proportionately more Diptera, and proportionately less Hymenoptera visited species with short flowering periods and few flowers present in the field. In a number of cases, the phenological and flower visitor patterns of the community of Mt Olympos deviated from those observed in other alpine environments suggesting a mediterranean influence even at high altitudes.     

Flower and Fruit Availability along a Forest Restoration Gradient

Frugivores and pollinators are two functional groups of animals that help ensure gene flow of plants among sites in landscapes under restoration and to accelerate restoration processes. Resource availability is postulated to be a key factor to structure animal communities using restoration sites, but it remains poorly studied. We expected that diverse forests with many plant growth forms that have less‐seasonal phenological patterns will provide more resources for animals than forests with fewer plant growth forms and strongly seasonal phenological patterns. We studied forests where original plantings included high tree species diversity. We studied resource provision (richness and abundance of flowers and fruits) of all plant growth forms, in three restoration sites of different ages compared to a reference forest, investigating whether plant phenology changes with restoration process. We recorded phenological data for reproductive plant individuals (351 species) with monthly sampling over 2 years, and found that flower and fruit production have been recovered after one decade of restoration, indicating resource provision for fauna. Our data suggest that a wide range of plant growth forms provides resource complementarities to those of planted tree species. Different flower phenologies between trees and non‐trees seem to be more evident in a forest with high non‐tree species diversity. We recommend examples of ideal species for planting, both at the time of initial planting and post‐planting during enrichment. These management actions can minimize shortage and periods of resource scarcity for frugivorous and nectarivorous fauna, increasing probability of restoring ecological processes and sustainability in restoration sites.     

REPRODUCTIVE PHENOLOGIES IN LOBELIA INFLATA (LOBELIACEAE) AND THEIR ENVIRONMENTAL CONTROL

Flowering and fruiting phenologies of individual plants and flowers of Lobelia inflata, a North American summer annual, were studied in the field and greenhouse to determine whether onset of flowering and fruit maturation were correlated, and the degree to which these reproductive phenologies were influenced by the environment. Within each of two field populations, larger plants flowered earlier and produced more flowers than smaller plants. Onset of flowering was positively correlated with onset of fruit maturation but not perfectly so. Two factors decreased the intensity of this correlation. First, at the flower level, the earlier a flower bloomed, the longer the resulting fruit took to develop. Second, fruit development times varied significantly among individual plants. In the greenhouse, individuals watered more frequently attained greater size and flowered earlier than individuals watered less frequently. Nutrient additions did not affect plant size or onset of flowering. These results indicate that for the summer annual Lobelia inflata, reproductive phenologies are phenotypically correlated, and that timing of reproduction is resource and size dependent, as it is for other monocarpic plant species.     

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.