How pollinator-mediated mating varies with population size in plants

Comparison of the pollination efficiency of Apis cerana with wild bees in oil-seed camellia fields

On Limits to Seed Production

Limitations to Fruit Production in Ionopsis utricularioides (Orchidaceae)

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Reports of sharply reduced feral bee populations (Apis mellifera) due to harsh winters and prevalence of several bee diseases have raised concerns that pollination and fruit set in pumpkin fields will be adversely affected. In 1995 and 1996, five and eight pumpkin (Cucurbita pepo) fields, respectively, were inventoried on three occasions per season for pollinator activity and percent fruit set. Pollen removal from male flowers was determined visually using a rating scale, while deposition of pollen on stigmata of female flowers was judged by rating fluorescence of pollen on the stigmatic surface under a “black light.” Samples were taken from 15 to 30 locations in each field, and female flowers tagged. These were considered set if they had enlarged to fist size within 14 days. In both years, the amount of pollen remaining on male flowers was negatively correlated with female flower fluorescence ratings. Neither pollen on male flowers nor stigma fluorescence were significantly correlated with percent fruit set. Fifty-two percent of tagged flowers set fruit in both years, with a range of 24% to 84%, and 17% to 78% in 1995 and 1996, respectively. Presence of bee hives in or near the fields had no effect on fruit set. The results indicate that the pollen removal and deposition ratings used were not reliable for predicting fruit set in farmers' fields. In these 2 years, bee hives were not needed in the sampled fields.

Fragmentation of plant communities is generally considered to be detrimental to the survival of the constituent species, especially those that are rare. We investigated the effects of fragmentation on reproductive biology in nine populations of the rare taxon Banksia nivea subsp. uliginosa, differing in size and habitat context. Small mammals were the main pollinator, with lower contributions from honeybees and birds (honeyeaters). There was no significant relationship between population size, fruit set and seed germinability. Fruit set was marginally highest (25.5 ± 3.4) in medium size populations and was variable over years. Lower fruit set (1.0 ± 0.6) in the smallest population may be due to inbreeding depression or lack of pollinators in a degraded habitat, but low fruit set was also observed in the largest population despite high levels of gene flow, possibly due to low pollinator visitation rate in a low-density population. Seeds from all populations had high germination success (>93.4%). Predation occurred in up to 56% of fruit and increased with increasing population size, but was not significantly different between populations of different size. Our study provided evidence that the reproductive output in species in a naturally fragmented landscape system may be resilient to reduced population size and other influences of anthropogenic fragmentation.

Background The relative influence of both conspecific and heterospecific plant density on the fecundity of generalist rewardless plants is unclear. Aims To test whether distance from both conspecific and heterospecific plants influence pollen removal, deposition, and fruit set in the nectarless orchid, Dactylorhiza sambucina that co-flowers with the rewarding Ranunculus apenninus. Methods I experimentally translocated D. sambucina inflorescences and measured pollen removal and deposition rates. I measured plant height and distance to D. sambucina and R. apenninus plants in 30 D. sambucina patches of varying density. I quantified the proportion of pollen removed, pollinia deposited, and fruit set and tested whether these vary according to plant height and distance to both D. sambucina and R. apenninus. Results Translocated inflorescences outside the population had a higher proportion of pollen removal than control plants within the population, while pollinia deposition rates did not differ. Increased D. sambucina height increased pollen removal rates, while increasing distance from R. apenninus resulted in both increased pollen deposition and fruit set. Conclusion Pollination and fecundity of a nectarless orchid may increase with increasing distance from rewarding heterospecifics. This study improves our understanding of how the fecundity of rewardless plants is influenced by co-flowering plants.

European honeybees (Apis mellifera) were less efficient pollinators ofGrevillea barklyana than nectar-feeding birds. Nectar-collecting honeybees did not contact reproductive parts of flowers. Pollen-collecting honeybees preferentially visited malestage flowers but rarely visited female-stage flowers. Fruit set on caged inflorescences that allowed access to honeybees but excluded birds was reduced by more than 50% compared to inflorescences that were visited by both types of visitors. Further, fruit set on caged inflorescences was less than on bagged inflorescences that excluded both birds and honeybees, indicating that pollen removal by bees decreased opportunities for delayed autonomous selfing in the absence of birds. Although fruit set was not pollen-limited at the study site, pollen removal by honeybees would decrease fruit set in small populations where birds are scarce. In addition, pollen removal by honeybees would reduce opportunities for outcrossing and reproductive success through male function. Although honeybees have been in Australia for insufficient time to have exerted selection on floral traits, evolutionary shifts in response to these animals are likely to occur in the future.

The period between the beginning of anthesis and flower senescence modulates the transport of pollen by pollinators among conspecific flowers, and its length may therefore influence reproductive success. This study evaluated whether floral longevity favours pollen removal from the anthers over fecundity (seed set) in an ornithophilous species that does not undergo pollen limitation. Field investigations were conducted on floral longevity, nectar production, pollinator behaviour, and variations in fruit set (FS), mean number of seeds per fruit (MSF) and pollen removal by hummingbirds (PR) during the anthesis of Salvia sellowiana in south-east Brazil. Anthesis of flowers exposed to pollinators lasted 4 d, as well as on flowers with pollen removed from the anthers or deposited on the stigma. The longevity of bagged flowers was significantly higher (approx. 9 d). FS and PR reached 87.2 and 90 %, respectively, in natural conditions. PR increased gradually over the period of anthesis; however, FS and MSF reached their maxima in the first hours of anthesis. Nectar production was continuous, but the secretion rate was reduced after pollination. The removal of nectar from non-pollinated flowers stimulated its production. The longevity of anthesis in S. sellowiana seems to be related to the mechanism of gradual dispensing of pollen, resulting in greater male reproductive success. This is in agreement with the pollen-donation hypothesis. The small number of ovules (four) of S. sellowiana and the high frequency and the foraging mode of its pollinators may favour the selection for floral longevity driven by male fitness in this system.

Pollination efficiency and reproductive success vary strongly among populations of most animal-pollinated plant species, depending on their size and local density, whereas individual plants within populations experience varying levels of reproductive output as a result of differences in floral display. Although most orchid species have been shown to be severely pollination limited, few studies have investigated the impact of the above-mentioned factors on pollination success and reproduction, especially in rewarding species. In this study, the impact of population size, local density of flowering plants, and floral display on the rates of pollinia export and fruit production was investigated in 13 natural populations of the rewarding terrestrial orchid Listera ovata. In addition, an emasculation experiment was set up to examine how floral display and local density of flowering plants affected the relative importance of cross- vs. geitonogamous pollination in determining fruit set. In the studied populations, pollination efficiency, pollen removal, and fruit set increased with increasing population size until a threshold value of 30–40 flowering plants was reached, above which pollination efficiency and reproductive output decreased again. On average, plants with large floral displays showed higher proportional pollinia removal and fruit set compared with smaller plants. Fruit production was also significantly and positively related to local plant density, whereas emasculation did not affect the relationship between local plant density and fruit set, suggesting that geitonogamous pollination did not affect the outcome of female function. The results of this study are discussed in the light of the flowering mechanism of the species and its generalized pollination system.

Pollinator-mediated selection and evolution of floral traits have long fascinated evolutionary ecologists. No other plant family shows as wide a range of pollinator-linked floral forms as Orchidaceae. In spite of the large size of this model family and a long history of orchid pollination biology, the identity and specificity of most orchid pollinators remains inadequately studied, especially in the tropics where the family has undergone extensive diversification. Angraecum (Vandeae, Epidendroideae), a large genus of tropical Old World orchids renowned for their floral morphology specialized for hawkmoth pollination, has been a model system since the time of Darwin. The pollination biology of A. cadetii, an endemic species of the islands of Mauritius and Reunion (Mascarene Islands, Indian Ocean) displaying atypical flowers for the genus (white and medium-size, but short-spurred) was investigated. Natural pollinators were observed by means of hard-disk camcorders. Pollinator-linked floral traits, namely spur length, nectar volume and concentration and scent production were also investigated. Pollinator efficiency (pollen removal and deposition) and reproductive success (fruit set) were quantified in natural field conditions weekly during the 2003, 2004 and 2005 flowering seasons (January to March). Angraecum cadetii is self-compatible but requires a pollinator to achieve fruit set. Only one pollinator species was observed, an undescribed species of raspy cricket (Gryllacrididae, Orthoptera). These crickets, which are nocturnal foragers, reached flowers by climbing up leaves of the orchid or jumping across from neighbouring plants and probed the most 'fresh-looking' flowers on each plant. Visits to flowers were relatively long (if compared with the behaviour of birds or hawkmoths), averaging 16.5 s with a maximum of 41.0 s. At the study site of La Plaine des Palmistes (Pandanus forest), 46.5 % of flowers had pollen removed and 27.5 % had pollinia deposited on stigmas. The proportion of flowers that set fruit ranged from 11.9 % to 43.4 %, depending of the sites sampled across the island. Although orthopterans are well known for herbivory, this represents the first clearly supported case of orthopteran-mediated pollination in flowering plants.

Conspecific density is an important factor in determining the fecundity of pollinator-dependent plants. However, the outcome of increased conspecific density can range from reduced per capita pollination rates and fecundity (competition) to increased per capita pollination rates and fecundity (facilitation). I tested whether local conspecific density influenced pollination and fecundity in the rewarding orchid, Gymnadenia conopsea. I measured the distance to each of the nearest five plants in sixty patches of varying density in a large population in the Italian Apennine mountains. I examined whether pollen removal, pollen deposition, fruit, and seed set varied according to mean distance from conspecifics and plant height. After controlling for plant position in the population, the proportion of pollen removal, pollen deposition, and fruit set decreased with increasing distance from conspecifics, while pollen removal and deposition, but not fruit set, increased with increasing plant height. The proportion of viable seeds per fruit did not vary according to conspecific distance. Decreases in pollen removal, pollen deposition, and fruit set occurred within an average of 1 m distance from conspecific plants, indicating that local aggregation is an important determinant of both pollination success and fecundity. These results suggest that despite being a geographically widespread orchid, G. conopsea may suffer reduced fecundity when it occurs in small, isolated populations. Such field investigations can potentially lead to better predictions for successful re-establishment of populations of orchids under conservation-aided restoration efforts.

Trapping trichomes, a common trait found in protogynous trap flowers, are thought to enable interaction with pollinators, promoting both pollen removal and fruit set. However, this hypothesis has yet to be empirically confirmed. In this study, we experimentally tested whether trapping trichomes favour male and female functions in trap flowers. Using protogynous South American Aristolochia esperanzae trap flowers, we observed the temporal variation in both sexual functions and associated them with the wilting of the flower and trapping trichomes. Next, we described the trapping trichomes features associated with the capture, entrapment and release of pollinators. Finally, we used manipulative experiments involving trapping trichome removal to quantify their influence on pollinator entrapment, pollen removal and fruit set. While flowers wilted after the end of the female phase, the trapping trichomes only wilted some hours after pollen presentation, promoting a delay in the exit of pollinators, so they carried pollen. Trapping trichomes features, such as a density gradient towards the interior of the tube, unidirectional orientation and epicuticular waxes, may be important in the capture and entrapment of pollinators. Trapping trichome removal decreased drastically the probability of pollinator entrapment and pollen removal, leading to no fruit set. These results indicate that the evolution and maintenance of trapping trichomes in trap flowers probably occurred through selective pressures from both sexual functions and highlight their role in the functioning of this highly phenotypically specialized pollination system. Read the free Plain Language Summary for this article on the Journal blog.

Taxonomically related species can differ in a number of reproductive traits, which may translate into a differential mating system and pollination success. Here we compare two hermaphroditic insect-pollinated Daphne species (D. rodriguezii and D. gnidium) which differ in distribution (island endemic vs. mediterranean) and floral traits (long- vs. short-tube corolla). We investigated their mating system and pollen limitation by means of hand-pollination experiments and quantified the diversity and abundance of flower visitors by direct observations. Plant size and five reproductive traits (flower production, proportion of viable anthers, pollen production, flower tube length and tepal area) were studied to assess how they contribute to reproductive success, measured as proportion of pollen grains germinated per stigma and fruit set. Selfing was very low and pollen limitation existed in both species, though was higher in D. rodriguezii probably due to the scarcity of flower visitors. The low fruit set in both species suggests that most of the pollen grains found on stigmas are self-pollen. Pollinators appeared to favour some floral traits (specifically, flower tube length or tepal area) in both species, although flower crop in D. rodriguezii was the only reproductive trait influencing fruit set. In both species, the highest variability in reproductive traits and pollination success was within individuals. Our findings suggest that despite both species showed similar mating system, dependency on outcrossing pollen and selection of floral traits, pollen limitation was higher in D. rodriguezii, probably as a higher proportion of self-pollen arrives to its stigmas.

Eremosparton songoricum (Litv.) Vass. is a rare, central Asian desert species which shows lower fruit set and seed set (<16%) than most hermaphroditic species. We hypothesized that fruit production was limited by pollen and resources. To evaluate potential fruit abortion due to pollen limitation, supplemental hand-pollination was undertaken, the mating system was investigated and the foraging behavior of pollinators was recorded. To investigate possibile resource limitation, flowers and young pods were artificially removed and fertilization were manipulated. The results showed that under natural pollination, the number of pollen deposited on the stigma greatly exceed the number of ovules per ovary. Mating system experiments showed that the species is self-compatible, but depended on pollinators to set seeds. Supplemental outcross pollination increased fruit set significantly. The most frequent effective pollinator Megachile terminate Morawitz, was observed pollinating many flowers of the same individual plant (74.5±1.3%). These results suggested that fruit production is affected by insufficient outcross pollen rather than by pollen quantity. Removal of 2/3 of the flowers and young pods led to significantly higher fruit set, as did addition of fertilizers (N–P–K: 0.025–0.05–0.013 g, N–P–K: 0.05–0.1–0.025 g) showing that reducing resource acceptors and increasing inorganic resources both helps to improve fruit set. We therefore conclude that reproductive success of E. songoricum is limited by both outcross pollen and available nutrients.