First record of Melampodium divaricatum (Asteraceae) in West Tropical Africa

A plant that showed morphological closeness to Aspilia africana (Pers) C. D. Adams (Asteraceae) was spotted and collected in 2015 along Afe Babalola University road, Ado‐Ekiti, Ekiti State, Nigeria with coordinates 7°36′59.99″N, 5°12′60.00″E. However, upon closer observation some distinct and peculiar characteristics that clearly distinguished it from Aspilia africana were revealed, e.g. sterility of the disc florets and production of achenes by ray florets only. Another striking character of the plant was total emptying of the capitulum after achene maturation, leaving an empty capitulum cup on the plant. Literature and herbarium searches revealed that the plant had neither been reported from West Tropical Africa nor collected in any herbarium in Nigeria before. The plant was eventually identified as Melampodium divaricatum (L.) which is an annual erect herb, distributed in tropical and subtropical regions but mostly restricted to Mexico, North America and Central America. Morphological, reproductive and cytological studies carried out on the plant revealed it to possess a highly branched erect pigmented stem, simple opposite sub sessile leaves with acute apex and distantly serrated margins, capitula with yellow unisexual disc and ray florets, sterile disc florets, fertile ray florets, relatively high pollen fertility (92.85%), a somatic chromosome number of 2n = 24 and regular formation of 12 bivalents, indicating the plant to be a diploid species. Further studies on Melampodium in Nigeria and a general revision of the flora of West Tropical Africa is suggested as well as the need to monitor M. divaricatum in the region since it appears to have the capacity to become invasive.     

Differential outcrossing rates in dispersing and non-dispersing achenes in the heterocarpic plant Crepis sancta (Asteraceae)

The association between mating systems and dispersal in plants has been studied mostly in cleistogamous species where, generally, seeds produced by cleistogamous (selfed) flowers are less dispersed than seeds produced by chasmogamous (potentially outcrossed) flowers. In heterocarpic Asteraceae, non-dispersing fruits (achenes) are produced at the periphery of the capitulum (outer florets) whereas dispersing achenes are produced by inner florets in the same capitulum. Since all the florets are protandrous, the outer floret developing first are in female phasis when anthesis of inner florets takes place. Thus, outer florets can be potentially selfed by the inner florets of the same capitulum whereas the latter must be pollinated by flowers of other capitula. Therefore outer florets should be more inbred than inner florets. To test this hypothesis, we measured the natural outcrossing rate in outer and inner florets using allozymes in three populations of the heterocarpic Crepis sancta. The results showed that the outcrossing rate was highest for non-dispersed achenes. Moreover, among the outcrossed achenes within a capitulum it was observed that the number of paternal parents of non-dispersing achenes was higher than for dispersing achenes. The pattern observed was therefore the opposite to the pattern of cleistogamous plants and contradicts the putative pollination mechanism we proposed for Asteraceae. The results agree with the predictions of sib competition theory which considers that outcrossing may minimize competitive interactions among relatives (sibs) falling near the mother plant. Higher outcrossing rate in outer florets could also occur because pollinators are more attracted to these florets.     

Variation in floral morphology within a population of Aster hispidus var. tubulosus (Asteraceae,Astereae)

The family Asteraceae has a particular inflorescence, the capitulum, consisting of ray florets and disc florets. The ray florets function as petals that attract pollinators. Marked variation in the ray floret morphology is known in a natural population of Aster hispidus var. tubulosus (Asteraceae). We analyzed the variation and found two distinct types in the ray florets, the long tubular ray floret and the ligulate ray floret. In this species, therefore, the variation in floral morphology among capitula, each of which is the basic pollination unit, is caused by the variation in the composition of the two ray floret types among capitula. We evaluated the sources of the observed variation in the floral morphology among capitula within a population using a hierarchical analysis that separated within‐individual (i.e. among capitula within each individual) and between‐individual components of the variation. We found that the main source of the variation lay at the between‐individual level, not at the between‐capitulum level nested within individuals. This finding will provide the basic knowledge that enables future study exploring whether the between‐individual variation in floral morphology caused by the compositional variation of the ray floret types leads to differential pollination success of individual plants in species of Asteraceae.     

Factors Contributing to Variation in Seed Production among Remnant Populations of the Endangered Daisy Gerbera aurantiaca1

The species‐rich mist‐belt grasslands of southern Africa have been severely reduced in extent as a result of commercial afforestation, thus confining many endemic plants and animals to small populations in habitat fragments. We investigated the influence of various environmental factors on seed production and seedling recruitment in remnant populations of the endangered grassland herb Gerbera aurantiaca (Asteraceae). Experiments with color traps showed that Eriesthis beetles, which appear to be the primary pollinators of G. aurantiaca, were most abundant in the two largest y extant populations. Seeds are produced in a very small proportion (typically <10%) of the ca 80 female florets in a capitulum. The mean number of seeds produced in undamaged capitula was found to be significantly lower in small than in large populations. Pollen limitation was evident from a significant overall increase in seed set after supplementalpollination in three populations over two seasons. The proportion of capitula containing seed predators did not differ markedly among populations or years, but lepidopteran larvae, which destroy all of the seeds in a capitulum, weremost abundant in the two largest populations in 2003. The presence of juvenile plants varied markedly amongpopulations, but this could not be linked firmly to estimates of seed production. Clonal growth is likely to contribute to the persistence of small isolated populations of G. aurantiaca, even when seed production is severely compromised by pollen limitation and predation.     

Style morphological diversity of some Asteraceae species from Argentina: systematic and functional implications

Stylar micromorphological diversity of 42 Asteraceae species from Argentina was analysed considering species phylogenetic membership and some floral reproductive functions (pollen presentation and pollen reception). In particular, the morphology and organisation of pollen presenter (sweeping hairs) and pollen receptive structures (stigmatic papillae) were described. Results showed that style morphology of the studied species is far more diverse than the categories previously established for Asteraceae, and that it is problematic to relate the sweeping-hair arrangement of species to the only three modes of pollen presentation described for the family, indicating that the hypothesised relationship could be more complex than was formerly thought. For all species with di- or trimorphic florets, the styles of female florets were more slender and without or with more reduced sweeping hairs than the styles of hermaphrodite florets, and divergences of sweeping hair arrangements and morphology were higher among phylogenetically related species. These results suggest that functional aspects of floral morphology seem to be more important than phylogenetic constraints as selective forces determining stylar pollen presentation structures. In contrast, stigmatic-area organisation as well as the morphology of stigmatic papillae remain identical between female and hermaphrodite florets and among phylogenetically related species. Thus, stigmatic papilla morphology seems to be a phylogenetically constrained character in the studied species. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

INTRASPECIFIC VARIATION IN POLLEN YIELD IN BROMEGRASS (POACEAE: BROMUS)

Variation in pollen production was measured within five hermaphrodite species of bromegrass (Bromus). Anther length is an excellent predictor of pollen production in this genus (R² = 0.97). Anther length varied considerably within each of the species, both among and within individual plants. Within plants, most of the variation occurred among florets within spikelets; florets in upper spikelet positions were smaller and produced less pollen. In B. inermis, pollen production was decreased by defoliation and increased in shoots that grew on thatching ant (Formica obscuripes) mounds. Whole-shoot pollen yield was determined by spikelet number, number of florets per spikelet, and pollen production per floret. All of these yield components must be considered in attempts to estimate pollen production accurately.     

Location of the stigmatic areas in Mutisia speciosa Aiton ex Hook. a new floral feature in Asteraceae

In Mutisieae species, the style branches are described as short and the stigmatic areas cover the inside of the style branches. As shown in preliminary observations, the Mutisia speciosa florets had long style branches (7 mm), bifurcate only at the apex (about 1.5 mm) and juxtaposed along the remaining length. The objective was to locate the stigmatic areas in M. speciosa. For this purpose, neutral red, 3% hydrogen peroxide and hand pollinations were used. For the pollinations, florets with cut apical portion of the branches (about 2 mm) and florets with intact branches were used. In the first group of florets, the pollen was deposited along the margins of the juxtaposed branches (about 5 mm), and in the second, the pollen was ventrally deposited on the bifurcated apical portion. Some of these florets were left on the plant until fruiting and others were analyzed under a fluorescence microscope to check pollen tube growth. The neutral red test defined two ventro-marginal bands fused at the apex of each style branch, consisting of papillose cells. Intense bubbling in the hydrogen peroxide test showed that only the bands are receptive. The pollinations resulted in fruit sets and growth of pollen tubes, confirming that the bands are receptive along their entire length. This result is new and indicates the need for further studies on the floral biology of tropical Asteraceae species to improve the understanding of their reproductive attributes.     

Pollen size in Hordeum L.: correlation between size,ploidy level,and breeding system

The size of pollen in the genus Hordeum (Poaceae) is correlated with ploidy level and breeding system. Generally, the pollen size increases with the ploidy level, and outbreeding species possess significantly larger pollen than inbreeders. In H. roshevitzii (2x), H. pusilplum (2x), H. murinum subsp. murinum (4x), and H. parodii (6x) pollen, heteromorphisms occur between the central and the lateral florets of the triplet. In all four taxa, pollen from florets in which the anthers are exserted is larger than pollen from florets where the anthers remain inside the floret. The biological consequences of heteromorphic pollen are discussed, and a model for the evolution of breeding strategies in Hordeum is suggested.     

Function and evolution of sterile sex organs in cryptically dioecious Petasites tricholobus (Asteraceae)

Background and Aims

Why are sterile anthers and carpels retained in some flowering plants, given their likely costs? To address this question, a cryptically dioecious species, Petasites tricholobus, in which male and female plants each have two floret types that appear pistillate and hermaphroditic, was studied. The aim was to understand the function of sterile hermaphroditic florets in females. In addition, the first examination of functions of sterile female structures in male plants was conducted in the hermaphroditic florets on males of this species. These female structures are exceptionally large in this species despite being sterile.

Methods

Differences in floret morphology between the sex morphs were documented and the possible functions of sterile sex organs investigated using manipulative experiments. Tests were carried out to find out if sterile female structures in male florets attract pollinators and if they aid in pollen dispersal, also to find out if the presence and quantity of sterile hermaphroditic florets in females increase pollinator attraction and reproductive success. To investigate what floret types provide nectar, all types of florets were examined under a scanning electron microscope to search for nectaries.

Key Results

The sterile female structures in male florets did not increase pollinator visits but were essential to secondary pollen presentation, which significantly enhanced pollen dispersal. Sterile pistillate florets on male plants did not contribute to floral display and disappeared in nearly half of the male plants. The sterile hermaphroditic florets on female plants attracted pollinators by producing nectar and enhanced seed production.

Conclusions

The presence of female structures in male florets and hermaphroditic florets on female plants is adaptive despite being sterile, and may be evolutionarily stable. However, the pistillate florets on male plants appear non-adaptive and are presumably in decline. Differential fates of the sterile sex organs in the species are determined by both the historical constraints and the ecological functions.     

Pollination and seed set in tropical wetland grasses

This study concerns 39 different grass species occurring in the wetland regions of Godavari delta in the west Godavari district (16°15′–17°30′N and 80°50′–81°55′E) of Andhra Pradesh, southern India. In each species, the sexual status of lemmas, the period of stigma receptivity, pollen/ovule ratio, pollen longevity and daily pollen release were examined. The functional sexual systems were determined after performing controlled pollinations and testing seed quality by weight and germination potential. The florets of 35 species are morphologically hermaphroditic. Among the other four species, Chionachne koenigii and Iseilema laxum are monoecious; Chrysopogon aciculatus and Pennisetum pedicellatum are andromonoecious. Examination of the sequential opening of florets and controlled pollinations revealed that C. koenigii and I. laxum are obligate outcrossers with incompatibility to geitonogamous pollen; C. aciculatus and P.pedicellatum have a high level of compatibility to xenopollen. These and another nine species were treated as predominantly outcrossing species. Yet another five species were predominantly self‐fertilized, while the remaining 21 species displayed a mixed breeding system. In a majority of the species the P/O ratios are not high and thus do not conform to the expected large P/O ratios in wind‐pollinated plants and postulated breeding systems except in a few species. The majority of these grasses shed pollen over a short period spreading from 2 am to 8 am, when usually low turbulent conditions exist, thus allowing short distance transport and restricted pollen shadows with higher concentration of pollen near the source. This appears to be the appropriate strategy for maximization of pollination in view of the short lifespan of pollen that extends mostly over 1–3 h period, and the patchy distribution of the grasses under study. The restricted pollen shadows facilitate the low levels of pollen production in the grass species under study to fertilize most or all the available ovules.     

Difference Analysis of ClCYC2-Like Genes Expression and DNA Methylation Between the Two Types of Florets in Chrysanthemum lavandulifolium

DNA methylation is an important epigenetic modification, that is involved in the regulation of gene expression and cell differentiation, and plays an important regulatory role in flower development in higher plants. There are two types of florets on the capitulum in the genus Chrysanthemum, the flower symmetry factor CYCLOIDEA (CYC) 2-like genes may be important candidate genes for determining the identity of the two types of florets. In this study, the diploid plant Chrysanthemum lavandulifolium was used as the research material, and qRT-PCR and bisulfite sequencing polymerase chain reaction (BSP) were used to identify the expression and DNA methylation pattern of CYC2-like genes in the two types of florets. Gene expression analysis showed that the six ClCYC2-like genes were significantly different in the two types of florets, and the expression levels of ClCYC2c, ClCYC2d, ClCYC2e and ClCYC2f in the ray florets were significantly higher than those in the disc florets. For the DNA methylation analysis of the three genes ClCYC2c, ClCYC2d, and ClCYC2e, it was found that the DNA methylation levels of these three genes were negative correlated with their expression levels, and the ways in which the three genes were regulated by the DNA methylation were different. It is speculated that the different DNA methylation of ClCYC2-like genes in the two types of florets may affect the differentiation and development of the two types of florets. This study provides new clues about epigenetics for the analysis of capitulum formation in Asteraceae.

     

Selective advantage of ray florets in Scalesia affinis and S. pedunculata (Asteraceae), two endemic species from the Galápagos

The presence of neuter ray florets in species within Asteraceae is generally believed to increase pollinator attraction. In the endemic Galápagos genus Scalesia (Asteraceae) a natural variation in the presence/absence of neuter ray florets is found. To evaluate whether the presence of ray florets plays a selective role on female reproductive success we chose two species of Scalesia, Scalesia affinis that carries ray florets and S. pedunculata that is rayless. On Santa Cruz Island capitula of S. pedunculata were equipped with fake ray florets while others were untouched. On Isabela Island ray florets were removed on half of the capitula of S. affinis. In S. affinis rayed capitula received more pollinators and more pollen, which resulted in a significantly higher embryo production. In S. pedunculata no effect on embryo production was found. The disagreement between the two species may correspond to a difference in visitation frequency, S. pedunculata receiving many more visit than S. affinis. Thus, ray floret development proved beneficial in pollinator-restricted localities.     

Reproductive biological characteristics potentially contributed to invasiveness in an alien invasive plant Bidens frondosa

Bidens frondosa L. (Asteraceae) is a widespread invasive weed in China. By experimental observation and bagging treatment, the reproductive biological characteristics of this species, such as phenology of flowering, floral syndrome, breeding system and seed germination characteristics, were studied to assess the association of these reproductive characteristics with invasiveness. Flowers of B. frondosa bloom from September to October every year in Ji'an city, Jiangxi province. The lifespan of a single capitulum is approximately 4 to 5 days, with 30 to 60 florets per capitulum. The capitulum diameter, anthocaulus length, floret length and width, stamen length and pistil length were 6.1, 30.9, 2.2, 0.6, 3.0 and 2.7 mm, respectively. The seed set percentage of 48.5% in the treatment of bagging flowers without emasculation suggests B. frondosa is self‐compatible; meanwhile, the percentage of 63.1% in the treatment of bagging with emasculation and manual xenogamy suggests it also is cross‐compatible. P/O ratio per capitulum of this species was 450.5, which suggests that the breeding system of B. frondosa is facultative xenogamous, and it needs pollinators to some degree. The main floral visitors were insects of Hymenoptera, Diptera and Lepidoptera. The 1000‐achenes weight was 1.97 g. The achenes emerged as a small germination peak from the fourth to fifth day, and had a high accumulated germination rate of up to 84.0% on day 20 after sowing. Our experimental findings suggested that the reproductive biological characteristics, such as the versatile mating system of self‐ and cross‐pollination, high seed production, a special method of achene dispersal and germination peak, accompanied by a high accumulated germination rate, might contribute to the invasive ability of B. frondosa.     

CAPITULUM PHYLLOTAXIS AND NUMERICAL CANALIZATION IN MICROSERIS PYGMAEA (ASTERACEAE: LACTUCEAE)

The positions at which floret primordia arise in developing capitulum buds of Microseris pygmaea D. Don have been mapped by computer-assisted light microscopy. The primordia can be assigned positions along a basic phyllotactic spiral with a divergence angle of about 137.5°. In addition, there are regular deviations from a spiral arrangement. Typically, the first 26 primordia in phyllotactic sequence are arranged in two concentric circles of 13 primordia with considerable deviations in the divergence angle and in the distances between primordia along a parastichy at positions 13 and 26. This arrangement can be simulated by geometric models that include nearest neighbor packing, together with spiral phyllotaxis. The circular arrangement of peripheral primordia at nearly equal radial distances from the center of the developing capitulum helps to explain the numerical constancy (canalization) of peripheral structures, especially the constant number of 13 inner phyllaries on heads with very different numbers of florets.     

Phenotypic Plasticity of Capitulum Morphogenesis in Microseris pygmaea (Asteraceae: Lactuceae)

Inbred populations of the annual Chilean species Microserispygmaea show phenotypic plasticity in the number of floretsper capitulum. In order to find out how this plastic variationmay arise during development, two inbred lines (A92 and C96)were grown under short day conditions. Groups of plants fromeach strain were transferred to long day conditions at about2 week intervals. In this way we introduced variation for plantsize (number of leaves per plant) at onset of flowering. Thenumber of florets per mature capitulum increased linearly withplant size. After transfer to long day conditions, plants wereharvested daily for light microscopic measurements of meristemwhole mounts. Only the first capitulum of each plant was analysed.All florets were formed after 12 d in strain C96 and after 14d in strain A92. In order to detect the effect of plant sizeon morphogenesis, we performed a multiple regression analysisof developmental parameters on time and number of leaves. Widthand height of the capitulum receptacle increased daily witha growth rate depending on the starting size. Differences inmeristem size were detected already in vegetative plants ofdifferent sizes. In contrast, there was no influence of plantsize on floret primordium size. We combined the multiple regressionmodels in one simple model for prediction of floret numbersfrom numbers of leaves per plant at onset of flowering. Predictionsof this model agree with observed relationships in both inbredlines.Copyright 1994, 1999 Academic Press Asteraceae, capitulum, inflorescence, meristem, meristic character, Microseris pygmaea, morphogenesis, phenotypic plasticity     

Comparative cyto-embryological investigations of sexual and apomictic dandelions (Taraxacum) and their apomictic hybrids

. In the autonomous apomictic Taraxacum officinale (common dandelion), parthenogenetic egg cells develop into embryos and central cells into endosperm without prior fertilisation. Unreduced (2n) megaspores are formed via meiotic diplospory, a nonreductional type of meiosis. In this paper, we describe the normal developmental pathways of sexual and apomictic reproduction and compare these with the development observed in the apomictic hybrids. In sexual diploids, a standard type of megasporogenesis and embryo sac development is synchronised between florets in individual capitula. In contrast, we observed that megasporogenesis and gametogenesis proceeded asynchronously between florets within a single capitulum of natural triploid apomicts. In addition, autonomous endosperm and embryo development initiated independently within individual florets. Parthenogenetic initiation of embryo development in outdoor apomicts was found to be temperature-dependent. Egg cells produced in natural apomicts were not fertilised after pollination with haploid pollen grains although pollen tubes were observed to grow into their embryo sacs. Both reductional and diplosporous megasporogenesis were observed in individual inflorescences of triploid apomictic hybrids. Embryo and endosperm development initiated independently in natural and hybrid apomicts.     

Differences in pollinator effectiveness of birds and insects visiting Banksia menziesii (Proteaceae)

Summary The effectiveness of nectarivorous birds, introduced honey bees and staphylined beetles as pollinators of Banksia menziesii was assessed. Staphylinids removed substantial amounts of pollen but did not deposit any onto stigmata. Abundance of beetles on inflorescences was related to the mean number of florets opening per day. Honey bees collecting pollen were more likely to effect pollination than those collecting nectar which only contacted stigmata when arriving or leaving an inflorescence. Nectar-foraging birds probed between florets 10.2±0.8 (±SE) times, contacting 8–16 stigmata during each probe. Bees visited inflorescences ten times more frequently than birds although they deposited only 25% of the pollen that birds did on stigmata. Fruit set was ten times greater on inflorescences visited by birds than on inflorescences visited by bees. Bees were capable of removing as much pollen as birds but, because of direct pollen transfer to birds when florets opened during foraging, actual removal was probably much less. Selection for floret opening during nectar foraging by birds may have resulted from pollen removal by non-pollinating animals, such as staphylinids.     

REGENERATION OF THE SUNFLOWER CAPITULUM AFTER CYLINDRICAL WOUNDING OF THE RECEPTACLE

Using the young capitulum of Helianthus annuus L., a cylindrical plug of undifferentiated receptacle tissue, 1 mm in diameter, was isolated from lateral communication with the rest of the receptacle surface by a vertical circular wound cut, while retaining continuity with the subapical meristem. Within 24 hr, active cell division was induced at the inner and outer surfaces of the wound and in the receptacle epidermis bordering the wound edges, creating a rounded rim at the top of the wound. Within 3–6 days, floral initials, spaced 133–166 μm apart appeared on the flanks of both rims and later on the top of the plug and surrounding receptacle surface. The first formed initials developed into involucral bracts or ray florets and the later ones into disc florets which were organized into contact parastichies, the number of which did not conform with the Fibonacci series. The base of the plug developed into a stem-like structure completing the regeneration of a fully formed functional capitulum. This operation was demonstrated for two sunflower cultivars and occurred in both long and short daylengths.     

Changes in Florets’ Vertical Direction within Inflorescence Affects Pollinator Behavior,and Fitness in <i>Trifolium repens</i>

Ecological interactions between flowers and pollinators greatly affect the reproductive success. To facilitate these interactions, many flowers are known to display their attractive qualities, such as scent emission, flower rewards and floral vertical direction, in a rhythmic fashion. However, less is known about how plants regulate the relationship between these flower traits to adapt to pollinator visiting behavior and increase reproduction success. Here we investigated the adaptive significance of the flower bending from erect to downward in Trifolium repens. We observed the flowering dynamic characteristics (changes of vertical direction of florets, flowering number, pollen grain numbers, pollen viability and stigma receptivity over time after blossom) and the factors affecting the rate of flower bending in T. repens. Then we altered the vertical direction of florets in inflorescence of different types (upright and downward), and compared the pollinator behaviors and female reproductive success. Our results showed that florets opened sequentially in inflorescence, and then bend downwards slowly after flowering. The bending speed of florets was mainly influenced by pollination, and bending angle increased with the prolongation of flowering time, while the pollen germination rate, stigma receptivity and nectar secretion has a rhythm of “low-high-low” during the whole period with the time going. The visiting frequency of all the four species of pollinators on upward flowers was significantly higher than that of downward flowers, and they especially prefer to visit flowers with a bending angle of 30°–60°, when the flowers was exactly of the highest flower rewards (nectar secretion and number of pollen grains), stigma receptivity and pollen germination rate. The seed set ratio and fruit set ratio of upward flowers were significantly higher than downward flowers, but significantly lower than unmanipulated flowers. Our results indicated that the T. repens could increase female and male fitness by accurate pollination. The most suitable flower angle saves pollinators’ visiting energy and enables them to obtain the highest nectar rewards. This coordination between plants and pollinators maximizes the interests of them, which is a crucial factor in initiating specialized plant-pollinator relationships.     

Pollinator visitation closely tracks diurnal patterns in pollen release

Premise

Animal-pollinated plants face a high risk of pollen loss during its transfer. To limit the negative effect of pollen losses by pollen consumption and heterospecific transfer, plant species may adjust and stratify their pollen availability during the day (i.e., “schedule” their pollen presentation) and attract pollinators in specific time frames.

Methods

We investigated diurnal patterns of pollen availability and pollinator visitation in three coflowering plant species: Succisa pratensis with open flowers and accessible pollen, pollinated mainly by pollen-feeding hoverflies; Centaurea jacea with open flowers and less accessible pollen, pollinated mainly by pollen-collecting bees; and Trifolium hybridum with closed flowers and pollen accessible only after the active opening of the flower, pollinated exclusively by bees.

Results

The three plant species differed in the peak pollen availability, tracked by the visitation activity of their pollinators. Succisa pratensis released all pollen in the morning, while pollinator activity was still low and peaked with a slight delay. In contrast, C. jacea and T. hybridum had distinct pollen presentation schedules, peaking in the early afternoon. The pollinator visitation to both of these species closely matched their pollen availability.

Conclusions

Stratifying pollen availability to pollinators during the day may be one of several mechanisms that allow coflowering plants to share their pollinators and decrease the probability of heterospecific pollen transfer.