A ballistic pollen dispersal system influences pollination success and fruit‐set pattern in pollinator‐excluded environments for the endangered species Synaphea stenoloba (Proteaceae)

The critically endangered Synaphea stenoloba (Proteaceae) has numerous scentless flowers clustered in dense inflorescences and deploys a ballistic pollen ejection mechanism to release pollen. We examined the hypothesis that active pollen ejection and flowering patterns within an inflorescence influence the reproductive success (i.e. fruit formation) of individual flowers within or among inflorescences of S. stenoloba in a pollinator‐excluded environment. Our results showed that: (1) no pollen grains were observed deposited on the stigma of their own flower after the pollen ejection system was manually activated, indicating self‐pollination within an individual flower is improbable in S. stenoloba; (2) fruit set in the indoor open pollination treatment and the inflorescence‐closed pollination treatment indicated that S. stenoloba is self‐compatible and pollen ejection can potentially result in inter‐floral pollination success; (3) fruit set in the inflorescence‐closed pollination treatment was significantly lower than that of indoor open pollination, indicating within‐ and between‐flower pollination events in an inflorescence are most likely limited, with pollination between inflorescences providing the highest reproductive opportunity; and (4) analysis of the spatial distribution of cumulative fruit set on inflorescences showed that pollen could reach any flower within an inflorescence and there was no functional limitation on seed set among flowers located at various positions within the inflorescence. These data suggest that the pollen ejection mechanism in S. stenoloba can enhance inter‐plant pollination in pollinator‐excluded environments and may suggest adaptation to pollinator scarcity attributable to habitat disturbance or competition for pollinators in a diverse flora. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 59–68.     

The relative importance of birds and bees in the pollination of Metrosideros excelsa (Myrtaceae)

Exclusion experiments were used to assess the effect of different pollinator groups on outcrossing and seed production in Metrosideros excelsa. The main study site was Little Barrier Island, New Zealand where indigenous bird and native solitary bees are the main flower visitors. Our results showed that native birds were more important pollinators of M. excelsa than native bees. Seed production was much higher in open pollination than in two exclusion experiments where either birds were excluded and native bees only had access to flowers, or where all pollinators had been excluded. The number of fertile seeds per capsule was 45% higher after open pollination than in treatments with bee visitation only and 28% higher than in treatments where all flower visitors were excluded. Estimated outcrossing rates were significantly higher (t_m = 0.71) for open pollination in the upper canopy (>4 m above‐ground level) where bird visitation is presumed to be more frequent than for a treatment with native bee access only (t_m = 0.40). Our results also suggest that a large proportion of seeds (66%) arise from autonomous self‐pollination when all pollinators are excluded. In four trees of a modified mainland population with predominantly introduced birds and a mixture of introduced and native bees there was no decrease in seed production for the treatment allowing bee access only, indicating that – in contrast to native bees – honeybees may be more efficient pollinators of M. excelsa. Observation of the foraging behaviour of both groups of bees showed that native bees contact the stigma of flowers less frequently than honeybees. This is likely to be a consequence of their smaller body size relative to honeybees.     

Effectiveness of short‐tongued bees as pollinators of apparently ornithophilous New Zealand mistletoes

Abstract The flowers of two species of threatened New Zealand mistletoes (Peraxilla tetrapetala and Peraxilla colensoi, Loranthaceae) have explosive buds that do not open unless force is applied by birds or two species of native short‐tongued bees. Opened flowers are visited by a variety of birds and insects. Although both species of Peraxilla conform to a pollination syndrome of ornithophily, bees may be effective alternative pollinators. We investigated the effectiveness of bees and birds as pollinators of P. colensoi at one site and P. tetrapetala at two sites in the South Island. Bees and other insects outnumbered birds as flower visitors at all three sites. By excluding birds with wire cages, we showed that two bee species regularly open flowers of P. tetrapetala, but only rarely open flowers of P. colensoi. Few pollen grains were deposited when either birds or bees opened buds, so opening buds was not by itself sufficient for adequate pollination. Instead, pollen continued to accumulate over the next 6 or 7 days, even inside cages that excluded birds. Both populations of P. tetrapetala were regularly pollen‐limited, but in different ways. At Ohau, opened flowers gained enough pollen to produce seeds, but many buds were not opened and hence failed to set seed. In contrast, at Craigieburn, nearly all buds were opened, but many of these did not receive enough pollen. These results demonstrate that native bees can partially replace birds as pollinators of mistletoes, despite their apparent ornithophilous syndrome. Ongoing reductions in New Zealand forest bird numbers means that the service bees provide may be important for the long‐term future of these plants.     

Severe pollen limitation in populations of the New Zealand shrub Alseuosmia macrophylla (Alseuosmiaceae) can be attributed to the loss of pollinating bird species

The loss of bird species following human colonization of New Zealand has raised concerns about the consequences for crucial ecosystem functions such as pollination. The understorey shrub Alseuosmia macrophylla (Alseuosmiaceae) exhibits characteristics typical of a bird pollination syndrome, but populations still persist in northern North Island forest remnants despite the local extinction of most endemic bird pollinators, leading to the suggestion that moths – rather than birds – may be the primary pollinators. The aim of this study was to quantify the importance of endemic birds as pollinators of A. macrophylla over several years by comparing plants on Little Barrier Island (LBI), where all extant endemic bird pollinators still occur, to plants at sites on the adjacent North Island in the Waitakere Ranges (WTK), where only one of these species remains common. Flowers on LBI were visited by endemic bellbirds (Anthornis melanura) and stitchbirds (Notiomystis cincta), while at WTK sites the most common visitors were the recently arrived silvereye (Zosterops lateralis) and the introduced honeybee (Apis mellifera), both of which acted principally as nectar robbers. Caged flowers on LBI had significantly lower fruit set than open flowers, and plants at WTK were significantly more pollen‐limited than plants on LBI. This provides evidence that the loss of endemic pollinating birds is the most likely reason for the high pollen limitation found in some North Island A. macrophylla populations, and the very low seed set of these populations could have serious implications for the long‐term persistence of this species.     

Effect of pollen and resource limitation on reproduction of Zygophyllum xanthoxylum in fragmented habitats

Limitations on pollen and resources may significantly affect plant reproduction in fragmented habitats. In this study, phenology and pollinator frequency and activity were investigated to estimate the role of pollinators in Zygophyllum xanthoxylum reproduction, and this species is ecologically important in northwest China. In addition, the relative impact of restrictive amounts of pollen and resources on the seed set per flower was evaluated. It was found that adding pollen boosted the size of the seed set per flower, but had no significant effect on the number of flowers. By contrast, the addition of resources increased flower numbers as well as had a slight impact on the seed set per flower. These results indicate the amount of available pollen is a limiting factor for reproductive success. Moreover, Apis mellifera was identified as the most effective pollinator of Z. xanthoxylum, and there were more overall pollinators and visitations in the control than in the fragmented habitats. Furthermore, the limitations in pollen were more restrictive in the fragmented area than in the control. This was due to increased pollinator visitations in the control that could ameliorate the effects of lower pollen levels. When there is a limited availability of suitable pollinators, self‐pollination is critical in fragmented habitats. Z. xanthoxylum has reproductive strategies that aid in adapting to harsh environments, including protogyny and delayed selfing.     

Pollination ecology of the pre-Pyrenean endemic Petrocoptis montsicciana (Caryophyllaceae): effects of population size

Petrocoptis montsicciana (Caryophyllaceae) is a threatened pre‐Pyrenean endemic that grows exclusively on caves and walls of limestone. We studied its pollination ecology by monitoring phenology and by evaluating pollen and nectar production, pollinator activity (frequency and behaviour of visitors), quantity and quality of pollination services, pollen/ovule ratio, and seed set in response to insect exclusion and self‐compatibility tests. We also analysed the effect of population size on reproductive mechanisms by comparing a large and a small population. Flowers of P. montsicciana produced nectar and were visited by Hymenoptera (79.7%), Diptera (11.5%), and Lepidoptera (8.8%). The most frequent pollinators (60.6% of total visits) were long‐tongued bees of the genus Anthophora. Both populations had a similar range of pollinators. We found a correlation between the number of visited flowers and the number of open flowers per census; 88.7% of pollen grains deposited on the stigmas were conspecific and the main competitor was another chasmophyte plant, Antirrhinum molle. Bagged flowers set seeds but significantly less so than hand‐self‐pollinated and control flowers. Thus, although self‐compatible and self‐pollinated, entomophilous pollination of P. montsicciana is required in order to explain c. 10–40% of total seed set, in accordance with P/O ratio estimations. Bagged flowers from the small population set significantly more seeds than the large one. Visitation rates were lower in the small population, but, unexpectedly, showed higher stigmatic pollen loads and similar or higher seed set. These results suggest an increase of spontaneous selfing rates in the small population, probably favoured by a smaller flower size, which can not only assure reproductive success when pollinators are scarce, but also provide additional potential to adapt to climatic changes. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 76 , 79–90.     

The effect of petal size manipulation on pollinator/seed-predator mediated female reproductive success of Hibiscus moscheutos

The effects of petal-size manipulations on the behavior of pollinators and pollen/seed predators, and on pollen removal and deposition, were studied in Hibiscus moscheutos (Malvaceae) populations. The ultimate effects on the female reproductive success of flowers, such as fruit set, seed predation rate, and final seed set were also measured. We applied three levels of petal removal (100%, 50%, and 0% size reduction in radius) to flowers in natural populations. Two pollinators (Bombus pennsylvanicus and Ptilothrix bombiformis) ignored flowers without petals, suggesting that pollinators use petals as a visual cue to locate flowers. Consequently, 100% petal removal reduced female reproductive success considerably, mainly through a higher rate of fruit abortion due to failure of pollen deposition on stigmas. No significant differences between the 50% petal removal treatment and uncut control were detected in any components of female success examined. The results, therefore, suggest that differences in petal size have little influence on female reproductive success of Hibiscus flowers at our study site. Final seed set varied considerably depending on the larval densities of two coleopteran seed predators (Althaeus hibisci and Conotrachelus fissunguis). A. hibisci responded to petal size, and a higher density of adults was found in flowers in which petal size had not been reduced. Because Althaeus feed on pollen as adults and no effect of petal size on seed predation was detected, the preference of Althaeus for larger flowers may represent a foraging strategy for adult beetles and may exert counteracting selection pressure on petal size through male reproductive success of flowers. Received: 30 November 1997 / Accepted:12 June 1998     

Evolution of oviposition strategies and speciation in the globeflower flies Chiastocheta spp. (Anthomyiidae)

Trollius europaeus (Ranunculaceae) is involved in an intimate interaction with several species of Chiastocheta flies (Anthomyiidae) that are both seed predators and pollinators. In this paper, we analyse the oviposition strategy of the six Chiastocheta species found to coexist on T. europaeus in 19 populations from the French Alps. We show that the species are not equivalent in their oviposition behaviour: C. rotundiventris usually deposits no more than one egg per flower in first-day flowers whereas C. dentifera aggregates its eggs on fruits and thus does not contribute to pollination at all; the four remaining species deposit eggs sequentially during the flowering period from the 2nd to the 7th day. Hence, the outcomes of the interaction in terms of net seed production for the plant greatly depend on the Chiastocheta species visiting it, ranging from a mutualistic to a purely parasitic interaction. We assessed mitochondrial divergence between Chiastocheta spp. by sequencing a 1320-bp mitochondrial DNA fragment. The low divergence observed between species (0–4.15%) suggests that genus diversification took place recently. Unlike in other plant–insect systems where diversification is usually thought to be driven by cospeciation or host shifts, we propose that Chiastocheta speciation took place within the host plant. Basal separation of a particularly mutualistic species provided favourable conditions for plant specialization on this seed-parasite as a pollinator early in the evolution of the association. The parasitic species ovipositing on fruits derived from a species ovipositing on flowers. Diversification of the intermediate strategies probably occurred in relation with the Pleistocene climatic events, reproductive isolation between species being reinforced by niche partitioning for oviposition and/or sexual selection.     

Asymmetrical nature of the Trollius–Chiastocheta interaction: insights into the evolution of nursery pollination systems

The mutualistic versus antagonistic nature of an interaction is defined by costs and benefits of each partner, which may vary depending on the environment. Contrasting with this dynamic view, several pollination interactions are considered as strictly obligate and mutualistic. Here, we focus on the interaction between Trollius europaeus and Chiastocheta flies, considered as a specialized and obligate nursery pollination system – the flies are thought to be exclusive pollinators of the plant and their larvae develop only in T. europaeus fruits. In this system, features such as the globelike flower shape are claimed to have evolved in a coevolutionary context. We examine the specificity of this pollination system and measure traits related to offspring fitness in isolated T. europaeus populations, in some of which Chiastocheta flies have gone extinct. We hypothesize that if this interaction is specific and obligate, the plant should experience dramatic drop in its relative fitness in the absence of Chiastocheta. Contrasting with this hypothesis, T. europaeus populations without flies demonstrate a similar relative fitness to those with the flies present, contradicting the putative obligatory nature of this pollination system. It also agrees with our observation that many other insects also visit and carry pollen among T. europaeus flowers. We propose that the interaction could have evolved through maximization of by‐product benefits of the Chiastocheta visits, through the male flower function, and selection on floral traits by the most effective pollinator. We argue this mechanism is also central in the evolution of other nursery pollination systems.     

Pollinator‐mediated phenotypic selection does not always modulate flower size and number in the large‐flowered Mediterranean shrub Cistus ladanifer (Cistaceae)

Flower size and number usually evolve under pollinator‐mediated selection. However, hot, dry environments can also modulate display, counteracting pollinator attraction. Increased pollen deposition on larger flower displays may not involve higher female fitness. Consequently, stressful conditions may constrain flower size, favouring smaller‐sized flowers. The large‐flowered, self‐incompatible Mediterranean shrub Cistus ladanifer was used to test that: (1) this species suffers pollen limitation; (2) pollinators are spatially–temporally variable and differentially visit plants with more/larger flowers; (3) increased visits enhance reproduction under pollen limitation; (4) stressful conditions reduce female fitness of larger displays; and (5) phenotypic selection on floral display is not just pollinator‐mediated. We evaluated pollen limitation, related floral display to pollinator visits and fruit and seed production and estimated phenotypic selection. Flower size was 7.2–10.5 cm and varied spatially–temporally. Visitation rates (total visits/50 min) ranged from 0.26 to 0.43 and increased with flower size. Fruit set averaged 80% and seed number averaged 855, but only fruit set varied between populations and years. Selection towards larger flowers was detected under conditions of pollen limitation. Otherwise, we detected stabilizing selection on flower size and negative selection on flower number. Our results suggest that selection on floral display is not only pollinator‐dependent through female fitness in C. ladanifer. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 540–555.     

Pollination and fruit‐set of Grevillea robusta in western Kenya

The floral visitors of silky oak, Grevillea robusta A. Cunn. ex R.Br., their foraging behaviour and their effects on fruit‐set were studied at Malava, western Kenya. Grevillea robusta is a popular tree for farm plantings in the eastern and central African highlands. Yield of seed has been disappointingly low in some areas and a lack of appropriate pollinators has been suggested as a possible cause. Investigations involved the monitoring of visitors on active inflorescences, assessment of the rewards available to potential pollinators, and exclusion experiments to establish the effects of various visitors on fruit‐set. The flowers are visited mainly by birds and insects. The likely pollinators of G. robusta are sunbirds (Nectarinia amethystina, N. cyanolaema, N. olivacea, N. superba and N. venusta) and white‐eyes (Zosterops kikuyuensis and Z. senegalensis). Very little aggressive behaviour between birds was recorded. No nocturnal pollinators were observed. Nectar was the major floral reward for pollinators, but is likely depleted by ants and honey bees, the foraging behaviour of which confirmed them to be nectar‐robbers. These insects hardly ever touched stigmas during their visits. Eighty‐nine per cent of bird visits were in the morning (07.00–10.00 hours) when nectar volume was highest. Inflorescences bagged to exclude birds set no fruits, and unmanipulated flowers and flowers bagged with self‐pollen set no fruits, indicating a self‐incompatibility mechanism. Control cross‐pollinated flowers displayed greatly increased fruit‐set (25.1%) compared with natural open‐pollination (0.9%). All these findings confirm the importance of cross‐pollen transfer to flowers and the necessity of pollinators for fruit‐set. Effective seed production requires activity of pollinators for self‐pollen removal and cross‐pollen deposition. Seed production stands for G. robusta should be established where flowering is prolific and bird pollinators are abundant.     

Interaction between floral color change and gender transition in the protandrous weed Saponaria officinalis

Natural selection has directed the evolution of floral traits so that pollinator visits are manipulated to maximize the fitness of individual plants by directing which other individual sires its seeds. In some plants, flowers change color over time and may have the ability to direct pollinators to rewarding flowers. In addition, by varying when pollen is available and when stigmas are receptive, protandrous plants can show variation in selfing rates. In this study, the association between color change and gender transition in flowers of Saponaria officinalis was examined. Anthocyanins were extracted from flowers of each gender stage to measure color using spectrophotometry. Female‐phase flowers were found to have significantly higher anthocyanin concentration than male‐phase flowers in both natural populations and experimental plots. This color change corresponded to a decrease in male sexual function, which was measured by the percentage of pollen grains stained as viable by lactophenol aniline blue and germinated on Brewbaker–Kwack media. Color change was phenotypically plastic. Plants grown in full sun had a more extensive color change than those grown in shaded experimental plots, and this effect was reversed the following year when the shading was removed. Pollinator observations documented both diurnal and nocturnal insect visitation. Fruit and seed set were equivalent on inflorescences bagged during daylight versus night, indicating that both diurnal and nocturnal insects are effective pollinators. If pollinators discriminate based on color, this could potentially reduce within‐plant floral visits and also geitonogamy. This study is the first to document flower color change and moth pollination in Saponaria officinalis.     

Variation in predation costs with Chiastocheta egg number on Trollius europaeus: how many seeds to pay for pollination?

1. In obligate plant/seed parasite–pollinator mutualisms, the plant is exclusively pollinated by an insect whose larvae are specific seed predators. Hence, outcomes of the interaction for the plant can vary with the number of eggs laid and the number of seeds eaten per larva. 2. In the work reported here, predation by Chiastocheta larvae on seeds of Trollius europaeus was analysed as a function of the number of eggs laid on the flower. Flowers with an increasing number of eggs were bagged in three populations and seeds were counted after the end of larval predation, in order to assess whether there was competition among larvae. 3. Seed predation on single‐egg flowers was high and variable (mean per population ranging from 15 to 40% of the developed seeds). Seed predation increased weakly with increasing egg load and was lower than gross seed production (always < 85%) whatever the number of eggs laid. This corresponds to a strong decrease in seed consumption per larva with increasing egg load, i.e. severe larval competition for resources. 4. The results suggest that both interference among Chiastocheta larvae and carpel dehiscence may protect T. europaeus seeds from total predation. Estimates of seed predation based on egg load observed in 20 natural populations in the French Alps typically ranged from 30 to 60%. The interaction was always beneficial for the plant and there was no risk of total seed destruction by Chiastocheta larvae, favouring stability of the mutualism.     

Deconstructing a floral phenotype: do pollinators select for corolla integration in Lavandula latifolia?

An experimental field study was undertaken to assess whether correlational selection generated by pollinators can account for observed phenotypic integration of the two corolla lips in the zygomorphic flowers of the insect‐pollinated Mediterranean shrub, Lavandula latifolia (Labiatae). The experiment consisted of manipulating the upper and lower corolla lips according to a two‐way factorial design with interaction, and involved the complete (lower lip) or extensive (distal two thirds of upper lip) excision of one or the two lips. Per‐flower male (proportion of pollen grains removed) and female (number of pollen tubes down the style) pollination success, and maternal fecundity (seed set), of experimental flowers were evaluated after exposure to natural pollination in the field. Artificially induced corolla variation had no significant effects on either male or female pollination success, and two aberrant floral morphologies, characterized by lack of corolla integration, had a maternal fecundity advantage over normal two‐lipped phenotypes. These results do not support a predominant role of correlational selection by pollinators in maintaining corolla integration in L. latifolia, but are instead consistent with the alternative view that integration is mainly the consequence of genetic and/or developmental factors.     

Pollinator activity and pollination success of Medicago sativa L. in a natural and a managed population

Medicago sativa L. is an important cash crop in the arid region of northwest China. Pollinator activity is an essential aspect of pollination success, but the relationships between pollinator visitation rate and seed set still need further study of M. sativa. We investigated the following characteristics of M. sativa in natural and managed populations: floral traits, pollinator activity, and breeding system. Our results indicated the management could affect the number of flowers produced; however, there was no detectable effect on the seed set per flower. We found the percentage of seeds among pollinated flowers in the managed population was significantly higher than that in the natural population. Moreover, the increase in the proportion of pollinated flowers could significantly increase seed set per flower, and pollinator visitation rate was the important limiting factor for seed set in both populations. Andrena lebedevi Popov was found to be the most frequent pollinator in both populations. Outcrossing was dominant in the breeding system and insect pollination played an important role in outcrossing. Our study suggested that proper management (artificial selection) could promote pollination success of M. sativa.     

Pollinator directionality as a response to nectar gradient: promoting outcrossing while avoiding geitonogamy

Plants with multiple flowers could be prone to autonomous self‐pollination and insect‐mediated geitonogamy, but physiological and ecological features have evolved preventing costs related to autogamy. We studied the rare perennial herb Dictamnus albus as a model plant, with the aim of describing the plant–pollinator system from both plant and pollinator perspectives and analysing features that promote outcrossing in an entomophilous species. The breeding system and reproductive success of D. albus were investigated in experimental and natural conditions, showing that it is potentially self‐compatible, but only intra‐inflorescence insect‐mediated selfing is possible. Nectar analysis showed gender‐biased production towards the female phase, which follows the male phase, and during flowering, full blooming is found in flowers at the bottom of the raceme. Among a wide spectrum of insect visitors, three genera (Bombus, Apis, Megachile) were found to be principal pollinators. A study of insect behaviour showed a tendency towards bottom‐to‐top flights for the most important pollinators Bombus spp. and Apis mellifera: upward movements on the racemes could be explained by foraging behaviour, from more to less rewarding flowers. In accordance with the ‘declining reward hypothesis’, bumblebees and honeybees leave the plant when gain of reward is low, after which few flowers are visited, reducing the chance of self‐pollen transfer among flowers. Intra‐flower self‐pollination is prevented in D. albus by protandry and herkogamy, while the nectar‐induced sequential pattern of pollinator visits avoids geitonogamy and tends to maximise pollen export, promoting outcrossing. All these features for preventing selfing benefit plant fitness and population genetic structure.     

Enhancement of fruit quality in Capsicum annum through pollination by Hypotrigona gribodoi in Kakamega,Western Kenya

A study was carried out in Kakamega forest, in the western region of Kenya, to evaluate the effectiveness of the stingless bee Hypotrigona gribodoi (Magretti, 1884) on the pollination of green pepper. Three treatments were applied and consisted of self‐pollination, pollination by feral pollinators in the open field and pollination by H. gribodoi in a net cage. The differences in fruit yield and seed quality were compared among treatments. Flowers pollinated by H. gribodoi produced the heaviest fruits with the highest seed numbers followed by feral pollinators and lastly self‐pollinated flowers. Moreover, seeds were significantly bigger in size in fruits resulting from flowers pollinated by H. gribodoi compared to fruits obtained from self‐pollinated flowers or flowers pollinated by feral insects. We, therefore, conclude that H. gribodoi is an efficient pollinator of green pepper in the tropical region of East Africa.     

Female reproductive success in gynodioecious Thymus vulgaris: pollen versus nutrient limitation and pollinator foraging behaviour

Gynodioecy is a dimorphic breeding system in which female individuals coexist with hermaphroditic individuals in the same population. Females only contribute to the next generation via ovules, and many studies have shown that they are usually less attractive than hermaphrodites to pollinators. Several mechanisms have been proposed to explain how females manage to persist in populations despite these disadvantages. The ‘resource reallocation hypothesis’ (RRH) states that females channel resources not invested in pollen production and floral advertisement towards the production of more and/or larger seeds. We investigated pollination patterns and tested the RRH in a population of Thymus vulgaris. We measured flower display, flower size, nectar production, visitation rates, pollinator constancy and flower lifespan in the two morphs. In addition, we measured experimentally the effects of pollen and resource addition on female reproductive success (fruit set, seed set, seed weight) of the two morphs. Despite lower investment in floral advertisement, female individuals were no less attractive to pollinators than hermaphrodites on a per flower basis. Other measures of pollinator behaviour (number of flowers visited per plant, morph preference and morph constancy) also showed that pollinators did not discriminate against female flowers. In addition, stigma receptivity was longer in female flowers. Accordingly, and contrary to most studies on gynodioecious species, reproductive success of females was not pollen limited. Instead, seed production was pollen limited in hermaphrodites, suggesting low levels of cross‐pollination in hermaphrodites. Seed production was resource limited in hermaphrodites, but not in females, thus providing support for the RRH. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 395–408.     

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.     

Who can open the flower? Assessment of the flower opening force of mammal‐pollinated Mucuna macrocarpa

Effective pollinators in Fabaceae species with papillionaceous flowers should have sufficient force to open the petals and expose the stamens and pistils from the keel petals. Here, we assessed the force that is required (operative force) to open the petals of the mammal‐pollinated Mucuna macrocarpa, in parallel with estimating the force of bee species. The operative force of M. macrocarpa was quantified by a digital force gauge and the force of three bee species that frequent these flowers was estimated from body mass. The operative force was 1.635 ± 0.146 (mean ± SD) newton (N). This force was far larger than that estimated for the largest bee in the study area. In addition, the operative force of M. macrocarpa was 16–409 times greater than that of bee‐pollinated plants. Our results demonstrate that the operative force of M. macrocarpa flowers is adapted to mammalian openers, preventing potential nectar robbers.