Non-skipper butterflies and skippers had the longest proboscides Table 4. Long-tongued bees were the visitor group with the widest proboscides 0. There was no significant difference in the number of pollen grains extracted in simulated visits of the three other diameters, using fishing line sizes 0. Table 4. Proboscis measurements mean and standard error for flower visitors to A. Proboscis width and length with the same letters are not significantly different with Tukey comparisons.
The visitor group column shows the type of insect that has passed interacted with A. As previously explained in Table 1, the letter "n" represents the number of each type of insect that had an interaction with A. Each of the insects had their proboscis measured, and the average length and width were then taken.
Each type of insect were then compared to determine if any big differences were noticeable. The proboscis compared are more or less similar in size. The Non-Skipper butterflies do have a significantly greater proboscis length than the rest of the insects. Mean and standard error of the number of pollen grains on fishing line of increasing width inserted into flowers of A. The diameter of 0. To aid interpretation, the number of ovules ranges from 46 to 76 ovules per flower. Diameters with the same letters are not significantly different with Kruskal—Wallis analysis.
Using nylon fishing lines to attract visitors, the amount of pollen collected was large along the width of plant. Average of number of pollen grains on the fishing line stimulator of increasing width inserted in flowers of A. Size 0. Mean and standard error of the length of the stigmatic surface stained with methylene blue. Diameters with the same letters are not signifi- cantly different with Tukey comparisons.
The stain is used to measure how much of the thread made contact. In other words, the stain is a visual stand-in for pollen. The mean width of the long-tongued bees' heads was 3. The final test of pollinator effectiveness, whether flowers visited by the different pollinators set fruit, provided clear results. Placing potted plants in the field, observing visits, bagging, tagging and following subsequent fruit set showed that only flowers visited by long-tongued bees set fruit following a single visit.
Of the 44 flowers visited by long-tongued bees, 16 None of the flowers visited by the other pollinator groups 4 by non-skippers butterflies, 19 by skippers and 2 by short-tongued bees produced any fruit. Though the sample sizes were small for non-skipper butterflies and short-tongued bees due to their lower rate of visitation Table 2 , we also found that these groups of visitors do not carry significant amounts of pollen on their proboscides; consequently, we conclude that these two groups of visitors are much less effective pollinators of A.
Pollination efficiency is a function of multiple interacting characters and behaviours, including flower shape and size as well as animal behaviour and morphology Ollerton et al.
In the pine rocklands of South Florida, A. Though many other species visit the flowers, the ones that are the most frequent i. Many studies of other plant—pollinator systems provide evidence that the morphological match between the pollination apparatus and the length of the proboscis is associated with pollination effectiveness Inouye ; Waser et al.
Our experiment with fishing lines of various thicknesses, however, suggests that the width of the proboscis of the pollinators of A. We found, surprisingly, that skippers and non-skipper butterflies did not carry much pollen on their proboscides, nor did they deposit pollen on stigmas of A.
Both skippers and non-skipper butterflies have been described as nectar thieves in other systems Adrienne et al. Hoc and Garcia found Lepidoptera to be nectar robbers for Phaseolus vulgaris. Castro et al. The complex flower morphology of A. The flowers restrict access to only those visitors with mouthparts long enough to reach the nectar at the base of the floral tube. Although Pascarella et al. Skippers were the most frequent and constant visitors, often visiting numerous flowers of the same species in a row.
We have observed that skippers and non-skipper butterflies often revisit the same flowers, while long-tongued bees rarely return to a previously visited flower.
Insects revisiting the same flowers could have negative consequences, as A. In many self-incompatible Apocynaceae, flower revisitation increases the probability that self-pollen is deposited onto the stigma, leading to ovule and fruit abortion Lipow and Wyatt , ; Wyatt et al. Abortion interferes with ovules in those fruits developing from cross-pollination and wastes those potential progeny Lipow and Wyatt ; Lopes and Machado The principal pollinators of A.
Even though their proboscides are slightly shorter than the length of the narrow portion of the floral tube i. Evidently, long-tongued bees push their mouthparts firmly against the anthers in their effort to reach the nectar at the bottom of the floral tube and thereby pick up much more pollen on the wide proboscis base than the narrow, longer mouthparts of Lepidoptera that apparently miss the reproductive parts of the flowers entirely. Many studies have highlighted a close match between the length of the flowers and the length of pollinator mouthparts of members of the family Apocynaceae Endress ; Proctor et al.
In contrast, we observed no association between the lengths of the proboscis of the pollinators with pollen removal, but we did find a correspondence between the width of pollinator mouthparts and pollination efficiency. The floral morphology of A. In our study, the long-tongued bees' heads were wider than the apical portion of the pollen chamber, causing the bees to touch the reproductive parts of the flowers; short-tongued bees, with their thinner heads, entered more deeply into the corolla, but did not contact the gynostegium.
The thicker fishing line used to simulate the size of the mouthparts of the long-tongued bees removed twice as much pollen as the thinner lines, and we observed a similar pattern with our pollen deposition experiments, in which the thicker fishing line touched the stigmatic surface much more extensively than thinner ones. Using a similar approach, Darrault and Schlindwein observed that proboscis width played an important role in pollen transfer efficiency in Hancornia speciosa Apocynaceae.
By looking more closely into the mechanics of pollen removal and deposition, and by allowing single visits by the four groups of visitors, our results demonstrate that pollen removal ability, and single-visit fruit set data, can be more important for determining effective pollinators than simple visitation frequency. Funding was provided to B.
The final "swarm" mm. In a flash of thunder? Only two sections are specifically identified as being representative of butterflies mm. The music is intended to support either interpretation. The tone poem is meant to serve as a kind of Escher portrait-in some places, one can assign to the same musical material either of the "opposing" programs bees or butterflies. Thus, the same music can represent either bees or butterflies, just as the foreground of an Escher print might shift from birds to fish, depending on the viewer's prospective.
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