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.     

Crab spiders (Thomisidae) attract insect flower‐visitors without UV signalling

1. Crab spiders (Thomisidae) indirectly affect insect flower‐visitor and flowering plant interactions by consuming and altering the behaviour of insects. 2. Although one expects insect flower‐visitors to avoid crab spiders actively, some crab spider species are known to attract flower‐visitors. Crab spiders may use UV signalling to lure potential prey to the flowers they occupy. 3. In the present study, a field experiment was conducted to examine the effects of crab spiders occupying three prairie plant species for the insect flower‐visitor community. Pollinating insects were significantly attracted to inflorescences with crab spiders compared to inflorescences without crab spiders for two plant species, and herbivorous insects were attracted to inflorescences with crab spiders for one of these plant species. The two flowering plant species with increased pollinator visitation showed increased seed weights for plants with crab spiders, indicating crab spider presence indirectly increased pollination. 4. To test the UV signalling hypothesis, inflorescences with crab spiders of one plant species were observed under both a UV‐blocking plastic and a clear plastic control. Contrary to our prediction, flower‐visitors were not more likely to land on inflorescences under the clear plastic; the UV signalling hypothesis was not supported. Other unknown explanations underlie prey attraction to inflorescences with crab spiders.     

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.     

The mating system of Hydrophyllum appendiculatum,a protandrous species

Summary The mating system of Hydrophyllum appendiculatum (Hydrophyllaceae), a protandrous, self-compatible, monocarpic plant was examined using progeny arrays assayed at three polymorphic allozyme loci. We were particularly interested in the effect of ecological factors on spatial and temporal variation in outcrossing rates. Multilocus estimates of outcrossing rates in three populations ranged from 0.62–0.81 indicating that the majority of seeds are produced via outcrossing. The population estimates did not differ significantly from each other indicating that there is little or no spatial variation in the mating system of this species. The estimates were, however, significantly less than unity, which demonstrates that although mainly outcrossing, a significant fraction of seeds are produced by selfing. Estimates suggested that biparental inbreeding occurred, although it was statistically significant in only one population. Individuals of H. appendiculatum may remain in anthesis 3–4 weeks and produce up to 30 inflorescences. As a result, the possibility exists for the mating system to vary through the flowering season. Although the mean outcrossing rate was highest in the middle of the flowering phenology than at the beginning or end, there was no evidence for statistically significant temporal heterogeneity. We were also interested in determining if the size of the floral display (number of inflorescences produced by a plant) influenced the outcrossing rate. The results obtained by two different statistical analyses were contradictory; there was a significant positive correlation between size of floral display and outcrossing rate, but the outcrossing rates of large plants (\s> 8 inflorescences) did not differ significantly from small plants (> 8 inflorescences).     

Should food‐deceptive species flower before or after rewarding species? An experimental test of pollinator visitation behaviour under contrasting phenologies

Many plant species reward their pollinators, whereas some species, particularly among orchids, do not. Similarity of floral cues between co‐flowering species influences how rapidly pollinators learn to avoid deceptive plants. This learning process, which affects the reproductive success of deceptive plants, may additionally depend on relative timing of flowering of sympatric rewarding and deceptive species. We tested the combined effects of corolla colour similarity and flowering order of rewarding and deceptive artificial inflorescences on visitation by naïve bumblebees. When deceptive inflorescences were offered after rewarding inflorescences, bumblebees visited them four times more often if both species were similar compared with when they were dissimilar. Pollinator visitation rate to deceptive inflorescences offered before rewarding inflorescences was intermediate and independent of similarity. Thus, early‐flowering deceptive species avoid the costs of dissimilarity with rewarding species. This mechanism may favour adaptive evolution of flowering phenology in deceptive species and explain why temperate deceptive orchids usually flower earlier than rewarding ones.     

A Developmental Pattern of Flowering in Colored <Emphasis Type="Italic">Zantedeschia</Emphasis> spp.: Effects of Bud Position and Gibberellin

The restricted flowering of colored cultivars ofZantedeschia is a consequence of developmental constraints imposed by apical dominance of the primary bud on secondary buds in the tuber, and by the sympodial growth of individual shoots. GA₃ enhances flowering inZantedeschia by increasing the number of flowering shoots per tuber and inflorescences per shoot. The effects of gibberellin on the pattern of flowering and on the developmental fate of differentiated inflorescences along the tuber axis and individual shoot axes were studied in GA₃ and Uniconazole-treated tubers. Inflorescence primordia and fully developed (emerged) floral stems produced during tuber storage and the plant growth period were recorded. Days to flowering, percent of flowering shoots and floral stem length decreased basipetally along the shoot and tuber axes. GA₃ prolonged the flowering period and increased both the number of flowering shoots per tuber and the differentiated inflorescences per shoot. Activated buds were GA₃ responsive regardless of meristem size or age. Uniconazole did not inhibit inflorescence differentiation but inhibited floral stem elongation. The results suggest that GA₃ has a dual action in the flowering process: induction of inflorescence differentiation and promotion of floral stem elongation. The flowering pattern could be a result of a gradient in the distribution of endogenous factors involved in inflorescence differentialtion (possibly GAs) and in floral stem growth. This gradient along the tuber and shoot axes is probably controlled by apical dominance of the primary bud. Online publication: 7 April 2005     

Resource-related variables drive individual variation in flowering phenology and mediate population-level flowering responses to climate in an asynchronously reproducing palm

Many tropical plant species show wide intra-population variation in reproductive timing, resulting in the protracted presence of flowering and fruiting individuals. Various eco-evolutionary drivers have been proposed as ultimate causes for asynchronous phenology, yet little is known about the proximate factors that control reproductive onset among individuals or that influence the proportion of trees producing new inflorescences within a population. We employed a nine-year phenological record from 178 individuals of the hyperdominant, asynchronously flowering canopy palm, Oenocarpus bataua (Arecaceae)¸ to assess whether resource-related variables influence individual- and population-level flowering phenology. Among individuals, access to sunlight increased rates of inflorescence production, while the presence of resource sinks related to current investment in reproduction—developing infructescences—reduced the probability of producing new inflorescences. At the population level, climate anomalies induced by El Niño Southern Oscillation (ENSO) affected the proportion of the population producing inflorescences through time. Moreover, the effects of ENSO anomalies on flowering patterns depended on the prevalence of developing infructescences in the population, with stronger effects in periods of low developing-infructescence frequency. Taken together, these results suggest that resource-related variables can drive phenological differences among individuals and mediate population-level responses to larger-scale variables, such as climate anomalies. Consequently, a greater focus on the role of resource levels as endogenous cues for reproduction might help explain the frequent aseasonal phenological patterns observed among tropical plants, particularly those showing high intra-population asynchrony.     

Reproductive biology of the rare and endangered Banksia brownii Baxter ex R. Br. (Proteaceae)

Abstract Banksia brownii is an endangered species, now limited to ～ 15 disjunct populations in southwestern Western Australia. Data on flowering phenology, plant size, fruit set, pollination and the mating system were gathered for two of these populations between March and October 1993. Flowering for both populations followed a similar pattern, with open flowers first evident in April, and the number of inflorescences with open flowers peaking in June. At both locations, trees differed considerably with respect to their size, the total number of inflorescences produced and the length of their flowering season. Fruiting success was typically low, with approximately half of all inflorescences failing to develop into infructescences. Only 1. 8% of the flowers originally present on inflorescences developed into follicles. The distribution of follicles along each infructescence was non-random, with most forming in the middle third of the infructescence for reasons relating to nutrient supply and pollinator behaviour. More flowers opened during the day than at night, although pollen was lost from individual flowers during both periods. Honeyeaters such as Phylidonyris novaehollandiae were common at the two study sites, and often carried large loads of B. brownii pollen. Though less frequently caught, the nocturnal mammals Rattus fuscipes and Tarsipes rostratus also bore substantial amounts of pollen. Most inflorescences from which these mammals and birds were excluded remained barren. Fruiting success was further reduced when invertebrates such as Apis mellifera were also prevented from visiting inflorescences. The ability of B. brownii to set at least some fruit in the absence of biotic poli-nators indicates that the species is partially self-compatible. Honeyeaters foraged preferentially at inflorescences with one to two thirds of their flowers open, probing mainly along the ‘advancing front’ of open flowers. These animals moved more frequently between inflorescences on the same plant than between those on different plants, and were often recaptured in the same locations. Mammals also appeared to be sedentary. Both B. brownii populations had mixed mating systems, with genetically determined outcrossing rates of ～0.7. The unusually high level of selfing in each population is presumably a reflection of the species’ self-compatibility and the foraging behaviour of its pollinators.     

The pollination ecology of Actaea spicata (Ranunculaceae)

Pollination ecology of Actaea spicata L. (Ranunculaceae) was studied at localities ranging from deciduous to spruce forest. The species is self–incompatible and possibly facultatively apomictic. No anemogamy seems to occur. In deciduous forest, the plant is primarily pollinated by pollen–foraging and mate–searching Byturus ochraceus (Scriba) (Coleoptera). It flowers during a period between emergence of the beetle and onset of anthesis of the prime pollen source and only larval host plant, Geum ur–banum L. (Rosaceae). The beetles sit on the suitably long anther filaments or on top of the stigma. Floral morphology promotes pollination–performing movements by the beetle. The complex floral fragrance seems to induce close–range behaviour, i.e. alighting and/or foraging. Flies become abundant visitors only shortly before anthesis is over, and their contribution is small. Large syrphids are relatively effective as pollinators, since they grasp the stamens near the base and often touch the stigma with pollen–dusted parts of their bodies. Due to lack of its food–plant, B. ochraceus is absent at spruce forest localities and northwards. At these localities flowering is delayed until the main fly appearance, and the flowers are heavily pollinated by several syrphids and muscids. At three deciduous–forest localities a second flowering took place with new, juxtaposed inflorescences on separate stalks. The distribution of such inflorescences within the population along with the distribution and abundance of pollinators among the two generations of flowers suggest that these inflorescences occur as a response to low rate of pollination in the first anthesis.     

Variation in sexual expression in the monoecious shrub Buxus balearica at different scales

Monoecy allows high plasticity in gender expression because the production of separate female and male flowers increases the ability to respond to specific environmental circumstances. We studied variation in sexual expression and its correlates in the monoecious shrub Buxus balearica, for two years, in six populations in the Balearic Islands and four in the Iberian Peninsula. Phenotypic gender varied among populations; while island populations showed slight variations around an average gender, mainland populations showed a broad range of variation in gender among individuals, always biased towards increasing maleness compared with the other populations. Within populations, gender was not related to plant size. Between-year changes were slight and mainly consisted of an increase in relative maleness in the mast year. Reproduction did not affect gender in the next year, as assessed by either observational or experimental methods. Most variation in gender expression occurred among individuals within populations (83.6 %), followed by variation among populations (13.6 %) and years (2.8 %). Our results suggest that male-biased gender at population and plant levels was related to stressful conditions and resource limitation, because: (1) maleness was higher in mainland populations, where summer drought was stronger; (2) maleness increased with elevation; (3) fruit set was positively correlated with femaleness; (4) the percentage of male inflorescences increased over the flowering period; and (5) male inflorescences were preferentially in lower parts of the branch nodes. Higher maleness in mast years, however, could be related to increased male success under synchronic flowering.     

Variation in multiple traits of vegetative and reproductive seagrass tissues influences plant–herbivore interactions

Plant–herbivore interactions have strong ecological and evolutionary consequences, but have been traditionally overlooked in marine higher plants. Despite recent advances in seagrass ecology that highlight the importance of herbivory, the mechanisms that regulate the feeding behaviour of seagrass consumers remain largely unknown. Herbivores have been shown to reduce the sexual reproductive success of seagrasses through direct consumption of inflorescences and seeds, but we know little about intraspecific variation in susceptibility to grazing of different seagrass tissues. We contrasted the relative palatability of reproductive and vegetative tissues of the temperate seagrass Posidonia oceanica in the field, and we assessed the feeding preferences among these tissues of the main consumers of the plant, the fish Sarpa salpa and the urchin Paracentrotus lividus. Moreover, we identified the plant traits that explained the observed feeding behaviour. We provide strong evidence for herbivore selectivity among seagrass tissues. In the field, 70–90% of inflorescences were damaged by herbivores compared to 3–60% of leaves of similar age. In feeding assays, the urchin P. lividus showed over a twofold preference for reproductive tissue at various stages of development. By contrast, we detected no feeding activity on either leaves or inflorescences from the fish S. salpa, which is known to migrate to deeper waters soon after flowering starts and during the period of fruit maturation. Despite being the preferred food of urchins, inflorescences were chemically defended, had higher levels of phenolics and lower nutrient and calorific content than leaves. We experimentally demonstrated that leaf structural defences are the primary factor in determining urchin feeding preferences. Removal of plant structure results in a drastic shift in urchin selectivity towards the most nutritious and less chemically defended leaf tissue, indicating that multiple mechanisms of defence to herbivory may coexist in seagrasses.     

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.     

The inheritance of partial self-compatibility inBrassica oleracea L.: results from a half diallel homozygous for a highly recessive S-allele

Summary In a study of partial self-compatbility inBrassica oleracea, flower number, seeded siliqua and seed production were recorded on self- and cross-pollinated inflorescences of the progenies of a half diallel between five unrelated inbred plants homozygous for the same recessiveS-allele.On cross-pollinated inflorescences significant amounts of additively controlled genetic variation were found for seed set per flowering site and its two components, seeded siliquae per flower and seeds per seeded siliquae. Considerable heterosis and gene interaction were also present and a simple additive dominance model did not fully explain the observed variation.On self-pollinated inflorescences, additive gene action was absent for the seed production variates although differences between progenies were highly significant and heterosis was present. Complex gene interactions were considered to be responsible. The characteristics of the method of assessment used and the relationship between self- and outcross seed production are discussed. It appears unlikely that the component lines could be selected for reduced self seed set, but selection for higher outcross seed set may be possible.     

Distribution of plant‐dwelling spiders: Inflorescences versus vegetative branches

Abstract The influence of the architecture of vegetative branches on the distribution of plant‐dwelling spiders has been intensively studied, and the effects on the aggregation of individuals in several spider species on plants include variation in prey abundance, availability of predator‐free refuges and smoother microclimate conditions. The emergence of inflorescences at the reproductive time of the plants changes branch architecture, and could provide higher prey abundance for the spiders. The distribution of spiders between inflorescences and vegetative branches was compared on four widespread plant species in a Brazilian savannah‐like system. Inflorescences attracted more spiders than vegetative branches for all plant species sampled. The influence of branch type (inflorescence and vegetative) on spider distribution was also evaluated by monitoring branches of Baccharis dracunculifolia DC. in vegetative and flowering periods for 1 year, and through a field experiment carried out during the same period where artificial inflorescences were available for spider colonization. Artificial inflorescences attached to B. dracunculifolia branches attracted more spiders than non‐manipulated vegetative branches for most of the year. However, this pattern differed among spider guilds. Foliage‐runners and stalkers occurred preferentially on artificial inflorescences relative to control branches. The frequencies of ambushers and web‐builders were not significantly different between treatment and control branches. However, most ambush spiders (65%) occurred only during the flowering period of B. dracunculifolia, suggesting that this guild was influenced only by natural inflorescences. The experimental treatment also influenced the size distribution of spiders: larger spiders were more abundant on artificial inflorescences than on vegetative branches. The hypothesis that habitat architecture can influence the spider assemblage was supported. In addition, our observational and experimental data strongly suggest that inflorescences can be a higher quality microhabitat than non‐reproductive branches for most plant‐dwelling spiders.     

Influence of plant species and plant growth stage on Frankliniella occidentalis pupation behaviour in greenhouse ornamentals

The influence of plant stage and plant species on the pupation behaviour of western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), was investigated in flowering and non‐flowering potted chrysanthemums and mini roses. On non‐flowering chrysanthemums and roses, 92–93% of the F. occidentalis pupated in the soil. On flowering roses, 87% pupated in the soil, and on flowering chrysanthemums only 60% chose the soil as a pupation site and 40% stayed on the plant. This means that, in the presence of flowers, especially complex inflorescences like chrysanthemums, a large proportion of F. occidentalis chooses stay on the plant to pupate.     

Significance of intra-inflorescence variation on flowering time of a spring ephemeral, <Emphasis Type="Italic">Gagea lutea</Emphasis> (Liliaceae), under seasonal fluctuations of pollinator and light availabilities

I studied the relationship between seed-set patterns within inflorescences and temporal variations in light and pollinator availabilities for 2 years in the spring ephemeral species Gagea lutea in a deciduous forest. Timing of canopy closure and seasonal trend of pollinator frequency did not synchronize with the annual fluctuation in flowering phenology. In the early snowmelt year, seed-set success reflected the seasonal pollinator abundance from early to middle flowering periods. In the late snowmelt year, however, seed-set rates were independent of pollinator activity and decreased with canopy closing even after hand pollination. The restricted seed production by defoliation and the increase in seed-set rates at the forest edge suggested that seed production was supported by current photosynthetic carbon gain. Thus, annual fluctuations of reproductive success can explain the variation in flowering phenology within a population although seasonal light deterioration would serve as a selective force for flowering in the early season.     

Studies on the Growth, Flowering, and Production of Female Sterile Flowers as Affected by Different Levels of Foliar Potassium in Solanum sisymbrifolium Lam.: I. EFFECT OF POTASSIUM CONTENT OF THE PLANT ON VEGETATIVE GROWTH AND FLOWERING

As compared to a high level of foliar potassium, a low but nota deficient level promotes shoot elongation and flowering inS.sisymbrifolium.This is accompanied by formation of flowers with rudimentaryovaries in which the megagametophyte in the ovules aborts atthe two- or the four-nucleate stage. Plants having a higherK content do not bear the female sterile flowers. Inflorescencesof the high-K plants with 20 per cent fewer buds have 23 percent higher dry weight than inflorescences of low-K plants.The ratio of organic nitrogen to dry weight is not affected,indicating that the inflorescences of the high-K plants arebetter supplied with metabolites in general.     

Relationship between floral longevity and sex allocation among flowers within inflorescences in Aquilegia buergeriana var. oxysepala (Ranunculaceae)

Understanding the fitness of plants with inflorescences requires examining variation in sex allocation among flowers within inflorescences. We examined whether differences in the duration of the male and female phases of flowering lead to variation in sex allocation and reproductive success among flowers within inflorescences. In 2002 and 2003, we quantified floral longevity, floral sex allocation, and reproductive success between the first and the second flowers within inflorescences in a protandrous species, Aquilegia buergeriana var. oxysepala. Floral longevity was greater in the first flowers than in the second ones in both years. The male phase lasted longer, and the initial number of pollen grains and the number of pollen grains removed were greater in the first flowers than in the second ones in both years. Within first flowers, the number of pollen grains removed was greater in flowers that had longer male phases, thus duration of the male phase may positively affect male reproductive success in the first flowers. The female phase lasted longer and the number of ovules was greater in the first flowers than in the second only in 2002. However, seed production per flower and female phase duration in both years were not significantly related. The variation in the number of pollen grains among flowers in this species may be caused by the variation in male phase duration.     

The influence of fire on the phenological behaviour of Mediterranean plant species in Bas-Languedoc (southern France)

Fire is an ancient ecological factor influencing the Mediterranean vegetation of southern France. The study was carried out on three areas to determine the phenological behaviour of plants with regard to fire. First we studied the flowering responses of perennials in relation to the time since fire: in a Quercus coccifera garrigue most species flower during the year following burning. In comparing species by species between burned and unburned areas most species did not show major differences in the phenological stages. However, fire did increase the number of inflorescences of grasses. A phenological synthesis showed that differences at the community level existed for the flowering stages between the burned areas and the unburned control sites during the first and second years following fire. The growth of some woody species was also studied; the elongation and growth of the plants were biggest during the first or second year after fire. The lack of differences in phenological response between burned and unburned plants may be an adaptive trait to fire.     

Effect of pollen limitation, additional nutrients, flower position and flowering phenology on fruit and seed production in Salvia verbenaca (Lamiaceae)

Variation in fruit set, seed/ovule ratio and mean single-seed weight among plants, and within plants among inflorescences, was investigated in a population of Salvia verbenaca in northwest Spain. Mean single-seed weight varied significantly among plants which flowered at about the same time and mean single-seed weight was significantly greater for plants that flowered late in the season than for plants flowering early in the season. Within individual plants, mean single-seed weight increased steadily between the earlier opening basal inflorescences and the later opening upper inflorescences. However, neither fruit set nor seed/ovule ratio varied significantly among inflorescences. Previous work on this population indicates that outcrossing improves mean single-seed weight, so I suggest that the observed pattern of among-inflorescence variation in mean single-seed weight may be attributable to a gradual increase in nonself pollen receipt as the flowering season progresses.