Breeding system and factors limiting fruit production in the nectarless orchid Broughtonia lindenii

Low fruit set values in most orchids (especially epiphytic and tropical species) are normally thought to be the consequence of pollination constraints and limited resources. In particular, pollination constraints are modulated by pollinator visitation rates, pollinator visitation behaviour (promoting crossing or selfing), the type and number of pollinia deposited on stigmas (in the case of orchids with subequal pollinia) and the amount of pollen loaded per inflorescence. In order to assess to what extent these factors can affect fruit set in specific orchid-pollinator systems, the repercussions of some of these aspects on reproduction of Broughtonia lindenii were examined in a coastal population in western Cuba. The study focused on plant breeding system, importance of pollen load and type of pollinia on subsequent fruit and seed, limiting factors of seed production and interaction with pollinators. This species presents long-lasting flowers that senesce after all forms of effective visit. Pollinator dependence for fruit production was demonstrated, while hand-pollination experiments revealed self-compatibility and inbreeding depression at seed level. More pollinia on stigmas enhance the proportion of well-developed seeds. In contrast, the pollinia type used in pollination is not important for seed quality of fruits, suggesting that small pollinia are not rudimentary. Natural fruit set in two consecutive years was substantially affected by pollinator activity, and also by systematic depredatory activity of ants and a caterpillar. Considering that this orchid completely lacks nectar and that the local assemblage of pollinators and predators influenced its reproduction, a minor importance of resource constraints in this epiphyte (with long-lasting reserve structures) is confirmed at least for a short time.     

Pollinator limitaton in a deceptive orchid, Pogonia japonica, on a floating peat mat

Deceptive orchid species that offer no floral rewards to pollinators often experience pollinator limitation because they. The breeding system and fruit set were investigated in order to examine pollinator limitation for a population of a deceptive orchid, Pogonia japonica , on a floating peat mat in Mizorogaike Pond, western Japan. Fruit sets for outcross-pollinated, self-pollinated and pollinator-excluded flowers were 75%, 80%, and 0%, respectively. Thus, this species was self-compatible but neither autogamous nor apogamous. The fruit set for open-pollinated flowers was 22.9% and 17.4% in 1994 and 1995, suggesting that pollinator limitation did occur in the field. Larger flowers were more likely to develop into fruit than smaller ones in open pollination (36% vs 8%). Thus, flower size seems to affect fruiting of this species. No effective visits by major pollinators such as bees, which could carry pollinia, were observed during 16.5 h of observation during daytime. Instead, thrips were often found moving on gynostemium or pollinia and circumstantial evidence that suggests thrips may partly transfer pollen in pieces out of a granular pollinium of P. japonica was obtained. Pogonia japonica should come to use thrips as a pollinator to supplement low fruit set under limited flower visitation by major pollinators.     

Autonomous self‐pollination in the nectarless orchid Pogonia minor

The pollination biology of the nectarless orchid Pogonia minor was investigated in central Japan. The investigation revealed that the solitary flowers failed to attract pollinators, while high rates of fruit set were observed in the natural population. Comparable levels of fruit set were obtained in bagged, artificial self‐pollinated and artificial cross‐pollinated plants, indicating that the species is not pollinator‐limited for fruit set under natural conditions. Autonomous self‐pollination in P. minor resulted from a reduced rostellum, which allowed contact between the pollinia and the stigma. Self‐pollination is thought to be an adaptive response that provides reproductive assurance under conditions of pollinator limitation. Since pollen limitation is generally known to be frequent among deceptive orchids, strong pollen limitation is probably a driving force in the autonomous self‐pollination mechanism in the nectarless orchid P. minor.     

Effects of flowering plant density on pollinator visitation, pollen receipt, and seed production in Delphinium barbeyi (Ranunculaceae)

Variation in flowering plant density can have conflicting effects on pollination and seed production. Dense flower patches may attract more pollinators, but flowers in those patches may also compete for pollinator visits and abiotic resources. We examined how natural and experimental conspecific flowering plant density affected pollen receipt and seed production in a protandrous, bumble bee-pollinated wildflower, Delphinium barbeyi (Ranunculaceae). We also compared floral sex ratios, pollinator visitation rates, and pollen limitation of seed set from early to late in the season to determine whether these factors mirrored seasonal changes in pollen receipt and seed production. Pollen receipt increased with natural flowering plant density, while seed production increased across lower densities and decreased across higher flower densities. Experimental manipulation of flowering plant density did not affect pollinator visitation rate, pollen receipt, or seed production. Although pollinator visitation rate increased 10-fold from early to late in the season, pollen receipt and seed set decreased over the season. Seed set was never pollen-limited. Thus, despite widespread effects of flowering plant density on plant reproduction in other species, the effects of conspecific flowering plant density on D. barbeyi pollination and seed production are minor.     

开花时间与伴生种对鹤首马先蒿传粉和生殖成功的影响

开花物候是物种间相互作用的重要生活史特征和适合度因子,在全球气候变化的背景下而备受关注.为探讨开花时间如何存种内和种间水平上影响植物的传粉和生殖成功,我们连续3年(2003-2005)对不同花期和伴生种存在情况下的鹤首马先蒿(Pedicularis gruina)的传粉者访花忠实度、受粉率、坐果率、单果种子产量和果实被啃食频率进行了比较研究.结果表明鹤首马先蒿的坐果率主要受其传粉环境的影响:在没有伴生种时,不同时期鹤首马先蒿的坐果率没有显著差异,34-38％的花可以坐果;在有伴生种存在时既可以显著提高其坐果率,也可显著降低其坐果率,这取决于传粉者类型以及伴生种密穗马先蒿(P.densispica)花期的差异.密穗马先蒿具有花蜜和花粉双重报酬,在群落中可以作为主体物种吸引传粉者并间接促进与其伴生的鹤首马先蒿的传粉和生殖成功.同样无蜜的管花马先蒿(P.siphonantha)和鹤首马先蒿伴生,则是通过提高群落水平对传粉者的吸引力进而直接促进鹤首马先蒿的传粉和生殖成功.此外,研究结果也表明开花时间对坐果率没有显著影响,但是显著影响单果种子产量和果实被啃食的频率.在相同的传粉条件下早花期植株单果种子产量显著高于晚花期的种子产量,同时早花期的果实被啃食的频率显著增加.     

Effective pollinia transfer by settling moths’ legs in an orchid Habenaria aitchisonii

A great diversity of flower morphology in orchids has long been thought to be selected by diverse pollinators. Habenaria Willd. (Orchidaceae) species are generally characterized by long nectar spurs and pollinated by long‐tongued insects (Lepidoptera), the mechanical fit between the spur and pollinator proboscis length being supposedly caused by “arms race” reciprocal selection. Here, we report that flowers of Habenaria aitchisonii Rchb. f. with nectar spurs (approximately 9 mm) were pollinated by three species of settling noctuid moths whose proboscises varied in length from 10 to 16 mm. When a settling moth crawled on the spikes and probed the flowers for nectar, pollinia were placed on the moths’ legs rather than on other body parts. Our 5‐year survey of pollinia movement and 3‐year supplemental pollination experiments indicated that fruit and seed production in this orchid were not often pollen‐limited at flower level. In a natural population in Shangri‐La, Southwest China, the proportions of pollinia removal and deposition on stigmas by moth legs were 93.8% and 83.5%, respectively. This finding of efficient pollen transfer by the pollinators’ legs in H. aitchisonii adds a new example of diverse pollinia placement on pollinators (here settling moths) in the Orchidaceae.     

Intrinsic and extrinsic factors acting on the reproductive process in alpine‐snowbed plants: roles of phenology,biological interaction,and breeding system

Pollinator activity and competition for pollinators lead to quantitative and qualitative pollen limitations on seed production and affect the reproductive success of plant species, depending on their breeding system (e.g., self‐compatibility and heterospecific compatibility) and genetic load (e.g., inbreeding depression and hybrid inviability). In alpine ecosystems, snowmelt regimes determine the distribution and phenology of plant communities. Plant species growing widely along a snowmelt gradient often grow with different species among local populations. Their pollinators also vary in their abundance, activity, and behavior during the season. These variations may modify plant–pollinator and plant–plant interactions. We integrated a series of our studies on the alpine dwarf shrub, Phyllodoce aleutica (Ericaceae), to elucidate the full set of intrinsic (species‐specific breeding system) and extrinsic factors (snow condition, pollinator activity, and interspecific competition) acting on their reproductive process. Seasonality of pollinator activity led to quantitative pollen limitation in the early‐blooming populations, whereas in the late‐blooming populations, high pollinator activity ensured pollination service, but interspecific competition for pollinators led to qualitative and quantitative pollen limitation in less competitive species. However, negative effects of illegitimate pollen receipt on seed‐set success might be reduced when cryptic incompatibility systems (i.e., outcross pollen grains took priority over self‐ and heterospecific pollen grains) could effectively prevent ovule and seed discounting. Our studies highlight the importance of species‐specific responses of plant reproduction to changing pollinator availability along environmental gradients to understand the general features of pollination networks in alpine ecosystems.     

The independent and combined effects of floral traits distinguishing two pollination ecotypes of a moth‐pollinated orchid

Identifying traits and agents of selection involved in local adaptation is important for understanding population divergence. In southern Sweden, the moth‐pollinated orchid Platanthera bifolia occurs as a woodland and a grassland ecotype that differ in dominating pollinators. The woodland ecotype is taller (expected to influence pollinator attraction) and produces flowers with longer spurs (expected to influence efficiency of pollen transfer) compared to the grassland ecotype. We examined whether plant height and spur length affect pollination and reproductive success in a woodland population, and whether effects are non‐additive, as expected for traits influencing two multiplicative components of pollen transfer. We reduced plant height and spur length to match trait values observed in the grassland ecotype and determined the effects on pollen removal, pollen receipt, and fruit production. In addition, to examine the effects of naturally occurring variation, we quantified pollinator‐mediated selection through pollen removal and seed production in the same population. Reductions of plant height and spur length decreased pollen removal, number of flowers receiving pollen, mean pollen receipt per pollinated flower, and fruit production per plant, but no significant interaction effect was detected. The selection analysis demonstrated pollinator‐mediated selection for taller plants via female fitness. However, there was no current selection mediated by pollinators on spur length, and pollen removal was not related to plant height or spur length. The results show that, although both traits are important for pollination success and female fitness in the woodland habitat, only plant height was sufficiently variable in the study population for current pollinator‐mediated selection to be detected. More generally, the results illustrate how a combination of experimental approaches can be used to identify both traits and agents of selection.     

Interspecific pollen transfer between two coflowering species was minimized by bumblebee fidelity and differential pollen placement on the bumblebee body

Aims When sympatric flowering plant species in a natural community share pollinators, study of plant–plant interactions via interspecific pollen transfer (IPT) is essential for understanding species coexistence. However, little is known about the extent of IPT between interactive species and its causes.Methods To explore how sympatric flowering plants sharing pollinators minimize deleterious effects of IPT, we investigated the pollination ecology of two endemic species, Salvia przewalskii and Delphinium yuanum, in an alpine meadow in the Hengduan Mountains, southwest China. We quantified conspecific and interspecific visits by shared bumblebee pollinators, amounts of pollen placed on different body sites of the pollinators and stigmatic pollen loads on open-pollinated flowers. To examine whether IPT affects female fitness, we measured pollen germination and seed production in the two species in an artificial pollination experiment.Important findings One bumblebee species, Bombus trifasciatus, was found to be the sole effective pollinator for the two coflowering species. Pollination experiments indicated that deposition of heterospecific pollen could significantly decrease seed set in both species. Experiments showed that S. przewalskii pollen could germinate well on stigmas of D. yuanum, inhibiting conspecific pollen germination in D. yuanum. However, seed set was not lower under open pollination than under cross-pollination within species, suggesting that no female fitness loss was caused by IPT. In foraging bouts with pollinator switches, switches from S. przewalskii to D. yuanum were relatively more frequent (8.27%) than the converse (1.72%). However, IPT from S. przewalskii to D. yuanum accounted for only 1.82% of total stigmatic pollen loads while the reverse IPT to S. przewalskii was 8.70%, indicating that more switches of bumblebees to D. yuanum did not result in higher IPT. By contrast, selection for reduced IPT to S. przewalskii would limit pollinator switches from D. yuanum. We found that a bumblebee generally carried pollen grains from both species but the two species differed in the position of pollen placement on the bumblebee's body; S. przewalskii ' s pollen was concentrated on the dorsal thorax while D. yuanum ' s pollen was concentrated ventrally on the head. This differential pollen placement along with pollinator fidelity largely reduced IPT between the two species with a shared pollinator.     

Do pollen carryover and pollinator constancy mitigate effects of competition for pollination?

Pollinator constancy and pollen carryover are both thought to mitigate competitive effects that result when shared pollinators cause loss of pollen to heterospecific flowers. I present analytical and simulation models to investigate how pollinator constancy and pollen carryover interact with each other and with the relationship between pollen receipt and seed set to determine pollination success in competitive environments. With inconstant pollinators, increased pollen carryover reduces variance in pollen receipt without affecting average pollen receipt. Consequently, for flowers requiring at least a threshold quantity of pollen for success, rare flowers with inconstant pollinators benefit from reduced carryover, especially for high pollen receipt thresholds, whereas common flowers benefit from increased carryover, especially for low receipt thresholds. Pollinator constancy is predicted to increase pollen receipt, especially if pollen carryover rates are low. As a result, increased pollinator constancy reduces the range of pollen receipt thresholds for which carryover is beneficial. Similarly, for flowers whose pollination success is a convex function of pollen receipt, carryover is expected to increase fecundity if pollinators are inconstant, but with even a low degree of pollinator constancy, carryover reduces fecundity. These results predict that rare plants with many ovules per flower benefit from dispersing aggregations of pollen, especially if their pollinators exhibit constancy, whereas plants with inconstant pollinators and low thresholds of pollen receipt benefit from pollen grains dispersing individually to increase the number of flowers reached by the pollen.     

Pollen limitation and fruiting failure related to canopy closure in Calypso bulbosa (Orchidaceae), a northern food‐deceptive orchid with a single flower

Natural fruit set is constrained by pollen limitation and fruiting failure, and pollen limitation is expected to be especially severe in deceptive orchids. We performed hand cross‐pollinations in ten populations of a food‐deceptive orchid, Calypso bulbosa, under sparse and dense canopies in three non‐consecutive years. We explored the relationships between natural fruit set, pollen limitation and fruiting failure. Mean natural fruit set over the years was 60%, which is exceptionally high for a deceptive orchid. On average, hand cross‐pollination increased fruit set by 23%. Among open‐pollinated plants that did not set a fruit, 55.5% were estimated to be pollen limited and 44.5% to be limited by fruiting failure, i.e. inability to set a fruit after pollination. In species with high natural fruit set, hand cross‐pollination experiments may not always detect statistically significant pollen limitation. In our case, pollen limitation tended to become significant when the natural fruit set dropped below 60%. Canopy cover had a significant effect on fruiting failure, which was more severe under a dense canopy. Although our results demonstrate pollen limitation in many cases, they also highlight the fact that food deception can be a very effective pollination strategy. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013 , 171 , 744–750.     

Pollinator‐mediated interactions between cultivated papaya and co‐flowering plant species

Many modern crop varieties rely on animal pollination to set fruit and seeds. Intensive crop plantations usually do not provide suitable habitats for pollinators so crop yield may depend on the surrounding vegetation to maintain pollination services. However, little is known about the effect of pollinator‐mediated interactions among co‐flowering plants on crop yield or the underlying mechanisms. Plant reproductive success is complex, involving several pre‐ and post‐pollination events; however, the current literature has mainly focused on pre‐pollination events in natural plant communities. We assessed pollinator sharing and the contribution to pollinator diet in a community of wild and cultivated plants that co‐flower with a focal papaya plantation. In addition, we assessed heterospecific pollen transfer to the stigmatic loads of papaya and its effect on fruit and seed production. We found that papaya shared at least one pollinator species with the majority of the co‐flowering plants. Despite this, heterospecific pollen transfer in cultivated papaya was low in open‐pollinated flowers. Hand‐pollination experiments suggest that heterospecific pollen transfer has no negative effect on fruit production or weight, but does reduce seed production. These results suggest that co‐flowering plants offer valuable floral resources to pollinators that are shared with cultivated papaya with little or no cost in terms of heterospecific pollen transfer. Although HP reduced seed production, a reduced number of seeds per se are not negative, given that from an agronomic perspective the number of seeds does not affect the monetary value of the papaya fruit.     

Floral resources,pollinators and fruiting in a threatened tropical deciduous tree

Aims Nothapodytes nimmoniana (Family Icacinaceae) is a deciduous tree species distributed in Asia facing severe population decline. Wood chips from the tree are a source of camptothecin, a globally sought-after alkaloid with cancer-treating properties, and are harvested unsustainably in natural forests. We studied the pollination ecology of the species and asked if there are constraints in pollination and fruiting success in its natural populations. We also discuss the potential effects of wood extraction on pollinators and reproductive success in the population.Methods The study was carried out during the flowering season in two populations with varying exploitation levels, both located in the Protected Area Network in the Western Ghats of Karnataka State. We assessed floral resource availability and measured pollinator diversity and activity in the canopy from the perspective of the biology of the species. We quantified pollinator abundance and percent fruit set as a function of population density.Important findings Flowers belong to the fly pollination syndrome and are open to visits by generalists such as Apis dorsata and Trigona iridipennis. Fruiting success did not vary between exploited and unexploited sites, but there was a significant difference in pollinator assemblage. In a lean male flowering year, fruit set was low, suggesting pollen limitation in the population. No fruiting was recorded in the second year at the site where trees were felled soon after the flowering period. We argue that when male floral resources are altered in exploited populations, pollinators of generalist nature may show a shift in foraging pattern. Also, generalist plants may be as susceptible to pollination loss as are specialist plants. The need for outcrossing in the species would probably exacerbate this vulnerability.     

Orthoptera, a new order of pollinator

Background and Aims

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.

Methods

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).

Key Results

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.

Conclusions

Although orthopterans are well known for herbivory, this represents the first clearly supported case of orthopteran-mediated pollination in flowering plants.     

Reproductive system and fitness of Vriesea friburgensis,a self‐sterile bromeliad species

Reproductive biology and plant fertility are directly related to many aspects of plant evolution and conservation biology. Vriesea friburgensis is an epiphytic and terrestrial bromeliad endemic to the Brazilian Atlantic rainforest. Hand‐pollination experiments were used to examine the reproductive system in a wild population of V. friburgensis. Plant fertility was assigned considering flower production, fruit and seed set, seed germination, and pollen viability. Self‐sterility observed from spontaneous selfing and manual self‐pollination treatments may be the consequence of late‐acting self‐incompatibility. Hand‐pollination results indicated no pollen limitation in the population studied. Floral biology features such as a few daily open flowers, nectar production, and sugar concentration corroborate hummingbirds as effective pollinators, although bees were also documented as pollinators. Components of fitness such as high flower, fruit, and seed production together with high seed and pollen viability indicate that this wild population is viable. From a conservation point of view, we highlight that this self‐sterile species depends on pollinator services to maintain its population fitness and viability through cross‐pollination. Currently, pollinators are not limited in this population of V. friburgensis. Conversely, the maintenance and continuous conservation of this community is essential for preserving this plant–pollinator mutualism.     

Orchid sexual deceit provokes ejaculation

Sexually deceptive orchids lure pollinators by mimicking female insects. Male insects fooled into gripping or copulating with orchids unwittingly transfer the pollinia. The effect of deception on pollinators has been considered negligible, but we show that pollinators may suffer considerable costs. Insects pollinating Australian tongue orchids (Cryptostylis species) frequently ejaculate and waste copious sperm. The costs of sperm wastage could select for pollinator avoidance of orchids, thereby driving and maintaining sexual deception via antagonistic coevolution or an arms race between pollinator learning and escalating orchid mimicry. However, we also show that orchid species provoking such extreme pollinator behavior have the highest pollination success. How can deception persist, given the costs to pollinators? Sexually-deceptive-orchid pollinators are almost exclusively solitary and haplodiploid species. Therefore, female insects deprived of matings by orchid deception could still produce male offspring, which may even enhance orchid pollination.     

Experimental reduction of pollinator visitation modifies plant–plant interactions for pollination

The strength of interactions between plants for pollination depends on the abundance of plants and pollinators in the community. The abundance of pollinators may influence plant associations and densities at which individual fitness is maximized. Reduced pollinator visitation may therefore affect the way plant species interact for pollination. We experimentally reduced pollinator visitation to six pollinator‐dependent species (three from an alpine and three from a lowland community in Norway) to study how interactions for pollination were modified by reduced pollinator availability. We related flower visitation, pollen limitation and seed set to density of conspecifics and pollinator‐sharing heterospecifics inside 30 dome‐shaped cages partially covered with fishnet (experimental plots) and in 30 control plots. We expected to find stronger interactions between plants in experimental compared to controls plots. The experiment modified plant–plant interactions for pollination in all the six species; although for two of them neighbourhood interactions did not affect seed set. The pollen limitation and seed set data showed that reduction of pollinator visits most frequently resulted in novel and/or stronger interactions between plants in the experimental plots that did not occur in the controls. Although the responses were species‐specific, there was a tendency for increasing facilitative interactions with conspecific neighbours in experimental plots where pollinator availability was reduced. Heterospecifics only influenced pollination and fecundity in species from the alpine community and in the experimental plots, where they competed with the focal species for pollination. The patterns observed for visitation rates differed from those for fecundity, with more significant interactions between plants in the controls in both communities. This study warns against the exclusive use of visitation data to interpret plant–plant interactions for pollination, and helps to understand how plant aggregations may buffer or intensify the effects of a pollinator loss on plant fitness.     

Reproductive interactions mediated by flowering overlap in a temperate hummingbird–plant assemblage

Pollinator‐mediated competition through shared pollinators can lead to segregated flowering phenologies, but empirical evidence for the process responsible for this flowering pattern is sparse. During two flowering seasons, we examined whether increasing overlap in flowering phenology decreased conspecific pollination, increased heterospecific pollination, and depressed seed output in the seven species composing a hummingbird–plant assemblage from the temperate forest of southern South America. Overall trends were summarized using meta‐analysis. Despite prevailing negative associations, relations between phenological overlap and conspecific pollen receipt varied extensively among species and between years. Heterospecific pollen receipt was low and presumably of limited biological significance. However, our results supported the hypothesis that concurrent flowering promotes interspecific pollen transfer, after accounting for changes in the abundance of conspecific flowers. Seed output was consistently reduced during maximum phenological overlap during the first flowering season because of limited fruit set. Responses varied more during the second year, despite an overall negative trend among species. Relations between estimated effects of phenological overlap on pollination and seed output, however, provided mixed evidence that conspecific pollen loss during pollinator visits to foreign flowers increases pollen limitation. By flowering together, different plant species might benefit each other's pollination by increasing hummingbird recruitment at the landscape level. Nevertheless, our results are mostly consistent with the hypothesis of pollinator‐mediated competition shaping the segregated flowering pattern reported previously for this temperate plant assemblage. The mechanisms likely involve effects on male function, whereby pollen‐transport loss during heterospecific flower visits limit pollen export, and more variable effects on female function through pollen limitation.     

Dose-response relationships between pollination and fruiting refine pollinator comparisons for cranberry (Vaccinium macrocarpon [Ericaceae])

Comparisons of pollinator efficacy using pollen received on stigmas can be refined by incorporating experimental dose-response relationships for pollen deposition and fruiting responses. A range of discrete pollen doses applied to cranberry stigmas resulted in decelerating curvilinear responses for fruiting, berry size, and seed set. Minimum thresholds and maximum asymptotes bounded reproductive responses to incremental stigmatic pollen loads. Four bee species were compared for their pollination efficacies on commercial cranberries, using counts of pollen received by stigmas during single bee visits to previously virgin flowers. Differences between these bee species were found to be exaggerated when raw pollen counts were used for comparison because foragers of some species often delivered pollen in excess of that needed to maximize fruit and seed production. Sixfold differences between species in mean pollen deposition translated into 1.5-2-fold differences in predicted cranberry fruit set and size. Implications for pollen tube competition and agricultural production are discussed.     

What We Think We Know vs. What We Need to Know About Orchid Pollination and Conservation: <Emphasis Type="Italic">Cypripedium</Emphasis> L. as a Model Lineage

While Darwin (1862, 1877) showed that reproductive success in orchid populations depended on adaptive floral morphology coupled with pollinator visitation a more recent review of the literature (Tremblay et al., 2005) confirmed that many out-breeding species are pollinator-limited because most orchid species showing low fecundity also lack rewards. The absence of rewards depresses both pollinator fidelity and the frequency of pollinator visits to an orchid population even though orchid flowers that lack rewards retain the same interlocking floral structures for precise pollinia removal and deposition found in related species that offer rewards. Using the genus, Cypripedium, as a model lineage of non-rewarding flowers this study also shows that the correlation between low fruit set in a Cypripedium sp. and its specific pollinator(s) is insufficient to predict specific frequencies of low fecundity. Annual rates of fruit set often vary broadly between populations of the same species and within the same population over several seasons. We speculate that fruit-set rates also decline when orchid demography and additional biotic and abiotic factors interrupt rates of pollinator activity (pre-zygotic) and fertilization/fruit maturation (post-zygotic). We suggest that that traditional field studies on pollination ecology and breeding systems be combined with data sets recording genetic variation and orchid flower demography in relation to seasonal variation in climate. We also propose that the same information be collected in regard to genetic variation, demography and phenology of populations of known orchid pollinators and co-blooming angiosperm species native to orchid habitats.