Seasonal complementary in pollinators of soft-fruit crops

Understanding the relative contributions of wild and managed pollinators, and the functional contributions made by a diverse pollinator community, is essential to the maintenance of yields in the 75% of our crops that benefit from insect pollination. We describe a field study and pollinator exclusion experiments conducted on two soft-fruit crops in a system with both wild and managed pollinators. We test whether fruit quality and quantity is limited by pollination, and whether different pollinating insects respond differently to varying weather conditions. Both strawberries and raspberries produced fewer marketable fruits when insects were excluded, demonstrating dependence on insect pollinators. Raspberries had a short flowering season which coincided with peak abundance of bees, and attracted many bees per flower. In contrast, strawberries had a much longer flowering season and appeared to be much less attractive to pollinators, so that ensuring adequate pollination is likely to be more challenging. The proportion of high-quality strawberries was positively related to pollinator abundance, suggesting that yield was limited by inadequate pollination on some farms. The relative abundance of different pollinator taxa visiting strawberries changed markedly through the season, demonstrating seasonal complementarity. Insect visitors responded differently to changing weather conditions suggesting that diversity can reduce the risk of pollination service shortfalls. For example, flies visited the crop flowers in poor weather and at the end of the flowering season when other pollinators were scarce, and so may provide a unique functional contribution. Understanding how differences between pollinator groups can enhance pollination services to crops strengthens the case for multiple species management. We provide evidence for the link between increased diversity and function in real crop systems, highlighting the risks of replacing all pollinators with managed alternatives.     

Comparison of two cacao (Theobroma cacao L.) clones for the effect of pollination intensity on fruit set and seed content

We compared the influence of pollination intensity (PI) on fruit set and seed number per pod in two cacao (Theobroma cacao L.) clones, IFC5 (Forastero Lower-Amazon Amelonado), which is self-compatible and known to produce a high number of seeds per pod under open pollination, and SCA6 (Forastero Upper-Amazon), which is self-incompatible and known to produce a low number of seeds per pod under open pollination. With both clones, PI had a positive effect on fruit set, with the maximum rate requiring more than 150 pollen grains per pod. One-half of the maximum rate of fruit set was reached with 39 pollen grains per pod for SCA6, and 78 for IFC5. With SCA6, a significant positive effect of PI on seed number per pod was also observed, with maximum seed number requiring more than 200 pollen grains per pod. In contrast, seed numbers with IFC5 were approximately equal over the PI range 100–800 pollen grains per pod. Patterns of seed number per pod were compared after: (1) open insect pollination, (2) low-intensity hand pollination and (3) high-intensity hand pollination. The patterns obtained with IFC5 under open pollination showed a peak of 40–50 seeds per pod, whereas the distribution patterns were more even with SCA6. The pattern of seed number under open pollination was similar to that obtained with high-intensity hand pollination for IFC5, and with low-intensity hand pollination for SCA6. We concluded that the high number of seeds per pod observed with IFC5 under natural insect pollination may be explained by a high number of compatible self-pollen grains on the stigma and by a severe drop of low-pollinated flowers eliminating potential lowfilled fruits. With SCA6, however, the number of compatible pollen grains deposited on the stigma was probably low under open pollination, and the flowers required lower pollen quantities to set fruit, which resulted in a high frequency of low seed numbers per fruit. This difference in the capacity to set low-seeded fruits might be considered as an adaptive trait related to the mode of reproduction.     

Impact of insect pollinators on yield and fruit quality of strawberry

Investigations were conducted to determine the insect pollinators visiting strawberry blossoms and their impact on fruit production. Various pollinators observed during the blooming period of strawberry were viz. Apis mellifera, Apis cerana, Apis florea. Apis dorsata, soil nesting solitary bees such as Andrena leana and A. ilerda, butterflies, houseflies, syrphid flies and some beetles. The percentage of fruit set was much higher in open pollinated plants than control. There was 11.20 per cent malformed fruit in open pollinated plots as compared to 17.44 per cent in controlled one. Further the fruits obtained from the open pollinated plants were of good quality and large as compared to the controlled plants. Growers are recommended to take advantage of the several pollinators, either the honey bee or the native pollinators (Syrphidae and native bees). The importance of diversifying pollination sources, avoiding the dependence on a single specific group is stressed. This study also suggests measures which envisage the conservation, establishment and increase of native pollinators’ populations in the typical agro-ecosystem of region.     

Interacting effects of grass height and herbivores on the establishment of an encroaching savanna shrub

Shrub encroachment is a widely observed problem in Southern African savannas. Although the effects of herbivory and grass height on woody species recruitment have been studied individually, little information exists about how these factors interact. In this study seeds and seedlings of the encroaching shrub Dichrostachys cinerea were planted in clipped and unclipped grass plots, with and without large herbivores present. Seed germination, seedling survival and seedling predation were monitored for 8 months. Germination started earlier in plots where herbivores were excluded. Overall, the earlier the seeds germinated, the longer the seedlings survived. Clipping positively affected the number of germinated seeds, seedling growth and survival but effects varied among herbivore exclusion treatments and sites. Invertebrates caused the majority of the seedling damage. We conclude the recruitment of D. cinerea is influenced by the interplay of grass height and herbivory. In this study, the presence of large herbivores early in the wet season, and the absence of simulated grazing later on, affected the regeneration of D. cinerea negatively. However, differences in effects among sites suggest that the mechanisms found here may work differently in other habitats.      

Flowering Phenology, Pollination, and Fruit Set of Cypripedium macranthos var. rebunense, a Threatened Lady's Slipper (Orchidaceae)

Cypripedium macranthos var. rebunense, was preliminarily studied on Rebun Island, Hokkaido, Japan, for conservation of this threatened orchid. The functional longevity of individual flowers varied from 0 to 18 days, a determining factor being the wind conditions at the growing sites. The flower was self-compatible but needed a pollen vector for fruit production. It lacked nectar and adopted a deceptive pollination system. Queens of a bumblebee, Bombus pseudo-baicalensis, were found to be pollinators but their visits were infrequent. Fruit set was 8.3% at one study site but only 1.2% at another, probably due to interference from the many visitors who came to see the flowers. Based on the results, conservation and management of lady's slipper were discussed. The need for less human interference and for preservation of the interactions between lady's slipper and its pollinator or other plant species with floral rewards is stressed. Received 1 November 2000/ Accepted in revised form 15 March 2001     

Impact of two specialist insect herbivores on reproduction of horse nettle,Solanum carolinense

The frequency of coevolution as a process of strong mutual interaction between a single plant and herbivore species has been questioned in light of more commonly observed, complex relationships between a plant and a suite of herbivore species. Despite recognition of the possibility of diffuse coevolution, relatively few studies have examined ecological responses of plants to herbivores in complex associations. We studied the impact of two specialist herbivores, the horse nettle beetle, Leptinotarsa juncta, and the eggplant flea beetle, Epitrix fuscula, on reproduction of their host, Solanum carolinense. Our study involved field and controlled-environment experimental tests of the impact on sexual and potential asexual reproduction of attack by individuals of the two herbivore species, individually and in combination. Field tests demonstrated that under normal levels of phytophagous insect attack, horse nettle plants experienced a reduction in fruit production of more than 75% compared with plants from which insects were excluded. In controlled-environment experiments using enclosure-exclosure cages, the horse nettle's two principal herbivores, the flea beetle and the horse nettle beetle, caused decreases in sexual reproduction similar to those observed in the field, and a reduction in potential asexual reproduction, represented by root biomass. Attack by each herbivore reduced the numbers of fruits produced, and root growth, when feeding in isolation. When both species were feeding together, fruit production, but not root growth, was lower than when either beetle species fed alone. Ecological interactions between horse nettle and its two primary herbivores necessary for diffuse coevolution to occur were evident from an overall analysis of the statistical interactions between the two herbivores for combined assessment of fruit and vegetative traits. For either of these traits alone, the interactions necessary to promote diffuse coevolution apparently were lacking.     

Effects of florivory on floral volatile emissions and pollination success in the wild parsnip

In addition to reducing fitness by consuming reproductive structures, florivores may also reduce plant fitness by altering interactions with pollinators. To date, the effects of florivore activity on the volatile profile of flowers and subsequent attractiveness to pollinators have not been extensively investigated. In this study, we had three specific objectives: to determine the impact of florivory by the parsnip webworm Depressaria pastinacella on the floral volatile profile of the wild parsnip Pastinaca sativa, to ascertain the mechanisms by which florivory changes the volatile profile, and to estimate the consequences of florivory on visitation by pollinators and eventual seed set. An overall indirect effect of webworms on seed set, that is, the effect of infestation on pollination success, was not detected. However, this overall lack of indirect effect masks the heterogeneity among individual plants. For seven of 14 plants examined, pollination success was altered by webworms, and in four of these plants the alteration in pollination success was consistent with webworm-altered visitation. Webworms significantly altered floral fragrance, in particular causing disproportionate increases in the emissions of octyl esters. Additionally, volatiles from webworm frass, which contains large amounts of the octyl ester metabolite n-octanol, may alter the floral fragrance in ways that change attractiveness of flowers to pollinators. This study suggests that the effects of florivores on plant fitness are not limited to the removal of floral units but may also involve alterations in floral volatile composition, through damage-induced release and detoxification of particular constituents, that affect visitation and pollination success. Handling Editor: Steve Johnson. An erratum to this article can be found at     

梨不同品种花粉生活力测定及授粉特性研究

以19个梨品种为试材,用离体培养法测定在不同贮藏温度下花粉生活力,并调查6个杂交组合和15个梨品种自交的结实特性。结果表明,在培养基中加入硼酸和赤霉素可提高花粉生活力;通过镜检发现大慈梨、八月红、黄金梨3个品种花粉败育;在不同的贮藏条件下,温度越低花粉发芽率下降越缓慢,因此在低温条件下适合花粉长久贮藏;在贮藏过程中,均存在"短期被迫休眠现象",且随温度的降低被迫休眠恢复时间向后延伸。6个杂交组合的花朵坐果率和花序坐果率平均值为82.54%和94.82%,与自然授粉没有明显差异,因此金二十世纪可以作为鸭梨、雪花梨的授粉树,雪花梨、鸭梨、红安久、红茄梨可以作为黄金梨的授粉树;15个梨品种自交结实率较低,而大慈梨、八月红、黄金梨、秦丰、秀玉、新星、雪花梨、丰水8个品种不结实,在生产上均需配置授粉树才能达到产量需求。     

Effect of pollination time,the hour of daytime,pollen storage temperature and duration on pollen viability,germinability, and fruit set of date palm (Phoenix dactylifera L.) cv "Deglet Nour"

Success artificial pollination with viable pollen is crucial process in the production chain of date palms. This study evaluated the impact of pollen storage temperature and duration, pollination time following spathe cracking, and the hour of daytime on pollen viability, germinability, fruit set and yield of 'Deglet Nour' date palm cultivar. In in vitro tests, fresh pollen showed the maximum viability (96.3%) and germination (85%) but it decreased thereafter upon the storage temperature (28, 4 and ?30 °C) and duration (3, 6, 9 and 12 months). In this respect, pollen stored at ?30 °C retained highest viability and germinability followed by those stored at 4 and then at 28 °C. In filed experiments, fruit set was 85, 75, 65, and 45% with pollination using fresh pollen, or pollen stored at ?30, 4 and 28 °C, respectively. Fruit set was 95%, 75%, and less than 50%, for pollination performed on the same day of spathe cracking, 6 and 12 days later, respectively. The highest fruit set percentage and yield/bunch were obtained with pollination performed between 12.0 pm and 15.0 pm in contrast to 8.0–11.0 am or 16.0–17.0.     

Large herbivores affect forest ecosystem functions by altering the structure of dung beetle communities

Dramatic increases in populations of large mammalian herbivores have become a major ecological issue, particularly in the northern hemisphere, due to their substantial impacts on both animal and plant communities through processes such as grazing, browsing, and trampling. However, little is known about the consequences of these population explosions on ecosystem functions. Here, we experimentally investigated how the population density of sika deer (Cervus nippon) in temperate deciduous forest areas in Japan affected the decomposition of mammal dung by dung beetles, which is a key process in forest ecosystems. We measured a range of environmental variables (e.g., vegetation cover, soil hardness) and the dung decomposition rate, measured as the amount of deer dung decomposed during one week, and sampled dung beetles at 16 study sites with three different deer densities (high/intermediate/low). We then used structural equation modeling to investigate the relationships between deer density, environmental variables, the biomass of dung beetles (classified into small or large species), and the dung decomposition rate. We found that the biomass of small species increased with increasing deer density, whereas that of large species was not related to deer density. Furthermore, the dung decomposition rate was positively related to the biomass of small species but unrelated to that of large species. Overall, our results showed that an increase in deer density affects the decomposition rate of mammal dung by changing the structure of dung beetle communities (i.e., increasing the number of small dung beetles). Such an understanding of how increases in large herbivore populations affect ecosystem functions is important for accurately evaluating the ecological consequences of their overabundance and ultimately managing their populations appropriately.     

Metabolomic analysis of the interaction between plants and herbivores

Insect herbivores by necessity have to deal with a large arsenal of plant defence metabolites. The levels of defence compounds may be increased by insect damage. These induced plant responses may also affect the metabolism and performance of successive insect herbivores. As the chemical nature of induced responses is largely unknown, global metabolomic analyses are a valuable tool to gain more insight into the metabolites possibly involved in such interactions. This study analyzed the interaction between feral cabbage (Brassica oleracea) and small cabbage white caterpillars (Pieris rapae) and how previous attacks to the plant affect the caterpillar metabolism. Because plants may be induced by shoot and root herbivory, we compared shoot and root induction by treating the plants on either plant part with jasmonic acid. Extracts of the plants and the caterpillars were chemically analysed using Ultra Performance Liquid Chromatography/Time of Flight Mass Spectrometry (UPLCT/MS). The study revealed that the levels of three structurally related coumaroylquinic acids were elevated in plants treated on the shoot. The levels of these compounds in plants and caterpillars were highly correlated: these compounds were defined as the ‘metabolic interface’. The role of these metabolites could only be discovered using simultaneous analysis of the plant and caterpillar metabolomes. We conclude that a metabolomics approach is useful in discovering unexpected bioactive compounds involved in ecological interactions between plants and their herbivores and higher trophic levels.     

Relative importance of diurnal and nocturnal pollinators for reproduction in the early spring flowering shrub Stachyurus praecox (Stachyuraceae)

Generalized pollination systems may be favored in early spring flowering plants, as during this period pollinator activity is unpredictable. Many previous studies have concentrated on the importance of diurnal visitors in pollination, and consequently, information on the contribution of nocturnal visitors to pollination in early spring is limited. This study was conducted to evaluate the relative importance of diurnal and nocturnal pollinators in the early spring flowering dioecious shrub Stachyurus praecox (Stachyuraceae), in two temperate forests in central Japan. Visitors to the female and male flowers were observed during day and night, and their relative contributions to seed set were compared. The pollinator observations revealed that the diurnal and nocturnal insects visited both male and female flowers, and that the main flower visitors were diurnal small bees and flies as well as nocturnal settling moths. The diurnal and nocturnal flower visitors also acted as pollinators, as the pollen grains of S. praecox were attached to the insects collected from the female flowers. Pollination experiments demonstrated that the contributions of diurnal pollinators to the seed set were higher than those of the nocturnal pollinators. The results of this study indicate that S. praecox has a generalized pollination system, comprising both diurnal insects and nocturnal settling moths. Although the roles of diurnal insects are more important in the pollination of S. praecox, nocturnal settling moths may have a complementary role in early spring.     

Population size,pollinator visitation and fruit production in the deceptive orchid Calypso bulbosa

It has been proposed that in non-rewarding animal-pollinated plants the pollination intensity should decrease with increasing population size and should increase with increasing local abundance of reward-producing plants. To test these hypotheses, we examined how population size, local abundance of Salix caprea, and tree cover were related to pollen removal and fruit production in 16 populations of the deceptive, early-flowering and bumblebee-pollinated orchid Calypso bulbosa in northern Sweden in 3 consecutive years. To determine whether fruit production was limited by pollinator visitation, supplemental hand-pollinations were performed in three populations in 3 years. Finally, to examine whether increased fruit production was associated with a reduction in future flower production, vegetative growth or survival, supplemental hand-pollination was repeated for 5 years in one population. The levels of pollen export, pollen deposition, and fruit set of C. bulbosa varied considerably among years and among populations. The proportion of plants exporting pollen was negatively related to population size, and positively related to density of S. caprea and to tree cover in 1 of the 3 years. In the other 2 years, no significant relationship was detected between proportion of plants exporting pollen and the latter three variables. In no year was there a significant relationship between fruit set and population size, density of S. caprea and tree cover. There was substantial among-year variation in the extent to which fruit production was limited by insufficient pollen deposition and in the amount of weather-induced damage to flowers and developing fruits. Fruit set was consistently higher in hand-pollinated than in open-pollinated plants, but this difference was statistically significant in only one of 3 years. Supplemental hand-pollination in 5 consecutive years increased cumulative fruit production 1.8 times, but did not affect flower production, plant size, or survival. Tree cover was negatively correlated with the incidence of frost damage in 1 year. The results indicate that life-time seed production may be pollen limited in C. bulbosa, and that variation in population size and local abundance of the early-flowering, nectar-producing S. caprea can only partly explain the extensive variation in pollinator visitation among populations of this species.     

Effect of pollination intensity on fruit and seed set in cacao (Theobroma cacao L.)

We studied the functional relationship between pollination intensity and fruit survival as well as the number of seeds per pod in the tropical tree Theobroma cacao L. on a Forastero Upper-Amazon clone (UPA 409) in Ivory Coast. Cutting the style 24 h after pollination allowed for counting the number of pollen grains deposited on a stigma without affecting fruit set and seed development. Forty-three pollen grains were necessary to reach 50% of maximum fruit set 28 days after pollination. Above 115 pollen grains, the proportion of developing ovaries reached a maximum of 88% 28 days after pollination and 75% at maturity. With fewer than 238 pollen grains per stigma, there was a close relationship between pollination intensity and number of seeds per pod; the pollenseed ratio increased from 1.61 to 3.81 for PI increasing from 30 to 238 pollen grains. For higher pollination intensities, the average number of seeds per pod reached a maximum of 58. The relationship between pollination intensity and seed content was modelled. Results are consistent with the hypothesis that ovules attracted pollen tubes in a similar way regardless of whether or not they had already been reached by another pollen tube.     

Extinction threats of a narrowly endemic shrub, Stachyurus macrocarpus (Stachyuraceae) in the Ogasawara Islands

Stachyurus macrocarpus is a narrowly endemic endangered shrub of which only one population remains on Chichi-jima Island (Ogasawara). We surveyed its population dynamics and reproductive states for 4 years and analyzed the population viability. In a survey of its entire distribution range, a total of 87 S. macrocarpus individuals have been found and 68 individuals have been confirmed in 2007. Thus, the actual population size is estimated to be less than 100 individuals. Environmental conditions and the results of a hand-pollination experiment suggest that low fruit setting in both sexes (female: 4.9–11.2%; hermaphrodite: 0.0%) may be caused by resource limitation. In addition, fruit predation by alien Rattus rattus was observed despite the rare fruit setting. The small effective population size (23.1–24.6 individuals) and hermaphrodite-biased sex ratio (46 hermaphrodites:12 females) would increase the risk of extinction. The habitat of S. macrocarpus was limited to dense scrubland and forest understory, and seedling regeneration was very scarce. During the survey period, 19 individuals (21.8%) were found dead, there were only two seedling recruitments, and the annual population growth rate was 0.979. The lack of occurrence on the south and west slopes and the shortness of shrubs in open spaces suggests that S. macrocarpus suffers stress from both dryness and frequent typhoons. However, S. macrocarpus also exhibited high mortality of shoot in forest understories. These findings suggest that the suitable habitat of S. macrocarpus is likely to be narrow in Ogasawara. While the recent increase of goat grazing has not affected individual mortality, 58.6% of shoots that had been grazed by goats were dead within 2 years. As elasticity analysis had shown that larger individuals make a greater contribution to the population growth rate, repeated goat grazing would impact the S. macrocarpus population in the near future by decreasing the vitality of larger individuals. Emergency measures for protecting the shrub from goat grazing and reinforcing the population through nursery cultivation were proposed.     

Sex, secondary compounds and asymmetry. Effects on plant–herbivore interaction in a dioecious shrub

We analysed the links between herbivory, anthraquinone content and developmental instability of leaves in Rhamnus alpinus, taking into account possible effects of sexual dimorphism. The amount of leaf loss caused by herbivores averaged 3%, rarely exceeding 25%. Leaf losses were evenly distributed in the shrubs, with highest variability among leaves of the same shoot, thus hiding possible shrub, sex or population effects. This pattern of herbivory implies a shifting of caterpillars from one leaf to another before consuming all readily available material. We suggest that this behaviour might be triggered by a short-term change in leaf palatability by means of an increase in the production of secondary compounds. Supporting this hypothesis, we have found a higher anthraquinone content in damaged leaves compared with undamaged ones. The leaves of male plants exhibited a higher concentration of anthraquinones than those of females, which contrasts with classic hypotheses. We relate this to the lower rate of biomass increase in males, which should allow them to allocate more resources to defence. Leaves showed fluctuating asymmetry (FA), but we did not find any relationship between the degree of asymmetry and sex, herbivory or anthraquinone content at any level considered. Therefore, FA cannot be considered as an indicator of susceptibility to damage by herbivores or of the ability to induce the production of defensive compounds in R. alpinus.     

Fruit maturation patterns of Carya spp. (Juglandaceae): an intra-crown analysis of growth and reproduction

Summary Fruit survival patterns, from fertilization to maturation, were examined for Carya ovata and C. tomentosa in a New Jersey USA forest. We observed fruiting and shoot growth characteristics over a 3-yr period to determine: (1) the patterns of fruit survivorship (from initiation to maturity) within and among years, (2) the relationships between shoot growth, fruit initiation, and fruit survival to maturity, and (3) the influence of phytophagous insects on fruit survival. We found that within years, smaller infructescences (1–2 fruits) exhibited greater relative survivorship than larger ones (3–4 fruits); however, absolute nut production was greatest for mid-sized infructescences (2–3 fruits). Among years, fruit survivorship varied considerably within populations. Across the 3-yr period we observed average fruit survivorship to be convex, linear, and concave, respectively. Likewise, shoot characteristics (length, width, number of leaves) varied concomitantly (decreasing fruit survivorship was accompanied by decreasing shoot length and number of leaves). Within years, we found no strong relationship between shoot characteristics and infructescence size and survival. The patterns of tree-to-tree variation suggested a strong genetic basis to shoot growth and fruit maturation. However, patterns of variation within and among years also indicated a strong environmental influence on these traits as well. Natural phytophagy by insects was observed to be low (<5%); however, shoot defoliations of 10–25% were not uncommon. Experimental defoliations (ambient, 10–15%, 20–40%, and 75–100%) did not result in reduced survival to maturity. Collectively, the data suggest that year-to-year variability in shoot growth has a greater influence on fruit maturation patterns than within year fruit-shoot relations.     

Optimal foraging and community structure: implications for a guild of generalist grassland herbivores

Summary A particular linear programming model is constructed to predict the diets of each of 14 species of generalist herbivores at the National Bison Range, Montana. The herbivores have body masses ranging over seven orders of magnitude and belonging to two major taxa: insects and mammals. The linear programming model has three feeding constraints: digestive capacity, feeding time and energy requirements. A foraging strategy that maximizes daily energy intake agrees very well with the observed diets. Body size appears to be an underlying determinant of the foraging parameters leading to diet selection. Species that possess digestive capacity and feeding time constraints which approach each other in magnitude have the most generalized diets. The degree that the linear programming models change their diet predictions with a given percent change in parameter values (sensitivity) may reflect the observed ability of the species to vary their diets. In particular, the species which show the most diet variability are those whose diets tend to be balanced between monocots and dicots. The community-ecological parameters of herbivore body-size ranges and species number can possibly be related to foraging behavior.     

The effect of wild radish floral morphology on pollination efficiency by four taxa of pollinators

The effects of floral morphology on rates of pollen removal and deposition by different pollinators in generalist plant species are not well known. We studied pollination dynamics in wild radish, Raphanus raphanistrum, a plant visited by four groups of pollinators: honey bees, small native bees, butterflies, and syrphyd flies. The effects of anther position and other factors on pollen removal during single visits by all four pollinator taxa were measured. Flowers with high anther exsertion (i.e., anthers placed higher above the opening of the corolla tube) tended to have the highest numbers of pollen grains removed, but this effect was strongest for honey bees and butterflies. For all pollinator taxa, pollen removal increased with the number of pollen grains available on a flower and whowed a positive, decelerating relationship with the duration of the visit. The effects of stigma position and other factors on pollen deposition during single visits by honey bees and butterflies were also studied. The nectar-feeding butterflies had a higher pollination efficiency (percentage of pollen grains removed from anthers that were subsequently deposited on a stigma) than the nectar- and pollen-feeding honey bees. Flowers with intermediate stigma exsertion had the highest numbers of pollen grains deposited on their stigmas by butterflies, but stigma exsertion had no effect on deposition by honey bees. For both butterflies and honey bees, pollen deposition on the recipient flower increased with the amount of pollen removed from the donor flower, and there was a positive, decelerating relationship between deposition and time spent at the flower; these results are analogous to those for pollen removal. The effects of anther and stigma exsertion on pollen removal and denosition did not fit predictions based on patterns of floral correlations, but results for morphology, pollen availability, time spent per visit, and pollinator efficiency are in broad agreement with previous studies, suggesting the possible emergence of some general rules of pollen transfer.