The effect of light environment, leaf area, and stored carbohydrates on inflorescence production by a rain forest understory palm

Variation in flowering by long-lived plants may be correlated with current resource availability. If, however, there are trade-offs between current and future reproduction, or between reproduction and storage or growth, then understanding variation requires a whole-plant, longer-term perspective. Inflorescence production by Calyptrogyne ghiesbreghtiana Linden ex. H. Wendl., an understory palm, was studied over 3 years. Annual inflorescence production varied greatly and was correlated with variation in plant size and light environment. There was no trade-off between past inflorescence production and the frequency of future inflorescence production. On the contrary, individuals that produced more inflorescences than predicted from their size and light environment tended to continue to do so in subsequent years also. I manipulated the resource environment of a subset of plants by removal of leaves and/or reproductive spikes. Leaf removal suppressed inflorescence production for the following 2 years, but spike removal had no effect. One year after leaf removal stored reserves were, on average, back to pre-treatment levels. There was, however, a negative effect of recent inflorescence production on storage. Plants with higher levels of storage had higher inflorescence production in the next 75 days. In C. ghiesbreghtiana the resource cost of reproduction is apparent in short-term variation in stored reserves. In contrast, annual inflorescence production does not follow a trade-off pattern between successive years, but consistently reflects both plant size and the light environment. Received: 20 October 1996 / Accepted: 25 January 1997     

Population structure and reproduction in the neotropical dioecious tree Compsoneura sprucei

Summary In rain forest study plots, the sexes of Compsoneura sprucei (Myristicaceae) were radomly distributed and similar in vegetative dimensions. The sex ratio among adults was estimated as 1.25 male: female. The population showed two flowering episodes per year, of unequal intensity. Sexual dimorphisms in order of increasing difference included the frequency of flowering, the number of flowers per inflorescence and the number of inflorescences per tree. Most females matured only 0–10 seeds per tree per flowering episode. Tree size was a better indicator of fecundity in males than females. Reproduction in both sexes was dominated by a very few prolific trees.     

Sexual dimorphism and sex ratios of two Australian dioecious species of alpine pineapple grass, Astelia alpina var. novaehollandiae and Astelia psychrocharis (Asteliaceae)

Sexual dimorphism and male biased sex ratios have been predicted for dioecious plants experiencing the limited conditions for growth and reproduction found in many alpine environments. To test these predictions, the reproductive ecology of two congeneric, co-occurring, dioecious, clonal, species was examined in the subalpine and alpine zones of Kosciuszko National Park, southeastern Australia. Specifically, plant size (vegetative cover of plants in quadrats), floral display (number of flowers per inflorescence, number of inflorescences per quadrat) and sex ratios (proportion of females in quadrats with flowers) were examined in ten populations of Astelia psychrocharis (Asteliaceae) and nine populations of Astelia alpina var. novae-hollandiae (Asteliaceae). Sexual dimorphism did occur, with males having more flowers per inflorescence (106% more flowers for A.alpina males and 12% more for A.psychrocharis males compared to females) and more inflorescences per quadrat than females (78% more inflorescences for A.alpina males and 46% more inflorescences for A.psychrocharis males compared to females). Plant size did not differ between male and female quadrats of either species, nor were there male biased sex ratios. However, plant size was related to flowering status in A.psychrocharis with the 65 quadrats that did not flower having lower vegetative cover than the 175 flowering quadrats indicating that there may be a minimum size/ cover required prior to flowering in this species. For A.alpina, all but two of the 185 quadrats randomly sampled flowered. There was no effect of altitude on plant size and very little effect of altitude on floral display for either species, apart from a slight increase in the number of inflorescences per quadrat with increasing altitude for A.psychrocharis, and slight decrease in number of flowers per inflorescence with increasing altitude for A.alpina females.     

Flower bud abortion influences clonal growth and sexual dimorphism in the understorey dioecious shrub Aucuba japonica (Cornaceae)

Sexual differences were investigated to determine the significance of flower bud abortion in the dioecious shrub Aucuba japonica Thunb. The mean number of flowers per inflorescence and the mean number of flowering inflorescences (as opposed to aborted inflorescences) per individual were greater in males than in females in 1997 and 1998. Reproductive investment by males was 0.4-times (1997) and 1.4-times (1998) that by females. In addition, females aborted 30.9% (1997) and 42.7% (1998) of their total flower buds without blooming, whereas no male flower buds aborted. One of the architectural traits of this shrub is that in the year that a flower bud is produced at the shoot apex, the shoot will branch into two or more shoots. Thus, there was less sexual difference in the number of current shoots per individual than there was in the number of flowering inflorescences. The relationship between annual growth and reproduction, and the probability of reproduction in the following year, suggested that the higher investment in female reproduction was manifested as a cost for reproductive frequency rather than as a cost for annual growth. The spatial distribution of both males and females was clumped, which may be the result of clonal growth. In addition, overall sex ratios were not skewed and the number of sprouts did not differ significantly between sexes. These results suggested that flower bud abortion by females might reduce sexual dimorphism in terms of clonal growth.     

I. LIFE HISTORY FEATURES OF THREE SEXUAL MORPHS OF ATRIPLEX CANESCENS (CHENOPODIACEAE) CLONES GROWN IN A COMMON GARDEN

Reproductive and growth parameters of Atriplex canescens were examined in a common garden study. Cloned ramets of male, female, and hermaphrodite individuals from two natural populations were planted in irrigated and nonirrigated portions of the garden. Sexual differences in growth and flowering schedules suggest tradeoffs in resource allocation between growth and reproduction. Males flowered with the greatest frequency and intensity while producing the smallest growth parameters. Females had the largest measures of vegetative growth, but flowered with the least frequency and intensity. Hermaphrodites appear to have a biology distinct from males and females, being more like females in growth and more like males in reproduction.     

迁地保护条件下两种沙冬青的开花物候比较研究

沙冬青属(Ammopiptanthus)植物是我国西北荒漠区唯一的常绿阔叶灌木。作者对吐鲁番沙漠植物园迁地保护的两种沙冬青的开花物候进行了详细的比较观察,旨在探讨它们在同一生境条件下开花特性的异同点及其影响因素。主要结果如下:(1)两种植物在开花频率、花序开放顺序、开花振幅曲线及单花寿命等开花参数上相似,但在始花时间、单株花期、花序的开花数及开放持续时间与频率分布、开花振幅等参数上明显不同;(2)在个体和群体水平上,蒙古沙冬青(A.mongolicus)始花时间均比新疆沙冬青(A.nanus)早,蒙古沙冬青开花全过程为20–21d,新疆沙冬青为13–14d;(3)蒙古沙冬青花序的开花数比新疆沙冬青多、开放持续时间长,两者在开花数(F=17.51,P<0.01)和持续时间(F=14.08,P<0.01)上均存在显著差异;(4)花序上的花大多从近基部向两端开放,开花振幅呈单峰曲线,但新疆沙冬青的开花振幅较高;(5)花序开放持续期的频率分布明显不同,新疆沙冬青较蒙古沙冬青更为集中,但两者的单花寿命稳定,均在7d左右;(6)花序上每天的开花数与其座果数呈正相关(蒙古沙冬青,r=0.885,P<0.05;新疆沙冬青,r=0.827,P<0.01),但其开花数和座果数与始花时间存在不同程度的相关关系,这些特点可能与开花对传粉者的吸引以及物种本身的遗传特性有关。对上述观察结果及其影响因素的分析表明,两种植物在开花参数上所表现出的一致性可能是受系统发育限制的,而彼此间的差异可能与其进化历史及所处的环境异质性有关,是在与环境的长期适应过程中分别形成的一些可遗传的变异;而不同年份间两种沙冬青在花序的开花数及开放持续时间上表现出的差异可能与环境温度的变化有关。这些结果对于探讨该属植物的繁殖生物学特性及其保护对策具有重要意义。     

Spatiotemporal variation in predispersal seed predation intensity

The effect of predispersal seed predation by Bruchus atomarius (Bruchidae, Coleoptera) on individual performance and population dynamics of the perennial forest herb, Lathyrus vernus (Leguminosae), was investigated in 11 permanent plots over 4 years. Seed predation and parameters describing intra-specific neighbour distance, plant size, inflorescence size, flowering phenology and current and previous herbivore damage were measured on all plants. In addition, demographic information from all plots was analysed using transition matrix population models in order to estimate the influence of seed predation on population growth rates. Predispersal seed predation rates differed significantly among years. Plot averages ranged from 0 to 83.7%. However, most of the variation occurred among individuals. Within individuals there was no consistency in predation rates among years. Exposure to herbivory, plant size and flowering phenology did not affect predation rates but individuals with larger inflorescences suffered from significantly higher predation. Seed predation in L. vernus was not influenced by neighbour distances of individual plants but it was positively correlated with the average density of seeds within plots, suggesting that seed predation is density dependent at the patch level. The reduction in population growth rate due to seed predation ranged from 0 to 7.6%. The sensitivity of population growth rate to reductions in seed production varied considerably among years and plots. This variation was mainly due to differences in the reproductive value of seeds and seedlings. The intensity of seed predation over the range found was not correlated with changes in population growth rate. The results of this study suggest that the influence of external factors, like seed predation, on population growth rate largely depends on the demographic transition rates in the investigated population.     

INFLORESCENCE TYPES AND FRUITING PATTERNS IN HAMLIN AND VALENCIA ORANGES AND MARSH GRAPEFRUIT

The structural, flowering, and fruit-setting patterns of inflorescences of mature Hamlin and Valencia orange and Marsh grapefruit trees were studied for three years. Several development patterns were found, some of which were relatively consistent for the different varieties or years of study. The sequence of anthesis on an inflorescence was: apical flower first, then basal flower, then the subapical flower. Terminals on which the earlier flowers appeared tended to have more inflorescences than those on which flowers appeared later. Inflorescences on which earlier flowers appeared also produced more flowers than those which began flowering later. Inflorescences that began flowering later were more likely to have leaves or have a greater number of leaves than earlier inflorescences. More than half of the inflorescences carried no leaves, and most of these had one flower. No evidence of a relationship between number of flowers and length of the inflorescence was found. Fruit set occurred primarily during the latter part of the flowering period. Many fruit were set on inflorescences without leaves, but on the basis of percent of flowers setting fruit, inflorescences with leaves were more productive. The greatest fruit set occurred in the subapical position on the inflorescence. With growth changes these fruit often appeared to be developing in the apical position. These patterns generally differed little from year to year. Variations may have been due to the differences in the number of flowers produced by the trees. Results were also similar between Hamlin and Valencia oranges. Patterns on Marsh grapefruit resembled those for the oranges but were frequently less consistent.     

Habitat protection,cattle grazing and density‐dependent reproduction in a desert tree

Anthropogenic activities usually trigger changes in the population density of plants. Thus, land management practices can influence density‐dependent demographic parameters and species interactions. We investigated plant‐pollinator interactions and reproduction in Prosopis flexuosa, the largest tree species in the Central Monte desert of Argentina, an important economic and cultural resource for humans and a functionally prominent species. We hypothesized that reproductive output of P. flexuosa would be limited at low densities, and that exclusion of catle grazing would enhance population density and consequently interaction frequency with pollinators and reproductive success. The study was conducted in and around Ñacuñán Biosphere Reserve (Mendoza, Argentina), where cattle grazing has been excluded for over 35 years. Working in five pairs of protected and cattle grazed 1‐ha plots, we recorded density of adult trees, pollinator visitation frequency to inflorescences and seeds per inflorescence in focal trees. Adult tree density was higher in protected plots than in cattle grazed plots. Density of reproductive trees was positively correlated with seed production, suggesting positive density dependence for reproduction (Allee effect). Pollinator visitation to inflorescences and seed production was higher in protected plots compared with plots under cattle grazing. Suppression of anthropogenic degradation has resulted in higher adult tree density in protected plots, indirectly higher pollinator visitation to inflorescences and higher reproductive success of trees. Increased frequency of plant‐pollinator interactions and tree reproduction suggest success of management practices aimed at protecting P. flexuosa woodlands.     

Population ecology in natural<Emphasis Type="Italic">Syneilesis palmata</Emphasis> stands: I. relationship between flowering and plant size

In this study, the relationship was explored between flowering potential and plant size in a population ofSyneilesis palmata (THUNB.) MAX., Usannamul. For this perennial shade-tolerant herb, the flowering rate, plant size, and changes that occurred in the next year after flowering were monitored in two different types of natural stands: moderately shaded (MS; for 11 years) and severely shaded (SS; 7 years). The correlation coefficients (CC) between inflorescence number and other factors, i.e., widths of the first leaf, second leaf, or total leaf breadth, were significant at the 1% level in both stands. In particular, the CC was >0.9 between the number of inflorescences and total leaf breadth. Although the mean flowering rates averaged 6.8% in MS and 8.6% in SS, those values varied widely, ranging from 0 to 53% among years. These rates were not related to petiole length but did increase in proportion to the width of the leaf blade. Flowering began when plants moved beyond the medium size class; the rate in the MS environment was 100% for the largest-sized plants. For plants in the SS, however, the highest rate was 35%. Over the 11 years, 62.8% of all plants in the MS never flowered, while another 30.8% bloomed only once. In comparison, plants observed for 7 years in the SS showed a non-flowering rate of 78.1% and a one-time-only rate of 19.2%. In the next year after flowering, 57.6% of the MS plants exhibited reduced growth while 25.8% of those individuals each split into two plants. In the SS, 37.8% had decreased development the following year, and a mortality rate of 37.8%.     

Flowering dynamics in Arum italicum (Araceae): relative role of inflorescence traits, flowering synchrony, and pollination context on fruit initiation

We studied the relative role of inflorescence traits, flowering synchrony, and pollination context for infructescence and fruit initiation in two Spanish populations of Arum italicum, a species in which inflorescences are the pollination unit. In this species, a specialized inflorescence organ, the appendix, is important for pollinator attraction. However, the short floral longevity and the production of mostly one inflorescence per plant make its pollination potentially dependent on strong flowering synchrony and on external factors not controlled by the plant (the pollination context). The flowering period in both sites lasted >3 mo. Day-to-day variation in simultaneous antheses was high, and 11-50% of antheses occurred on days during which no pollen donor was present. Inflorescence traits, flowering synchrony, and between-plant distance all influenced infructescence and fruit initiation, but their relative importance differed between sites. In one large population, infructescence initiation was positively related to inflorescence traits; in a smaller population infructescence initiation increased with the number of donor inflorescences. In both sites, percentage of fruits initiated per infructescence was dependent on a combination of inflorescence traits, flowering synchrony, and between-plant distance. Plants producing 2-4 inflorescences had higher probability of infructescence initiation and overlapped their antheses with more plants than single-inflorescence ones.     

Evidence for stabilising selection acting on flowering time in Arum maculatum (Araceae): the influence of phylogeny on adaptation

The relationship between flowering time and reproductive success was investigated in the fly-pollinated, monoecious perennial herb Arum maculatum L. (Araceae). This species temporarily traps its principle pollinator, a psychodid midge. Probability of fruit set was analysed in relation to early, peak and late periods of the flowering phenology of four British populations between 1992 and 1997. In three out of five cases, plants which flowered during early and late periods were significantly less likely to set fruit. In addition, one population showed a similar relationship for percentage fruit set of individual inflorescences, and seeds from peak-flowering plants were significantly heavier. There was no variation in number of female flowers per inflorescence over the flowering season. Probability of fruit set appears to be mediated by the likelihood of trapping psychodid midges that have previously been trapped and picked up pollen, an unlikely event during early and late flowering periods when few inflorescences are open. The majority of plants in all populations produce only one inflorescence which means that timing of flowering may be crucial to reproductive success. We interpret our findings as evidence that stabilising selection may be acting on some populations and/or during some years. The ultimate cause, however, can be related to the very short (12–18 h) female phase of each inflorescence, a phylogenetically conservative trait within the Araceae. Received: 19 August 1998 / 15 February 1999     

Pollination limitation,costs of capsule production and the capsule-to-flower ratio in Dendrobium monophyllum F. Muell. (Orchidaceae)

Abstract In perennial plants, life-history theory suggests that natural selection should result in the optimization of fruit-to-flower ratios within the limits imposed by the trade-offs between resource allocation for present reproduction and future growth and reproduction. The tropical orchid Dendrobium monophyllum F. Muell., an epiphyte or lithophyte, offers no nectar rewards, is self-incompatible and has a capsule-to-flower ratio of about 1:14. The influence of pollination limitation and the costs of capsule production on capsule-to-flower ratios were assessed using experimental and field studies in which individual plants were observed for 3 years. Pollinators visited about 80% of flowers, and capsule production was significantly related to inflorescence size and pollinaria removal. About nine pollinator visits occurred per capsule. Pollinator visitation and capsule production did not vary significantly between years. The inflorescence size classes most successful in capsule production were also the most frequent in natural populations. The experimental supplementation of outcross pollen to flowers increased capsule set over controls by 45% within a year, but was limited to about 53%. A capsule-to-flower ratio of 1:2 in experimental plants significantly decreased the subsequent growth and flowering of individuals relative to controls. A capsule-to-flower ratio above 1:10 in naturally pollinated plants decreased flowering in the subsequent year. Thus, it is suggested than an increase in capsule production above 10% would not necessarily correlate with greater reproductive fitness because of the increased cost of capsule production. The capsule-to-flower ratio recorded in this study could be evolutionarily stable because of trade-offs between selection for pollinator attraction and the cost of capsule production. The production of surplus flowers appears to function in pollinator attraction and increases fitness through male function.     

Tiller dynamics and assimilate partitioning in Lolium perenne with particular reference to flowering

The fate of 100 seedling plants of Lolium perenne L. was studied over a period of 2 years in a field plot. The birth and death of tillers and the production of inflorescences was followed, and the components of seed yield were recorded in detail in the first year. The pattern of distribution of ¹⁴CO₂ assimilated by the main shoot was examined at monthly intervals and during the first flowering season the distribution of ¹⁴C-assimilate from individual leaves and from the inflorescence was also studied. The capacity of individual tillers to assimilate ¹⁴CO₂ prior to flowering and the re-distribution of previously accumulated assimilate during seed growth were also assessed. Plants died at a more or less constant rate with time and only 54 survived to the end of the 2–yr period. First year mortality was associated with severe grazing or cutting but in the second year the death of ungrazed plants was observed. There was great variability in the production of tillers by surviving plants. In both years the number of live tillers per plant increased from July to the end of April with particularly rapid tillering in March and April establishing the maximum value for each year. There was a similar phase of rapid tillering after flowering in July. The number of live tillers per plant declined by 50% during stem elongation and inflorescence emergence and the majority of dead tillers were young secondary (in the first year) and tertiary (in the second year) tillers with a mean age of 40 days. Such tillers had poor assimilatory capacity prior to the onset of death and were not supplied with assimilate from the main shoot. Most of the plants surviving at the end of the experiment flowered in both years and one quarter of the maximum number of live tillers per plant recorded in April of each year produced inflorescences. The earlier a tiller was produced the greater was its chance of flowering and the greater its production of seed. The greater weight of seed produced was associated with the development of more seed-bearing florets per spikelet. There was relatively little export of “C-assimilate from the flowering main shoot, and the lower internodes formed the major sink for post-anthesis assimilate. The growth of seeds appeared to be relatively independent of the leaves for current assimilate. There was some evidence that assimilate accumulated in lower internodes was remobilised and utilised in the growth of seeds and new tillers. Overall, the results confirm the view that the grass plant is a dynamic population of short-lived tillers and indicate that increasing competition for assimilate at flowering exerts a major influence on the production and survival of tillers.     

Genetic interactions in the control of flowering time and reproductive structure development in tomato (Solanum lycopersicum)

Different tomato (Solanum lycopersicum) mutants, affected in flowering time, reproductive structure or plant architecture, were crossed to produce double mutants in order to investigate gene interactions in flowering regulation in this autonomous species with a sympodial growth habit. The compound inflorescence: uniflora, uniflora: self pruning, uniflora: blind, and jointless: uniflora double mutants all produced solitary flowers like their uniflora parent, instead of inflorescences. All double mutants were late flowering. uniflora: blind and uniflora: self pruning had flowering times intermediate between those of their two parents. jointless: uniflora and compound inflorescence: uniflora flowered later than uniflora, the mutant with the most delayed flowering. All double mutants developed strong lateral shoots at node levels approximately corresponding to the level at which their parent cultivars initiated their first reproductive structure, which is a typical trait of uniflora. These results suggest that the UNIFLORA gene acts upstream of the other investigated genes in controlling flowering in tomato, and that floral transition of the primary shoot and floral transition of sympodial segments are regulated differently.     

Additive and non-additive effects of simulated leaf and inflorescence damage on survival, growth and reproduction of the perennial herb Arabidopsis lyrata

Herbivores may damage both leaves and reproductive structures, and although such combined damage may affect plant fitness non-additively, this has received little attention. We conducted a 2-year field experiment with a factorial design to examine the effects of simulated leaf (0, 12.5, 25, or 50% of leaf area removed) and inflorescence damage (0 vs. 50% of inflorescences removed) on survival, growth and reproduction in the perennial herb Arabidopsis lyrata. Leaf and inflorescence damage negatively and independently reduced flower, fruit and seed production in the year of damage; leaf damage also reduced rosette size by the end of the first season and flower production in the second year. Leaf damage alone reduced the proportion of flowers forming a fruit and fruit production per plant the second year, but when combined with inflorescence damage no such effect was observed (significant leaf × inflorescence damage interaction). Damage to leaves (sources) caused a greater reduction in future reproduction than did simultaneous damage to leaves and inflorescences (sinks). This demonstrates that a full understanding of the effects of herbivore damage on plant fitness requires that consequences of damage to vegetative and reproductive structures are evaluated over more than 1 year and that non-additive effects are considered.     

Evolution of floral display in Eichhornia paniculata (Pontederiaceae): genetic correlations between flower size and number

The evolution of floral display is thought to be constrained by trade‐offs between the size and number of flowers and inflorescences. We grew in the glasshouse 60 maternal families from each of two Brazilian populations of the annual herb, Eichhornia paniculata. We measured flower size, daily flower number, and total flower number per inflorescence, and two indices of module size, leaf area and age at flowering. We also assessed the size and number of inflorescences produced over 6 weeks. All floral traits exhibited significant heritable variation, some of which was due to genetic variation in module size. Genetic (maternal family) correlations between daily and total flower number did not differ from 1.0, indicating that display size (daily flower number) cannot evolve independently from total flower number per inflorescence. Genetic correlations between flower size and daily flower number ranged from negative to positive (r=–0.78 to +0.84), depending on population and inflorescence. Positive correlations occurred when variation in investment per inflorescence was high so that some families produced both larger and more flowers. These correlations became zero when we controlled for variation in module size. Families that flowered later produced fewer, larger inflorescences (r=–0.33, –0.85). These data support theoretical predictions regarding the combined effects of variation in resource acquisition and allocation on traits involved in trade‐offs, and they emphasize the hierarchical organization of floral displays. Our results imply that patterns of resource allocation among inflorescences influence evolutionary changes in flower size and number per inflorescence.     

Seasonal variation in honeyeater foraging behaviour,infloresence abundance and fruit set in Banksia spinulosa (Proteaceae)

Seasonal variation in the foraging behaviour of honeyeaters and the production of fruit were examined in relation to the flowering intensity of Banksia spinulosa over two flowering seasons. The abundance of inflorescences was greater in the mid than in the early and late periods of the flowering season. In the mid period, many plants were blooming and each plant had many flowering inflorescences. Inflorescences received most visits by honeyeaters in the early flowering period; the visitation rate declining as flowering progressed. Eastern spinebills were the most common floral visitors at all times during the season. The number of foraging probes made at inflorescences by eastern spinebills did not differ throughout the season. Foraging movements between inflorescences on the same plant were more frequent in the mid period than in the early and late periods. Long distance movements between plants (more than 10 m apart) were promoted by aggressive interactions between honeyeaters. Inflorescences flowering in the late period were less likely to develop follicles because there were fewer visits by birds and/or because resources had been allocated to inflorescences pollinated earlier in the season. The number of follicles produced per infructescence did not differ between flowering periods. Overall, the number of inflorescences produced per plant, the number of visits received per inflorescence and the proportion of inflorescences that developed follicles were greater in 1987 than in 1988.     

Is analysing the nitrogen use at the plant canopy level a matter of choosing the right optimization criterion?

Herbivores can have strong deleterious effects on plant growth, reproduction, and even survival. Because these effects might be strongly interrelated, the direct consumptive effects of herbivores and a variety of indirect effects are difficult to untangle. Reductions in growth, for example, may strongly impact the flowering behaviour of plant species in the current season, but at the same time incur costs to survival, growth and reproduction in the next growing season(s). To get better insights in the effects of herbivory on the flowering behaviour of the long-lived polycarpic grassland herb Primula veris L., flowering patterns were monitored over ten consecutive years under two treatments (grazing and control mowing regimes). We tested the hypothesis that the size at flowering was affected by the presence of herbivores, and whether this translated into costs to future reproduction and survival. Overall, grazed plants were significantly smaller than control plants, and the size at which plants flowered was also significantly smaller when herbivores were present. The transition probability of flowering and of surviving into the next year was significantly smaller for all plants in the current year if they had been grazed than if they had been mown, indicating that herbivory incurred costs to both flowering and survival. Grazed plants also needed longer to start flowering, had fewer flowers and flowered less frequently, causing a significantly lower proportion of flowering adults in the population. These results suggest that the observed regression in plant size due to herbivory does not allow plants to capture enough resources to guarantee regular flowering in the longer run.