Constraints on flowering phenology in a phryganic (East Mediterranean shrub) community

The phenological pattern of flowering at the community level was studied in a Greek phryganic ecosystem near Athens for 4 consecutive years. Flowering is strongly seasonal: 80% of the insect-pollinated flora, which consists of 133 species, blooms between February and June. There is a variably expressed secondary flowering period in autumn. The pollinating fauna follows a strongly correlated pattern of abundance. Two types of plants were distinguished: pauciflorous species bearing <10 flowers that are large compared to the plant body, and multiflorous species with many small flowers. Pauciflorous species flower in the winter half of the year, while multiflorous species flower mainly in the summer half. The mean flower life spans are 9 and 3 days, respectively. The duration of flowering (DF) for each species is 55 days on average, which is long compared to other communities. The DF shows year-to-year variations, concomitant with the vicissitudes of the climate. The start of flowering of a species is statistically correlated with the temperature in the previous month, not with rainfall; its end date of flowering only partly compensates for the time gained or lost. DF is maximal in winter. The average flower life span of species flowering at any given date varies strongly and independently of the average DF. We tested the hypothesis that flowering phenology is set by phylogenetic and life form constraints. This could not be corroborated for phylogeny, evidently because of the overriding influence of the mediterranean climate, and probably also for biogeographical reasons. In contrast, life forms and multiflorous and pauciflorous species show strong differences. Many (51) of the species are therophytes; we tested the hypothesis that because of their annual habit they would be more dependent on pollination than perennials. Thus we anticipated that therophytic species would be differentiated from perennials in their flowering phenologies. This is not corroborated. We therefore conclude that the seed bank plays a role that is analogous to that of a perennial plant body.     

Directional selection on initial flowering date in Phlox drummondii (Polemoniaceae)

The fitness effects due to initial flowering date in Phlox drummondii were determined for three populations in central Texas (USA) over 3 yr (1990-1992). Mean fitness (seed set) always decreased with the later initiation of flowering. The likelihood of a plant fruiting differed with flowering date in five of the six instances (population by year combinations). Though plants that initiated flowering later tended to have spent more time in the vegetative stage and tended to die later in the year than did earlier flowering plants, this was not sufficient to overcome the reproductive penalties of flowering late. Plants that initiated flowering later in the season spent less time in the adult phase and were smaller. The mean number of flowers, fruits, and seeds per flowering plant always decreased with later flowering. Fruit set was negatively correlated with flowering date in four of the six population by year combinations. Nonparametric fitness functions were used to summarize predicted fitness among different initial flowering dates for each population on a yearly basis. Predicted mean fitness always declined nonlinearly with later flowering; the earliest flowering plants always had the highest predicted fitness. These fitness functions describe directional selection for the early initiation of flowering.     

Flowering dynamics of Orchis morio L. and Herminium monorchis (L.) R.Br, at two sites in eastern England

Information taken from two long-term demographic studies on Orchis morio L. and Herminium monorchis (L.) R.Br, is used to explore some of the factors which influence flowering. The proportion of plants which flowered each year varied considerably between species, flowering in O. morio exceeding 40% in all years except one over an 18 year period; over a 30 year period (1966–95), the number of plants of Herminium in flower never exceeded 36% of the population and no inflorescences were produced in 1977 and 1991. The relationship between flowering in Herminium in a given year and the monthly rainfall and temperature for the current and 3 previous years was analysed using logistic regression. Best fits were obtained using data for the summer months in the previous year, with an increasing flowering rate with rainfall and a decline with temperature. It is hypothesized that drought and high temperatures in the summer reduce leaf area and cause premature senescence and the death of leaves, with the result that not enough carbohydrates are stored to enable plants to support or initiate inflorescences the following year. For species such as Orchis morio which produce leaves in the autumn and remain green, summer drought causes no problems as they have no above ground organs. Factors which influence flowering in this species are as yet unknown.     

Phenological Sensitivity of Early and Late Flowering Species Under Seasonal Warming and Altered Precipitation in a Seminatural Temperate Grassland Ecosystem

Shifts in flowering phenology of plants are indicators of climate change. The great majority of existing phenological studies refer solely to gradual warming. However, knowledge on how flowering phenology responds to changes in seasonal variation of warming and precipitation regimes is missing. We report the onset of 22 early (flowering before/within May) and 23 late flowering (flowering after May) species in response to manipulated seasonal warming (equal to + 1.2°C; last 100-year summer/winter warming), additional winter rainfall, and modified precipitation variability (including a 1000-year extreme drought event followed by heavy rainfall) over the growing season in two consecutive years for a species-rich temperate grassland ecosystem. The average onset of flowering (over 2 years) was significantly advanced 3.1 days by winter warming and 1.5 days by summer warming compared to control. Early flowering species responded to seasonal warming in both years, while late-flowering species responded in only 1 year to summer warming. The average onset of early flowering species was significantly advanced, 4.9 days by winter warming and 2.3 days by summer warming. Species-specific analysis showed that even within the early flowering community there were divergences. A positive correlation between plant height and shift in flowering onset was detected under winter warming (R² = 0.20, p = 0.005). The average onsets of early and late flowering community were affected by neither winter rain nor growing season precipitation variability. Seasonal differences in warming, and particularly winter warming, might alter community dynamics among early and late flowering species which can cause shifts in the seasonal performances of temperate ecosystems.     

Time measurement and the control of flowering in plants

Many plants are adapted to flower at particular times of year, to ensure optimal pollination and seed maturation. In these plants flowering is controlled by environmental signals that reflect the changing seasons, particularly daylength and temperature. The response to daylength varies, so that plants isolated at higher latitudes tend to flower in response to long daylengths of spring and summer, while plants from lower latitudes avoid the extreme heat of summer by responding to short days. Such responses require a mechanism for measuring time, and the circadian clock that regulates daily rhythms in behaviour also acts as the timer in the measurement of daylength. Plants from high latitudes often also show an extreme response to temperature called vernalisation in which flowering is repressed until the plant is exposed to winter temperatures for an extended time. Genetic approaches in Arabidopsis have identified a number of genes that control vernalisation and daylength responses. These genes are described and models presented for how daylength might regulate flowering by controlling their expression by the circadian clock. BioEssays 22:38-47, 2000.     

Environmental influences on the phenology and abundance of flowering by Androsace septentrionalis (Primulaceae)

We studied the timing and abundance of flowering by Androsace septentrionalis L. (Primulaceae), an indeterminate winter annual or short-lived perennial, in 2 × 2 m plots at the Rocky Mountain Biological Laboratory in Colorado, USA, from 1982 to 2000. Flowers were counted every other day for most or all of the growing season in seven plots in a rocky meadow habitat and nine plots in a wet meadow habitat. The phenology and abundance of flowering were both highly variable, with mean dates of first flowering ranging from 16 May to 12 July and maximum daily counts of flowers ranging from 1 to 1187. Snowmelt date was the primary determinant of timing of flowering. For rocky meadow plots, the previous year's summer precipitation and the current year's average minimum temperature in May had significant effects on maximum number of flowers produced, but no environmental variable we considered was significantly correlated with flower abundance in the wet meadow plots. Length of flowering in individual plots ranged from 2 to 85 d, and many plot-years had both primary (about 1 mo) and secondary (about 10-12 d) flowering periods. The predicted increase in variability of precipitation accompanying climate change will affect negatively the long-term abundance and persistence of this species at our study site.     

Plant size and season of burn affect flowering and fruiting of the grasstree Xanthorrhoea preissii

Flowering and fruiting were assessed on 14 populations of the grasstree, Xanthorrhoea preissii Endl., occurring in the Darling Range near Perth, Western Australia. Independent of site, season of burn or year of flowering, there was a strong relationship between plant height, which varied from 0.1 to over 2 m, and the incidence of postfire flowering, which varied from 1% (winter burn) to 75% (summer burn) of grasstrees present. There was no relationship between inflorescence dimensions, or flower or fruit production on a spike basis, and plant size/age (height). When standardized for height, spring‐burnt populations produced 40% as many inflorescences as autumn‐burnt populations and 20% as many as summer‐burnt populations. Inflorescences produced by spring‐burnt plants were moderately smaller than those by summer–autumn‐burnt plants. Fruit density per spike in autumn‐burnt plants was 80% of that in spring–summer‐burnt plants. The net effect was an average of 70 000 fruits produced per 100 summer‐burnt plants, 22 000 in autumn‐burnt plants, and 14 000 in spring‐burnt plants. Ecophysiological explanations of these results and their implications for population dynamics have yet to be explored.     

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.     

Effects of experimentally imposed climate scenarios on flowering phenology and flower production of subarctic bog species

Climate scenarios for high‐latitude areas predict not only increased summer temperatures, but also larger variation in snowfall and winter temperatures. By using open‐top chambers, we experimentally manipulated both summer temperatures and winter and spring snow accumulations and temperatures independently in a blanket bog in subarctic Sweden, yielding six climate scenarios. We studied the effects of these scenarios on flowering phenology and flower production of Andromeda polifolia (woody evergreen) and Rubus chamaemorus (perennial herb) during 2 years. The second year of our study (2002) was characterized by unusually high spring and early summer temperatures. Our winter manipulations led to consistent increases in winter snow cover. As a result, average and minimum air and soil temperatures in the high snow cover treatments were higher than in the winter ambient treatments, whereas temperature fluctuations were smaller. Spring warming resulted in higher average, minimum, and maximum soil temperatures. Summer warming led to higher air and soil temperatures in mid‐summer (June–July), but not in late summer (August–September). The unusually high temperatures in 2002 advanced the median flowering date by 2 weeks for both species in all treatments. Superimposed on this effect, we found that for both Andromeda and Rubus, all our climate treatments (except summer warming for Rubus) advanced flowering by 1–4 days. The total flower production of both species showed a more or less similar response: flower production in the warm year 2002 exceeded that in 2001 by far. However, in both species flower production was only stimulated by the spring‐warming treatments. Our results show that the reproductive ecology of both species is very responsive to climate change but this response is very dependent on specific climate events, especially those that occur in winter and spring. This suggests that high‐latitude climate change experiments should focus more on winter and spring events than has been the case so far.     

Experimental icing affects growth,mortality, and flowering in a high Arctic dwarf shrub

Effects of climate change are predicted to be greatest at high latitudes, with more pronounced warming in winter than summer. Extreme mid‐winter warm spells and heavy rain‐on‐snow events are already increasing in frequency in the Arctic, with implications for snow‐pack and ground‐ice formation. These may in turn affect key components of Arctic ecosystems. However, the fitness consequences of extreme winter weather events for tundra plants are not well understood, especially in the high Arctic. We simulated an extreme mid‐winter rain‐on‐snow event at a field site in high Arctic Svalbard (78°N) by experimentally encasing tundra vegetation in ice. After the subsequent growing season, we measured the effects of icing on growth and fitness indices in the common tundra plant, Arctic bell‐heather (Cassiope tetragona). The suitability of this species for retrospective growth analysis enabled us to compare shoot growth in pre and postmanipulation years in icing treatment and control plants, as well as shoot survival and flowering. Plants from icing treatment plots had higher shoot mortality and lower flowering success than controls. At the individual sample level, heavily flowering plants invested less in shoot growth than nonflowering plants, while shoot growth was positively related to the degree of shoot mortality. Therefore, contrary to expectation, undamaged shoots showed enhanced growth in ice treatment plants. This suggests that following damage, aboveground resources were allocated to the few remaining undamaged meristems. The enhanced shoot growth measured in our icing treatment plants has implications for climate studies based on retrospective analyses of Cassiope. As shoot growth in this species responds positively to summer warming, it also highlights a potentially complex interaction between summer and winter conditions. By documenting strong effects of icing on growth and reproduction of a widespread tundra plant, our study contributes to an understanding of Arctic plant responses to projected changes in winter climatic conditions.     

Seed traits are pleiotropically regulated by the flowering time gene PERPETUAL FLOWERING 1 (PEP1) in the perennial Arabis alpina

The life cycles of plants are characterized by two major life history transitions—germination and the initiation of flowering—the timing of which are important determinants of fitness. Unlike annuals, which make the transition from the vegetative to reproductive phase only once, perennials iterate reproduction in successive years. The floral repressor PERPETUAL FLOWERING 1 (PEP1), an ortholog of FLOWERING LOCUS C, in the alpine perennial Arabis alpina ensures the continuation of vegetative growth after flowering and thereby restricts the duration of the flowering episode. We performed greenhouse and garden experiments to compare flowering phenology, fecundity and seed traits between A. alpina accessions that have a functional PEP1 allele and flower seasonally and pep1 mutants and accessions that carry lesions in PEP1 and flower perpetually. In the garden, perpetual genotypes flower asynchronously and show higher winter mortality than seasonal ones. PEP1 also pleiotropically regulates seed dormancy and longevity in a way that is functionally divergent from FLC. Seeds from perpetual genotypes have shallow dormancy and reduced longevity regardless of whether they after‐ripened in plants grown in the greenhouse or in the experimental garden. These results suggest that perpetual genotypes have higher mortality during winter but compensate by showing higher seedling establishment. Differences in seed traits between seasonal and perpetual genotypes are also coupled with differences in hormone sensitivity and expression of genes involved in hormonal pathways. Our study highlights the existence of pleiotropic regulation of seed traits by hub developmental regulators such as PEP1, suggesting that seed and flowering traits in perennial plants might be optimized in a coordinated fashion.     

Is the threshold size for flowering in Cynoglossum officinale fixed or dependent on environment?

In the monocarpic perennial Cynoglossum officinale L. the probability of flowering is related to the size of the plant. In previous work it was observed that this relation varies between years. We hypothesized that variable conditions during the winter, the period of vernalization, explain this variation.
We collected plants from the field in autumn and placed these under different simulated winter conditions in a climate room. In contrast to our hypothesis, the probability of initiating flowering at a given size was not affected by: ( a ) the temperature during the cold period, ( b ) the duration of the cold period, or ( c ) the application of a plant hormone (GA₃) or an inhibitor of gibberellin synthesis (paclobutazol) during the cold period. Winter cold is not necessary for floral initiation, and is only required for elongation of the inflorescence. It is unlikely that winter temperature affects the fraction of plants flowering.
Subsequent morphological investigation of flower development in material collected in the field showed that large plants had primordial inflorescences well before vernalization, sometimes as early as August. In plants grown from seeds under constant conditions in a climate room, the probability of initiating the inflorescence differed for plants grown at various temperatures (34·1% at 15°C, 100% at 20°C, and 95% at 25°C). Our results suggest that environmental conditions in August and September, up to 10 months before actual flowering, could affect the fraction of flowering plants.     

Rainfall and flowering synchrony in a tropical shrub: Variable selection on the flowering time ofErythroxylum havanense

Summary We tested the adaptive significance of flowering synchrony by means of a quantitative analysis of selection and by flowering induction experiments with the deciduous shrubErythroxylum havanense. Temporal schedules of flower and fruit production were determined for a local population (in three sites) in a Mexican seasonal forest for 2 years (1987–1988). The consequences of natural variation in flowering time (flowering initiation day) on maternal reproductive success (fecundity) were evaluated. We observed high levels of inter- and intraindividual flowering synchrony in 1987, but not in 1988 and this contrast was related to differences in rainfall patterns between the two years. A significant proportion (15.4%) of the phenotypic variation in flowering initiation day was accounted for by environmental variance. The expression of phenotypic variance of flowering time and, consequently, the opportunity for selection to act, are controlled by annual variation in rainfall. Despite the between-year difference in flowering synchrony, we detected a relatively intense directional selection on flowering initiation day in both years, but selection coefficients were of opposite sign (standardized directional gradients were –0.326 and 0.333 for 1987 and 1988, respectively). For both years there was a significant relationship between individual relative fitness and the number of neighbouring flowering plants in a given day, suggesting positive frequency-dependent selection.     

猕猴桃观花品种‘江山娇’与中华猕猴桃回交后代的性别分离与开花性状变异

以猕猴桃种间杂种品种‘江山娇’（Actinidia chinensis Planch×A.eriantha Benth）与中华猕猴桃（A.chinensis Planch）雄株杂交得到的杂交后代群体为实验材料,于2012、2013和2016年分别对该群体的雌雄性别比及其开花性状进行了调查。结果表明,杂交群体的雌雄性别比例小于1：1,即雄株偏多。杂交后代的花瓣颜色以红色为基础,但红色的分布区域、深浅及类型出现明显分离。聚类分析显示,这些杂交后代可通过花瓣CMYK色卡取值分为4个类群,其中猩红色与紫罗兰红色2个变异类型与观察表型完全一致。杂交后代群体的始花期、开花天数、花朵大小、开花量及单花花瓣数等性状均出现广泛分离,且因不同年份而出现变化。群体中杂交个体进入始花期的平均时间跨度为14 d,群体的始花期进入高峰时个体平均比例仅占群体的25.5%。2016年群体开花时间最长,最多有47.4%的个体开放10~13 d;2013年杂交群体的开放时间最短,有55.2%的后代开花3~5 d。本研究筛选出一批花瓣数多、花朵较大或单花序花朵较多的优良单株,并在后代群体中共发现21个含不同花数的花序组合类型。     

Environmental regulation of flowering

The timing of flower initiation is a highly plastic developmental process. To achieve reproductive success, plants must select the most favourable season to initiate reproductive development; this in turn requires continuous monitoring of environmental factors and a properly response. Environmental factors which change in a predictable fashion along the year, such as light and temperature, are the most relevant in terms of selection of the flowering season. In Arabidopsis and more recently in a few other species, molecular genetic analyses are providing a way to identify the genes involved in the regulation of flowering time. From gene sequences it is possible to develop hypotheses regarding molecular function and to infer some of the molecular mechanisms involved in the environmental regulation of flowering time. In this paper, we summarize recent discoveries concerning the mechanisms which plants use to perceive and respond to major environmental factors (light and temperature) and their different components. We focus mainly on annual plants and especially on Arabidopsis because most of the available molecular and functional data come from this species. However, additional information arising from other plant systems is also considered.     

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

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

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.     

Seasonal pattern of dry matter allocation in Dactylorhiza lapponica (Orchidaceae) and the relation between tuber size and flowering

A population of the tuberous orchid Dactylorhiza lapponica was sampled from June 2000 to June 2001 in the Sølendet Nature Reserve, Central Norway. Dry matter of aerial shoots, old tubers and new (replacement) tubers was measured, as well as reproductive status during 1999–2001. The biomass of the new tuber was found to continue to increase after the assimilation from photosynthesis had ceased in August. It is suggested that the increase is caused by mycotrophic activity and reallocation of nutrients from the aerial shoots. There was a clear relationship between tuber size and flowering behaviour. Individuals with flower primordia had the largest replacement tuber, whereas those that flowered in the sampling season or remained vegetative throughout 1999–2001 had the smallest. Individuals that flowered in the sampling season had the largest old tuber. Those, which had not flowered for at least two years, but had developed flower primordia, had the second largest, and those that remained vegetative throughout 1999–2001 had the smallest one. Individuals with a replacement tuber less than 0.22 g in October, have a very low probability of flowering the following season. Flowering entails a cost in terms of reduced biomass of the replacement tuber compared to vegetative individuals with old tubers of similar size. Allometric analyses revealed that above-ground biomass and biomass of replacement tubers increased with the biomass of old tubers in vegetative individuals. For generative individuals, however, above-ground biomass was only weakly related to below-ground biomass.     

Between-year variation in flowering and fruit set in frost-prone and frost-sheltered populations of dioecious Rubus chamaemorus

Summary The flowering and fruiting patterns of the dioecious perennial herb Rubus chamaemorus L. were studied in frost-prone (open) and frost-sheltered (Shaded) habitats in northern Sweden over 6 years. The number of ramets with flower buds, the proportion of flower buds that opened, and fruit set varied markedly between years. In the frost-prone populations, the occurrence or absence of detrimental frosts during the development of flowers and fruit could explain much of the variation, both in the proportion of flower buds that developed into flowers, and in fruit set. In the frost-sheltered populations, most female flowers that did not develop into fruit aborted without any signs of physical damage and before any ovules had begun to enlarge. Flower mortality caused by herbivores feeding on reproductive parts was commonly low, but reached values higher than 10% in one of the shaded populations. Hand-pollination increased the proportion of ovules producing seeds in the mature fruits by about 20%, and in one year also increased fruit set significantly in one population. Fruit-producing female ramets had a higher mortality and a lower probability of flowering in the subsequent year than male ramets and non-fruiting female ramets. In R. chamaemorus, the conditions for fruit maturation are highly unpredictable at the time of flower initiation. It is suggested that the apparent over-initiation of flower buds is advantageous, as it allows the plant to attain a high reproductive success in years favourable for flowering and fruit development.     

A rare continual flowering strategy and its influence on offspring quality in a gynodioecious plant

The majority of angiosperms have a single annual bout of reproduction; species that flower continually throughout the year are rare. Ochradenus baccatus is a gynodioecious, desert shrub whose principal flowering period is associated with the winter rains, although large individuals also remain in flower during the hot, dry summer. The goal of this study was to examine the reproductive consequences of continual flowering in a large population of O. baccatus in Israel. Over the two years of this study, 60% of individuals flowered continuously. The number of fruit and seeds per fruit were greater in winter. Winter seeds were ～12% heavier than summer seeds and had significantly higher germination rates (80 vs. 60%, respectively). Although summer seeds were smaller and less likely to germinate, we propose that the benefit derived from their production lies in their ability to capitalize on the first winter rains. These early rain events provide a head start on establishment and growth in the hostile desert environment. Plants that delay reproduction until the onset of rains risk having their offspring face the dry conditions of spring and summer.