Flowering seasonality and flower characteristics ofLoranthus acaciae Zucc. (Loranthaceae): Implications for advertisement and bird-pollination

The flowering biology and pollination ecology ofLoranthus acaciae was studied at Hazeva in the northern Arava Valley in Israel. Flowers at anthesis had red anthers, a red stigma and a green corolla which turned red as a postfloral phenomenon. Their flowering period was approximately 10 months long (from mid-June until mid-April) during which time two main flowering patterns were distinguished. Some plants flowered twice a year, with separate summer and winter flowering periods; other plants flowered continuously, with two peaks, one in the summer and one in the winter. Several significant differences between summer and winter flowering and fruiting were found: (1) the summer flowering period was shorter than that of winter, (2) flowering synchrony between individual plants was lower in summer than in winter, (3) in summer the plants produced a larger proportion of female flowers, whereas in winter most of the plants produced a larger proportion of hermaphrodites, (4) in summer a limited number of plants produced smaller flowers while the majority produced normal-sized flowers, whereas in winter the entire population produced only normal-sized flowers, and (5) fruit set percentage was lower in summer than in winter.L. acaciae was found to be self-compatible, but, since it was not spontaneously self-pollinated, it showed high dependence on pollinator activity. In summer the flowers were visited by a wide spectrum of pollinators, both birds and insects, while in winter flowers were visited almost exclusively by the orange-tufted sunbird (Nectarinia osea osea, Nectariniidae). These seasonal changes in flowering characteristics and pollinator activity could explain why reproductive success is higher in winter than in summer.     

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

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

Relationships between flowering phenology,plant size and reproductive success in shape Lotus corniculatus (Fabaceae)

Over three years the flowering phenology of individuals of Lotus corniculatus has been studied in relation to fruit set and seed predation to determine the relationships between four components of flowering time, plant size and reproductive success. Timings of first and peak flowering, and duration and synchrony of flowering differed between individuals in the same years. Between years, timing of first flowering was highly correlated for the same individuals, and was closely correlated with plant size and duration of flowering–larger plants flowered earlier and for a longer period. Peak flowering and synchrony were not correlated between-years for individuals.Fruit production and seed predation were correlated with some of the components of flowering phenology in some years, but not in others. The inconstancy of these relationships suggests that directional or stabilising selection is not acting consistently on the aspects of reproductive success studied in this work. The inconstancy of selection may result in the rather asynchronous flowering phenologies of individuals of L. corniculatus observed.We emphasize the importance of studying different components of flowering phenology in relation to individual plant size over several seasons. This work has shown that plant size not only has a direct effect on individual plant fecundity but also can influence flowering time and hence indirectly affect reproductive output.     

Fruit to flower ratios and trade-offs in size and number

Summary Hermaphroditic flowering plants commonly produce many more flowers than fruits. To interpret this observation, I suggest an explanation based on trade-offs occurring between the size and the number of flowers and fruits that mature when resources are limiting. I make this suggestion concrete with the aid of a phenotypic model analysed using the ESS approach. The model includes resource allocations to male and female reproductive structures at the time of flowering and to fruit maturation. The formulation allows for non-linear relations between opportunities for fertility gain on the one hand and flower and fruit size and number on the other. Results of the model indicate that fewer fruits will be matured than flowers produced, even in the absence of previously suggested factors such as pollinator limitation and bet-hedging in the face of environmental variability. Model results emphasize the distinction between flower versus inflorescence contributions to male and female reproductive success. Analysis also shows that the fruit to flower ratio in hermaphroditic plants will be lower than that in female plants, in accord with broad taxonomic surveys. Finally, results obtained lead to empirically testable predictions about the relationship between flower size, fruit size and the fruit to flower ratio. Methods to test these predictions are discussed.     

VARIATION IN INDIVIDUAL FLOWERING TIME AND REPRODUCTIVE SUCCESS OF AGALINIS STRICTIFOLIA (SCROPHULARIACEAE)

Field studies on two populations of Agalinis strictifolia were conducted over a 3-year period to investigate the relationship between flowering time of individuals and plant size, flowering duration, flower and fruit production, fruit predation, and growth rate. Seasonal patterns of pollinator visitation were compared with those of individual flowering time, flower density, percent fruit production, and mean seeds/fruit. In general, early and middle flowering individuals (as determined by either first flowering date or peak flowering) were larger, flowered longer, and produced more flowers and fruits than late flowering individuals. Early and middle flowering individuals (based on first flowering date) also grew faster than late flowering individuals. Although early and middle flowering individuals produced more fruits, fruit predators did not damage a disproportionate number of fruits compared to late flowering individuals. Patterns of bee visitation showed no association with seasonal patterns of flower density, percent fruit production, mean seed/fruit, or individual flowering time. In populations of A. strictifolia, it would seem that biotic or environmental determinants of growth rate (hence size and reproductive success) may be more important in generating variation in individual flowering time than patterns of pollinator visitation or fruit predation.     

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.     

SIZE-DEPENDENT GENDER CHANGE IN GREEN DRAGON (ARISAEMA DRACONTIUM; ARACEAE)

Green dragon (Arisaema dracontium; Araceae) is a perennial woodland herb capable of switching gender from year to year. Small flowering plants produce only male flowers but when larger they produce male and female flowers simultaneously. Distinct male and monoecious phenotypes (referred to hereafter as plants) share a single underlying cosexual genotype. Four populations in southern Louisiana were sampled to determine frequencies and size distributions of male and monoecious plants, and to determine the relationship of plant size with male and female flower production in monoecious plants. Male plants were significantly smaller than monoecious plants and made up 34%–78% of flowering plants within populations. Flower number (average = 120) was weakly positively correlated with size. Monoecious plants produced an average of 169 flowers (90 female) and had 100% fruit set, with individual berries containing an average of 2.5 ovules and 1.3 filled seeds. Male flower number was negatively correlated, and female flower number positively correlated, with basal stem diameter. Extrapolation of regression slopes suggested that green dragon should become completely female at a size 20% larger than the largest plant observed in this study. A simple model of inflorescence development is presented to illustrate how the reproductive system of green dragon is related to that of jack-in-the-pulpit (A. tnphyllum), which exhibits a more distinct switch between male and female phenotypes.     

Consumer pressure and seed set in a salt marsh perennial plant community

Summary Seed predation can be an important determinant of plant success, but has received little attention in wetland plant communities. Here, we examine the role of flower and seed predators in limiting the seed production of the dominant perennial plants in a salt marsh plant community. Of the four perennial investigated, direct ovule loss to consumers ranged from 51 to 80%, resulting in seed set reductions ranging from 50% to over 20-fold. Most losses were due to generalist grazing by the grasshopper, Conocephalus spartinae. More species-specific losses were inflicted by planthoppers, and microlepidopteran and dipteran larval seed parasites.Insect abundance and consumer pressure on flowers and seeds increased over the early summer, peaked in the middle of July, and declined through August, and this temporal pattern was reflected in the natural consumer damage incurred by each of the marsh perennials. Juncus gerardi flowers earlier than other marsh perennials and largely escapes heavy consumer losses. Spartina patens and Distichlis spicata flower in the middle of the summer during the peak consumer activity and incur extremely heavy seed losses. Spartina alterniflora flowers late in the summer as consumer pressure is subsiding, which appears to minimize its seed loss. In addition to destroying seeds directly, consumers also markedly reduce the frequency and affect the timing of sexual expression in these plants. In particular, predation drastically reduces the frequency of male flowers, which could lead to pollen limitation of seed set.Intense flower and seed predation on these marsh perennials may be an important determinant of the success of marsh plant populations as well as a potent selective force on their flowering phenologies and reproductive effort.     

Growth schedule of Xanthium canadense: Does it optimize the timing of reproduction?

Summary The effects of nutrition on the timing of reproductive initiation of a short-day annual plant Xanthium canadense (cocklebur) were examined with the following hypotheses in mind: If the plant always follows an optimal growth schedule, low-nutrient plants will initiate reproductive growth earlier than high-nutrient plants. On the other hand, if the plant flowers in response to photoperiodic stimuli, both plants will initiate reproductive growth on the same day. The sand-culture experiment showed that high-nutrient plants flowered earlier than the low-nutrient plants, leading to rejection of the first hypothesis. The predicted optimal flowering time is 2 days later than the actual flowering time in high-nutrient plants and 10 days earlier in low-nutrient plants. These deviations from the optimal times reduced the reproductive yield by 0.1% and 2.3%, respectively. The ratio of the final reproductive yield to the vegetative mass at flower initiation was 1.10 in high-nutrient plants and 0.63 in low-nutrient plants. Since the expected ratio for the optimal growth schedule is 1.0, high-nutrient plants followed the opitmal growth schedule more closely than the low-nutrient plants. Cocklebur is a fast-growing annual which is common in relatively nutrient-rich environments. This study suggests that cocklebur adapts itself to such environments through its photoperiodic response.     

Refuse attracts? Effect of refuse dumps of leaf‐cutting ants on floral traits

The nutrient‐rich organic waste generated by ants may affect plant reproductive success directly by enhancing fruit production but also indirectly, by affecting floral traits related with pollinator attraction. Understanding how these soil‐nutrient hot spots influence floral phenotype is relevant to plant–pollination interactions. We experimentally evaluated whether the addition of organic waste from refuse dumps of the leaf‐cutting ant Acromyrmex lobicornis (Hymenoptera: Formicidae: Attini) alters floral traits associated with pollinator attraction in Eschscholzia californica (Ranunculales: Papaveraceae), an entomophilous herb. We analysed flower shape and size using geometric morphometric techniques in plants with and without the addition of refuse‐dumps soil, under greenhouse conditions. We also measured the duration of flowering season, days with new flowers, flower production and floral display size. Plants growing in refuse‐dumps soil showed higher flower shape diversity than those in control soil. Moreover, plants in refuse‐dumps soil showed bigger flower and floral display size, longer flowering season, higher number of flowering days and flower production. As all these variables may potentially increase pollinator visits, plants in refuse‐dumps soil might increase their fitness through enhanced attraction. Our work describes how organic waste from ant nests may enhance floral traits involved in floral attraction, illustrating a novel way of how ants may indirectly benefit plants.     

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.     

Flowering phenology,pollen flow and fruit production in the andean shrub Befaria resinosa

Summary In the eastern Andes of Colombia, the shrub Befaria resinosa (Ericaceae) has peaks of flowering that are separated by extended periods of low flower production. The effect that these fluctuations in flower production have on pollen flow was investigated by using fluorescent dye as a pollen analog. Dye applied to open flowers was dispersed over long distances more often during low flower production than during high flower production. Whether enhanced pollen dispersal during flowering lows is of benefit to individual plants is not clear. The proportion of flowers that set fruit is positively correlated with flower abundance, negating the possibility that increased pollen dispersal results in a higher rate of fruit production due to outbreeding effects. It is also difficult to attribute the pattern of fruit production to changes in pollinator visitation rates, which are negatively correlated with flower abundance in the case of hummingbirds and not correlated at all with flower abundance in the case of insects. An opportunistic, large-bodied hummingbird (Colibri coruscans) visits B. resinosa during high flowering and may be a particularly effective pollinator, accounting for some of the increase in the proportion of flowers setting fruit. Rainfall is positively correlated with flower production and may be an important factor in shaping flowering phenology, but it is not significantly correlated with the proportion of flowers setting fruits. The possibility that low-level flowering may counteract inbreeding that results from peak flowering is discussed.     

Flowering phenology and reproductive success of native shrub Berberis darwinii Hook. along different light environments

Studies evaluating flowering phenology and reproductive success are necessary when we want to direct a domestication project in a species with a potential productive value. We studied flowering phenology and reproductive success of Berberis darwinii growing under different light conditions in its native distribution area in the Andean Patagonian forests of Argentina. We test the hypothesis that plants grown under conditions of high-light availability exhibit advanced phenology and higher reproductive success than those grown under conditions of lower light availability. Phenology and reproductive success were determined in three contrasting light conditions at two forest sites, which were, canopy, gap and forest edge. Plants did not bloom under the forest canopy. Flowering and fruiting period lengths were similar in both sites and light conditions of gap and forest edge during spring and summer. Although gap plants had more racemes per shoot, racemes of edge plants had more flowers, fruits and a higher proportion of flowers producing ripe fruit. We show that B. darwinii reproduction studied in the Andean Patagonian forests is conditioned by the canopy openness. Regarding reproductive success, edge plants invest less resources in flower production than gap plants to have similar fruit production.     

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%).     

PATTERNS OF FRUIT PRODUCTION IN A NEOTROPICAL ORCHID: POLLINATOR VS. RESOURCE LIMITATION

Patterns of growth, and of flower and fruit production, were monitored over two years in a population of the orchid Aspasia principissa Reichb. f. in central Panama. Observations and experimental manipulations were used to determine the relative importance of pollinator and resource limitation on fruit production. Within a season, fruit production was limited by pollinator availability. Fruit set for hand-pollinated flowers was over six times greater than that for naturally pollinated flowers. However, in plants that produced more than one fruit, fruit size declined in subsequent fruits, indicating that resources could limit seed production within a season. Plants producing fruits in 1986, on average, produced smaller shoots and inflorescences in 1987 relative to plants that flowered but produced no fruit in 1986. Thus, plants are likely to be resource limited over their lifetimes. Most individuals of reproductive size (82.5%) did not produce fruit over a three-year period. The reproductive dominance of a few individuals in this population of Aspasia principissa may have important implications for understanding the population structure of the species and the high species diversity of orchids.     

FACTORS LIMITING FRUIT AND SEED PRODUCTION OF A TEMPERATE SHRUB,STAPHYLEA TRIFOLIA L. (STAPHYLEACEAE)

Staphylea trifolia L., the bladdernut, is a self-incompatible temperate woodland shrub that flowers in May in Illinois. Factors limiting reproduction were studied at four levels: 1) Seeds/fruit. Seed production in open-pollinated fruit was frequently limited by too few fertilized ovules. Seed production in hand-cross-pollinated fruit was limited by resources or dispersal constraints: seed abortion rates were higher in hand-cross-pollinated fruits than in open-pollinated fruits. 2) Fruits/flower and 3) fruits/inflorescence. The number of fruits set and matured per flower and per infloresence in the open-pollinated treatment was limited by the number of flowers naturally cross-pollinated. In hand-cross-pollinated inflorescences, fruit set was not limited by resources even though fruit set was ten times greater than in the open-pollinated treatment. Evidence that resources limited fruit maturation in the hand-cross-pollinated inflorescences was equivocal. In hand-cross-pollinated flowers, fruit set was lowest when cold nights followed pollination, suggesting that cool temperatures limited postpollination physiological processes. 4) Fruits/individual. Early-flowering individuals matured fewer fruits than later-flowering individuals. Within the latter group, fruit production increased with plant size, although a relatively small individual matured the maximum number of fruits. Flowering phenologies and size of individuals varied among patches, resulting in differential reproductive success of patches.     

Flowering phenology and female fitness: impact of a pre-dispersal seed predator on a sexually polymorphic species

In order to produce seeds, animal-pollinated plants must flower synchronously with and be attractive to their pollinators while avoiding antagonists. Here, we explore temporal and inter-individual patterns in pollination and pre-dispersal seed predation of Dianthus sylvestris by Hadena moths, within and among three sex morphs. We scored plants that started flowering at different periods in 2001 and 2003 and found that fruit set decreased and predation rates increased over one season, and most of the other season, granting a female reproductive advantage to early flowering plants, though, we found no morph-specific temporal patterns. Female plants set more fruits, and more of their fruits escaped predation in one year, but this did not grant them a reproductive advantage since they produce fewer flowers per plant than the other morphs. Instead, mixed plants showed a clear female reproductive advantage. We also examined predation types by Hadena and seed production in attacked and intact fruits of individually marked flowers. Though female Hadena moths laid eggs preferentially into perfect flowers, flower sexes suffered similar predation by itinerant caterpillars. Attacked fruits contained fewer and lighter seeds than un-attacked ones. We conclude that pre-dispersal seed predation by Hadena may select on flowering onset of this sexually polymorphic species.     

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

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

The roles of female and hermaphroditic flowers in the gynodioecious–gynomonoecious Silene littorea: insights into the phenology of sex expression

Some gynodioecious species have intermediate individuals that bear both female and hermaphroditic flowers. This phenomenon is known as a gynodioecious–gynomonoecious sexual system. Gender expression in such species has received little attention in the past, and the phenologies of male and female functions have also yet to be explored. In this study, we examined variations in gender patterns, their effects on female reproductive success and sex expression in depth throughout the flowering period in two populations. The studied populations of Silene littorea contained mostly gynomonoecious plants and the number of pure females was very low. The gynomonoecious plants showed high variability in the total proportion of female flowers. In addition, the proportion of female flowers in each plant varied widely across the flowering season. Although there was a trend towards maleness, our measures of functional gender suggested that most plants transmit their genes via both pollen and ovules. Fruit set and seed set were not significantly different among populations; in contrast, flower production significantly varied between the two populations – and among plants – with consequent variation in total seed production. Conversely, gender and sex expression were similar in both populations. Plants with higher phenotypic femaleness did not have higher fruit set, seed set or total female fecundity. The mating environment fluctuated little across the flowering period, but fluctuations were higher in the population with low flower production. We therefore conclude that the high proportion of gynomonoecious individuals in our studied populations of S. littorea may be advantageous for the species, providing the benefits of both hermaphroditic and female flowers.