The pollination ecology of buzz-pollinated Rhexia virginica (Melastomataceae)

We examined the function of floral traits associated with buzz pollination through studies of Rhexia virginica (Melastomataceae) in the Muskoka region of Ontario, Canada. Controlled pollinations demonstrated that the species is self-compatible, but dependent on insects for pollen transfer. Bumble bees made 82 and 90% of observed insect visits to R. virginica in 1996 and 1997, respectively, and effectively buzzed flowers. Buzz pollination did not appear to be highly "specialized" since various species of bumble bee were capable of pollination, and pollen transfer efficiency appeared to be relatively low. Experimental manipulations provided little support for the hypothesis that the yellow color of melastome anthers mimics abundant pollen, thereby deceiving pollinators to visit regardless of whether most pollen has been removed. Fruit set averaged 52.6% among populations, owing largely to infrequent pollinator visits and pollen limitation. Flowers of R. virginica were infertile after a single day of anthesis, but petals were subsequently maintained for 1-2 d and stamens underwent a color change from bright yellow to red. Second-day flowers may function to increase floral display size and hence fertility, without a concomitant increase in pollen discounting. Studies of bumble bee foraging behavior and correlates of seed set provided indirect support for this hypothesis.     

Morphological specialization of heterantherous Rhynchanthera grandiflora (Melastomataceae) accommodates pollinator diversity

• The tropical Melastomataceae are characterized by poricidal anthers which constitute a floral filter selecting for buzz‐pollinating bees. Stamens are often dimorphic, sometimes with discernible feeding and pollinating functions. Rhynchanthera grandiflora produces nectarless flowers with four short stamens and one long stamen; all anthers feature a narrow elongation with an upwards facing pore.
• We tested pollen transfer by diverse foraging bees and viability of pollen from both stamen types. The impact of anther morphology on pollen release direction and scattering angle was studied to determine the plant's reproductive strategy.
• Medium‐sized to large bees sonicated flowers in a specific position, and the probability of pollen transfer correlated with bee size even among these legitimate visitors. Small bees acted as pollen thieves or robbers. Anther rostrum and pore morphology serve to direct and focus the pollen jet released by floral sonication towards the pollinator's body. Resulting from the ventral and dorsal positioning of the short and long stamens, respectively, the pollinator's body was widely covered with pollen. This improves the plant's chances of outcrossing, irrespective of which bee body part contacts the stigma. Consequently, R. grandiflora is also able to employ bee species of various sizes as pollen vectors.
• The strategy of spreading pollen all over the pollinator's body is rather cost‐intensive but counterbalanced by ensuring that most of the released pollen is in fact transferred to the bee. Thus, flowers of R. grandiflora illustrate how specialized morphology may serve to improve pollination by a functional group of pollinators.

     

The ecology and evolution of pollen odors

The literature is reviewed and new evidence presented that pollen produces odors, which serve multiple functions in pollination and defense. Pollen odor, which originates from pollenkitt, comprises volatiles that belong to the same chemical classes found in flower scents, that are in species-specific mixtures, and that contrast with odors of other floral parts. Pollen can also take up volatiles from surrounding floral odors, but this adsorption is selective and varies among species. Pollen odors are more pronounced in insect- than bird- or wind-pollinated plants, suggesting that volatile emission evolved in part under selection to attract pollinators. Pollen-feeding insects can perceive pollen odor and use it to discriminate between different pollen types and host plants. Pollen odor influences bee foraging, including the location of pollen sources, discrimination of flowers with different amounts of pollen, and hostplant recognition by pollen-specialist species. In the few wind-pollinated plants studied, odors of male flowers or pollen are comparatively high in -methyl alcohols and ketones; these volatiles may serve in pollen defense, with some known to repel insects. Pollen odor often includes chemicals with documented defense activity, which is probably aimed mainly at nonpollinator pollen-feeding insects and pathogens; an involvement in pollen allelopathy is also possible. Pollen volatiles comprise chemically diverse compounds that may play multiple roles, and their emission in pollen odor undoubtedly evolved under the principle, and often conflicting, selective pressures to both protect the male gametophyte and increase its dispersal by animals.     

The number of cells in the pollen of Melastomataceae (Myrtales)

The mature pollen of Melastomataceae is not three-celled, as reported earlier, but two-celled. Two-celled mature pollen is characteristic of all Myrtales, excluding Thymelaeaceae.     

Assessment of pollen reward and pollen availability in Solanum stramoniifolium and Solanum paniculatum for buzz‐pollinating carpenter bees

The two widespread tropical Solanum species S. paniculatum and S. stramoniifolium are highly dependent on the visits of large bees that pollinate the flowers while buzzing them. Both Solanum species do not offer nectar reward; the rewarding of bees is thus solely dependent on the availability of pollen. Flower visitors are unable to visually assess the amount of pollen, because the pollen is hidden in poricidal anthers. In this study we ask whether and how the amount of pollen determines the attractiveness of flowers for bees. The number of pollen grains in anthers of S. stramoniifolium was seven times higher than in S. paniculatum. By contrast, the handling time per five flowers for carpenter bees visiting S. paniculatum was 3.5 times shorter than of those visiting S. stramoniifolium. As a result foraging carpenter bees collected a similar number of pollen grains per unit time on flowers of both species. Experimental manipulation of pollen availability by gluing the anther pores showed that the carpenter bees were unable to detect the availability of pollen by means of chemical cues before landing and without buzzing. Our study shows that the efficiency of pollen collecting on S. paniculatum is based on large inflorescences with short between‐flower search times and short handling time of individual flowers, whereas that of S. stramoniifolium relies on a large amount of pollen per flower. Interestingly, large carpenter bees are able to adjust their foraging behaviour to drastically different strategies of pollen reward in otherwise very similar plant species.     

Specialist pollinators deplete pollen in the spring ephemeral wildflower Claytonia virginica

Pollinators that collect pollen – and specifically, pollen‐specialist bees – are often considered to be the best pollinators of a (host) plant. Although pollen collectors and pollen specialists often benefit host plants, especially in the pollen that they deliver (their pollination “effectiveness”), they can also exact substantial costs because they are motivated to collect as much pollen as possible, reducing the proportion of pollen removed that is subsequently delivered to stigmas (their pollination “efficiency”). From the plant perspective, pollen grains that do not pollinate conspecific stigmas are “wasted”, and potentially costly. We measured costs and benefits of nectar‐collecting, pollen‐collecting, and pollen‐specialist pollinator visitation to the spring ephemeral Claytonia virginica. Visits by the pollen‐specialist bee Andrena erigeniae depleted pollen quickly and thoroughly. Although all pollinators delivered roughly the same number of grains, the pollen specialist contributed most to C. virginica pollen delivery because of high visitation rates. However, the pollen specialist also removed a large number of grains; this removal may be especially costly because it resulted in the depletion of pollen grains in C. virginica populations. While C. virginica appears to rely on pollen transfer by the pollen specialist in these populations, nectar‐collecting visitors could provide the same benefit at a lower cost if their visitation rates increased. Pollen depletion affects a pollinator's value to plants, but is frequently overlooked. If they lower the effectiveness of future floral visitors, visits by A. erigeniae females to C. virginica may be more detrimental than beneficial compared to other pollinators and may, in some circumstances, reduce plant fitness rather than increase it. Therefore, A. erigeniae and C. virginica may vary in their degree of mutualism depending on the ecological context.     

Pollination ecology of Silene acutifolia (Caryophyllaceae): floral traits variation and pollinator attraction

BACKGROUND AND AIMS: The floral display influences the composition of pollinators interacting with a plant species. Geographic and temporal variation in pollinator composition complicates the understanding of the evolutionary consequences of floral display variation. This paper analyses the relationships between Silene acutifolia, a hermaphroditic perennial herb, and its pollinators, based on field studies in the north-west of Spain. METHODS: Studies were conducted over three years (1997-1999). Firstly, the main pollinators of this species were determined for two years in one population. Secondly, pollen limitation in fruit and seed production was analysed by supplementary hand pollinations, and counting the pollen grains and tubes growing in styles for two different-sized populations. Finally, the effect of flower size and number on the rate of visitation and total seed number was examined for 15 marked plants. RESULTS AND CONCLUSIONS: The primary pollinators were long-tongued insects, including Hymenoptera, Lepidoptera and Diptera, but the composition and visitation frequencies differed between years. Pollen limitation occurred in one of the years of study. There was between-population variation in the number of pollen grains and pollen tubes found in styles, suggesting pollen limitation in one population. Overall, pollinators visited plants with more open flowers more frequently, and pollinated more flowers within these plants. Conversely, petal and calyx sizes had no effect on insect visitation. Plants with higher rates of visits produced higher number of seeds, suggesting that pollinator-mediated limitation of seed and fruit production may be important in some years.     

Interannual fluctuations in floral longevity, pollinator visitation and pollination limitation of an alpine plant (Gentiana straminea Maxim., Gentianaceae) at two altitudes in the Qinghai-Tibetan Plateau

Prolonged floral longevity and bumblebees as dominate pollinators in alpine ecosystem have been suggested to overcome pollination limitation of alpine plants arising from the decrease of pollinator activity with increasing altitude. However, this conclusion has never been examined in the Qinghai-Tibetan Plateau (QTP), the highest and largest plateau in the world. In this study, we intended to test year-to-year correlations between floral longevity, visiting frequency and pollen limitation of this species between two populations (at 3200 m and 4000 m) of Gentiana straminea in this plateau. Pollinator exclusion elongated both male and female phases greatly at both sites, and durations of both male and female phases in natural condition varied greatly over three years. The visiting frequency of bumblebees varied greatly at the higher altitude, but seemed to be stable at the lower altitude. Seed production was pollination limited in both populations in most studied years. The floral durations, pollinator frequency and pollination limitation showed no significant and consistent variations with the increase of altitude. The previous hypothesis that the prolonged floral longevity of alpine plants can compensate for low levels of pollinator visitation therefore could not be confirmed, and our results further suggested that in the QTP platform, the altitude shows no consistent effect on the reproductive performance of this species, despite that the fluctuation of visit frequency intensified at the higher altitude.     

Development of the gametophytes, flower, and floral vasculature in Dichorisandra thyrsiflora (Commelinaceae)

The flowers of Dichorisandra thyrsiflora (Commelinaceae) are monosymmetric and composed of three sepals, three petals, six stamens, and three connate carpels. The anthers are poricidal and possess a wall of five cell layers (tapetum included). This type of anther wall, not previously observed in the Commelinaceae, is developmentally derived from the monocotyledonous type via an additional periclinal division and the persistence of the middle layers through anther dehiscence. Secondary endothecial thickenings develop in the cells of the two middle layers only. The tapetum is periplasmodial and contains raphides. Microsporogenesis is successive and yields both decussate and isobilateral tetrads. Pollen is shed as single binucleate grains. The gynoecium is differentiated into a globose ovary, hollow elongate style, and trilobed papillate stigma. Each locule contains six to eight hemianatropous to slightly campylotropous crassinucellar ovules with axile (submarginal) placentation. The ovules are bitegmic with a slightly zig-zag micropyle. Megagametophyte development is of the Polygonum type. The mature megagametophyte consists of an egg apparatus and fusion nucleus; the antipodals having degenerated. The floral vasculature is organized into an outer and inner system of bundles in the pedicel. The outer system becomes ventral carpellary bundles. All other floral vascular traces originate from the inner system.     

Character displacement of male nuptial colour in threespine sticklebacks (Gasterosteus aculeatus)

To study the isolation mechanism of two commonly intermingled louseworts, Pedicularis rhinanthoides and Pedicularis longiflora , pollination biology in three mixed populations with the two species was investigated during a 3-year project. The results indicated that higher flowering density could help to enhance pollinator activity, and thus increase reproductive output. Bumblebees are the exclusive pollinator for the two louseworts and are essential for their reproductive success. Reproductive isolation between the two species is achieved by a combination of pre- and postzygotic isolation mechanisms. Although both species are pollinated by bumblebees, the present study indicates they successfully avoid interspecific pollen transfer due to floral isolation. Mechanical isolation is achieved by the stigma in the two species picking up pollen from different parts of the pollinator's body, whereas ethological isolation occurs due to flower constancy. Additionally, strong postzygotic isolation was demonstrated by non seed set after artificial cross-pollination even with successful pollen tube growth. We describe the hitherto unreported role of variation in the tightness and direction of the twist of the corolla beak in maintaining mechanical isolation between Pedicularis species. Although floral isolation in Pedicularis is incomplete, it is considered to be an important mechanism in maintaining species boundaries in the genus.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 37–48.     

Ragweed pollen sampling in Gallarate (North-West of Milan) during four years (1987–1990)

Summary Ambrosia artemisifolia (elatior), an imported pollen, never sampled in the past in the atmosphere of our country, is beconning the most important Composita responsible for late-summer pollinosis in our area.The documented increased pollen concentration confirmed greater sensitization in people coming to our allergological centre.     

Among-species differences in pollen quality and quantity limitation: implications for endemics in biodiverse hotspots

Background and Aims

Insufficient pollination is a function of quantity and quality of pollen receipt, and the relative contribution of each to pollen limitation may vary with intrinsic plant traits and extrinsic ecological properties. Community-level studies are essential to evaluate variation across species in quality limitation under common ecological conditions. This study examined whether endemic species are more limited by pollen quantity or quality than non-endemic co-flowering species in three endemic-rich plant communities located in biodiversity hotspots of different continents (Andalusia, California and Yucatan).

Methods

Natural variations in pollen receipt and pollen tube formation were analysed for 20 insect-pollinated plants. Endemic and non-endemic species that co-flowered were paired in order to estimate and compare the quantity and quality components of pre-zygotic pollination success, obtained through piecewise regression analysis of the relationship between pollen grains and pollen tubes of naturally pollinated wilted flowers.

Key Results

Pollen tubes did not frequently exceed the number of ovules per flower. Only the combination of abundant and good quality pollen and a low number of ovules per flower conferred relief from pre-zygotic pollen limitation in the three stochastic pollination environments studied. Quality of pollen receipt was found to be as variable as quantity among study species. The relative pollination success of endemic and non-endemic species, and its quantity and quality components, was community dependent.

Conclusions

Assessing both quality and quantity of pollen receipt is key to determining the ovule fertilization potential of both endemic and widespread plants in biodiverse hotspot regions. Large natural variation among flowers of the same species in the two components and pollen tube formation deserves further analysis in order to estimate the environmental, phenotypic and intraindividual sources of variation that may affect how plants evolve to overcome this limitation in different communities worldwide.     

Resource limitation and pollen source (self and outcross) affecting seed production in two louseworts, Pedicularis siphonantha and P. longiflora (Orobanchaceae)

Two animal-pollinated hermaphrodite plants, Pedicularis siphonantha and P. longiflora , have been used to investigate factors limiting seed production in natural populations. To evaluate the potential seed abortion due to resources limitation, seed development has been observed and seed count conducted twice. Seed production per capsule has been compared when flowers have been removed and in a control group. Open pollination has been investigated and pollen supplementation undertaken to estimate the possibility of pollen limitation. Results show that seed abortion is frequent. Stigmatic pollen load is significantly higher than ovule number per ovary under open pollination for both species. Additional self and outcross pollen did not affect seed production. Flower removal significantly increases seed production per capsule, which indicates that seed production of the studied species is limited by available resources. To detect differences in seed production between flowers pollinated by self and outcross pollen, hand pollination of bagged flowers has also been conducted in natural populations of the two Pedicularis species. Compared with open pollination, hand-pollinating self-pollen decreases, while outcross pollen increases seed production per capsule. Such results suggest that inbreeding depression in the two self-compatible species may also result in partial seed abortion under open pollination if mixed pollen is deposited on the stigma. Our results also suggest that pollen interference plays an important role in low female fertility in the two Pedicularis species.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 83–89.     

露蕊乌头(毛茛科)不同海拔居群的传粉生态学

对青藏高原东北部两个不同海拔地点（互助,2460m和海北,3200m）的露蕊乌头Aconitum gymnandrum的传粉生态学进行了比较研究。研究结果表明高海拔居群的单花花期、雄性期持续时间、雌性期持续时间比低海拔长。露蕊乌头不存在无融合生殖现象,尽管高度自交亲和,但产生种子必须依赖传粉媒介。熊蜂是露蕊乌头的主要传粉昆虫,两个居群均存在传粉限制。熊蜂取食花蜜为主,在同一个花序上的访问顺序以由下向上为主,但在互助居群和海北居群有3．9％和2．7％的访花是由上向下进行的。统计发现有37．7％和29．3％的访问行为发生在同一植株内;因此,同株异花传粉造成的自交在露蕊乌头中仍然不能避免。低海拔居群的熊蜂种类和访花频率均高于高海拔居群,但自然状态下两个居群的结实率并没有显著差异。高海拔露蕊乌头居群可能具有较长的柱头持续时间,从而补偿了由于访花频率低带来的结实率降低。