Multiyear study of pollinator efficiency and importance of a wide array of pollinators in a field-cultivated strawberry plot.

Using wild pollinators to pollinate crops without introducing human-managed pollinators is cost-effective and friendly to native ecosystems. To maintain stable, good-quality yields in crops that mainly use wild pollinators, it is essential to determine which flower visitors are important pollinators and their degree of importance. In this study, we observed flower-visiting insects for 5 years in outdoor cultivated strawberries surrounded by a semi-natural environment in central Japan. We estimated the pollination effectiveness and efficiency of the 10 main flower-visiting insect taxa on strawberries by examining the relationship between the number of visits per flower and subsequent achene fertilization rates per berry. Finally, the pollinator importance (%) to the total pollination service was estimated for each of the 10 main taxa and for all others. Among the 10 main insect taxa, 6 were effective pollinators, i.e., they significantly increased achene fertilization rates by increasing their number of visits to a flower. Considering the 5-year mean, these six taxa accounted for the top six important pollinators. Andrena (subgenus Micrandrena) spp. were the most important and three other bee taxa, including Apis mellifera and Ceratina spp., were the next most important pollinators; one fly and one butterfly species were also important pollinators. This indicates that strawberry pollinators were diverse in the study area. The flower-visit frequency and importance of many pollinators fluctuated from year to year, implying that various pollinators pollinate strawberry flowers each year, and in field surveys of crop-pollinator communities multiyear investigations are needed to identify important pollinators and to estimate their importance. To the best of our knowledge, this is the first attempt to quantify the proportional importance of each pollinator to the total pollination service for a crop.

Long horns protect Hestina japonica butterfly larvae from their natural enemies.

Animals sometimes have prominent projections on or near their heads serving diverse functions such as male combat, mate attraction, digging, capturing prey, sensing or defence against predators. Some butterfly larvae possess a pair of long frontal projections; however, the function of those projections is not well known. Hestina japonica butterfly larvae have a pair of long hard projections on their heads (i.e., horns). Here we hypothesized that they use these horns to protect themselves from natural enemies (i.e., predators and parasitoids). Field surveys revealed that the primary natural enemies of H. japonica larvae were Polistes wasps. Cage experiments revealed that larvae with horns intact and larvae with horns removed and fitted with horns of other individuals succeeded in defending themselves against attacks of Polistes wasps significantly more often than larvae with horns removed. We discuss that the horns counter the paper wasps' hunting strategy of first biting the larvae's 'necks' and note that horns evolved repeatedly only within the Nymphalidae in a phylogeny of the Lepidoptera. This is the first demonstration that arthropods use head projections for physical defence against predators.

The choosing of sleeping position in the overnight aggregation by the solitary bees Amegilla florea urens in Iriomote Island of Japan.

In addition to the process of joining the sleeping aggregation, the choice of sleeping position is an important night-time behaviour of small diurnal insects because of the increased risk for predator attacks as well as bad weather. The aggregation behaviour of the solitary bee Amegilla florea urens was investigated to elucidate the choice of sleeping position on substrates. Male and female constructed single-sex aggregations on hanging leaves during May and June, respectively. Most individuals tended to form aggregations with other individuals while few individuals slept alone. During the aggregation forming, both the number of individuals that tried to join the aggregation and the completion time of aggregation increased with the number of sleeping individuals, whereas the success rate of joining was unaffected. The sleeping positions of subsequent arrivals on the substrates were higher than those of the first arrivals in female aggregations. Therefore, the first female to arrive tended to be located near the bottom of a hanging substrate. Dissecting sleeping females showed that they contained mature oocytes, indicating that sexually mature individuals formed aggregations. In male aggregations, however, we could not find a clear relationship between the position on substrates and the arrival sequence. We suggest that the purpose for sleeping in aggregations might be a dilution effect for nocturnal predation and that the females that finished both nesting and foraging quickly could choose the optimal positions in the aggregation when they arrived on the sleeping substrates.

Long Frontal Projections Help Battus philenor (Lepidoptera: Papilionidae) Larvae Find Host Plants.

Animals sometimes develop conspicuous projections on or near their heads as, e.g., weaponry, burrowing or digging tools, and probes to search for resources. The frontal projections that insects generally use to locate and assess resources are segmented appendages, including antennae, maxillary palps, and labial palps. There is no evidence to date that arthropods, including insects, use projections other than true segmental appendages to locate food. In this regard, it is noteworthy that some butterfly larvae possess a pair of long antenna-like projections on or near their heads. To date, the function of these projections has not been established. Larvae of pipevine swallowtail butterflies Battus philenor (Papilionidae) have a pair of long frontal fleshy projections that, like insect antennae generally, can be actively moved. In this study, we evaluated the possible function of this pair of long moveable frontal projections. In laboratory assays, both frontal projections and lateral ocelli were shown to increase the frequency with which search larvae found plants. The frontal projections increased finding of host and non-host plants equally, suggesting that frontal projections do not detect host-specific chemical cues. Detailed SEM study showed that putative mechanosensillae are distributed all around the frontal as well as other projections. Taken together, our findings suggest that the frontal projections and associated mechanosensillae act as vertical object detectors to obtain tactile information that, together with visual information from lateral ocelli and presumably chemical information from antennae and mouthparts, help larvae to find host plants. Field observations indicate that host plants are small and scattered in southern Arizona locations. Larvae must therefore find multiple host plants to complete development and face significant challenges in doing so. The frontal projections may thus be an adaptation for finding a scarce resource before starving to death. This is the first evidence that arthropods use projections other than true segmental appendages such as antennae, mouthparts and legs, to locate food resources.

Reward and non-reward learning of flower colours in the butterfly Byasa alcinous (Lepidoptera: Papilionidae).

Learning plays an important role in food acquisition for a wide range of insects. To increase their foraging efficiency, flower-visiting insects may learn to associate floral cues with the presence (so-called reward learning) or the absence (so-called non-reward learning) of a reward. Reward learning whilst foraging for flowers has been demonstrated in many insect taxa, whilst non-reward learning in flower-visiting insects has been demonstrated only in honeybees, bumblebees and hawkmoths. This study examined both reward and non-reward learning abilities in the butterfly Byasa alcinous whilst foraging among artificial flowers of different colours. This butterfly showed both types of learning, although butterflies of both sexes learned faster via reward learning. In addition, females learned via reward learning faster than males. To the best of our knowledge, these are the first empirical data on the learning speed of both reward and non-reward learning in insects. We discuss the adaptive significance of a lower learning speed for non-reward learning when foraging on flowers.

Interspecific and intersexual learning rate differences in four butterfly species.

Learning plays an important role in food acquisition for a wide range of insects and has been demonstrated to be essential during flower foraging in taxa such as bees, parasitoid wasps, butterflies and moths. However, little attention has been focused on differences in floral cue learning abilities among species and sexes. We examined the associative learning of flower colour with nectar in four butterfly species: Idea leuconoe, Argyreus hyperbius, Pieris rapae and Lycaena phlaeas. All butterflies that were trained learned the flower colours associated with food. The flower colour learning rates were significantly higher in I. leuconoe and A. hyperbius than in P. rapae and L. phlaeas. Among the four species examined, the larger and longer-lived species exhibited higher learning rates. Furthermore, female butterflies showed a significantly higher learning rate than males. This study provides the first evidence that learning abilities related to floral cues differ among butterfly species. The adaptive significance of superior learning abilities in the larger and longer-lived butterfly species and in females is discussed.

An invasive dandelion unilaterally reduces the reproduction of a native congener through competition for pollination.

The impact of invasive alien species on native species is of increasing global concern. Invasive plants have various negative effects on natives through competition; however, relatively little is known about competition for pollination. The relationship between Japanese native dandelions (Taraxacum spp.) and invasive congeners may be a typical case of such an interaction. For example, native dandelions are being replaced by invasive congeners, especially in urban and suburban areas of Japan. To explain this phenomenon, we hypothesized that when natives are mixed with attractive invasives, natives may suffer from reduced seed set because invasives deprive natives of pollinators or because pollinators frequently move between species, resulting in interspecific pollen transfer. To test this hypothesis, we studied the effect of the invasive dandelion T. officinale on the pollination and seed set of the native T. japonicum using artificial arrays of monospecific and mixed-species plots as well as natural populations. Taraxacum officinale attracted more pollinator visits, perhaps because it produced more nectar than T. japonicum. The number of pollinator visits to T. japonicum was reduced when the congeners were grown together, and pollinators moved frequently between the two species. The proportion of seed set for T. japonicum was reduced in the presence of T. officinale in both artificial arrays and natural populations. These results support our hypothesis that interspecific competition for pollination plays an important role in the recent replacement of native dandelions by invasive congeners in Japan. Because invasive dandelions are apomicts, negative effects are incurred only by sexual natives. Thus, this system can be recognized as a rare case of interspecific interaction through pollination.