Previous studies of the division of labor in colonies of eusocial Hymenoptera (wasps and bees) have led to two hypotheses regarding the evolution of juvenile hormone (JH) involvement. The novel-or single-function hypothesis proposes that the role of JH has changed from an exclusively reproductive function in primitively eusocial species (those lacking morphologically distinct queen and worker castes), to an exclusively behavioral function in highly eusocial societies (those containing morphologically distinct castes). In contrast, the split-function hypothesis proposes that JH originally functioned in the regulation of both reproduction and behavior in ancestral solitary species. Then, when reproductive and brood-care tasks came to be divided between queens and workers, the effects of JH were divided as well, with JH involved in regulation of reproductive maturation of egg-laying queens, and behavioral maturation, manifested as age-correlated changes in worker tasks, of workers. We report experiments designed to test these hypotheses. After documenting age-correlated changes in worker behavior (age polyethism) in the neotropical primitively eusocial wasp Polistes canadensis, we demonstrate that experimental application of the JH analog methoprene accelerates the onset of guarding behavior, an age-correlated task, and increases the number of foraging females; and we demonstrate that JH titers correlate with both ovarian development of queens and task differentiation in workers, as predicted by the split-function hypothesis. These findings support a view of social insect evolution that sees the contrasting worker and queen phenotypes as derived via decoupling of reproductive and brood-care components of the ancestral solitary reproductive physiology.behavioral development ͉ division of labor ͉ methoprene ͉ worker polyethism

Summary 1.Interactions between bees and flowers are historically a key topic of research in biology. Apis mellifera is known to forage on a wide variety of flowers to collect nectar and/or pollen. Until now, resource collection strategies were investigated focusing on the influence of single factors on foraging decisions, such as hive requirements, surrounding vegetation, resource availability, seasonality and interactions with other bees. Despite the vast amount of literature on A. mellifera, we lack a comprehensive view of its flower selection. 2. Our aim was to go beyond specific circumstances in order to identify general criteria for flower selection at species level. Applying data mining techniques (Market Basket Analysis), we found that bees exert a precise flower selection choice mainly driven by their pollen requirements. The bees foraged all over the study area with no obvious preference for the close surrounding vegetation. They selected a restricted number of all the available sources, for nectar as well as for pollen. Apis mellifera chose some floral sources more regularly than others. The most recurrent sources changed in time according to subsequent flowering peaks and often corresponded to anemophilous species. The compelling need for pollen revealed the constant demand of a few distinct floral sources. Plants providing huge quantities of pollen, such as wind-pollinated species, were preferred. When the pollen source also provided a good quantity of nectar, foragers collected both resources on the same species. 3. This work on A. mellifera helps to clarify some of the complex aspects of its floral selection processes, which emerge from studies on single colonies, areas or environmental factors influencing individual foraging behaviours. It also suggests that some established theories on pollinator-flower interactions, such as the role of wind-pollinated species, need revision.

Questions: In animal-mediated pollination, pollinators can be regarded as a limiting resource for which entomophilous plant species might interact to assure pollination, an event pivotal for their reproduction and population maintenance. At community level, spatially aggregated co-flowering species can thus be expected to exhibit suitable suites of traits to avoid competition and ensure pollination. We explored the problem by answering the following questions: (i) are co-flowering species specialized on different guilds of pollinators? (ii) do co-flowering pollinator-sharing species segregate spatially? (iii) do co-flowering pollinator-sharing species that diverge in anther position spatially aggregate more than those that converge in anther position? Study site: Euganean Hills (NE Italy).Methods: Plant composition, flowering phenology and interactions between each entomophilous plant species and pollinating insects were monitored every fifteen days in 40 permanent plots placed in an area of 16 ha. We quantified the degree of flowering synchrony, pollinator-sharing and spatial aggregation between each pair of entomophilous species. We then tested the relationship between the degree of co-flowering, pollinator-sharing and spatial aggregation, and between spatial aggregation and anther position.Results: Entomophilous species converged, at least partially in flowering time, and the phenological synchronization of flowering was significantly associated with the sharing of pollinator guilds. Coflowering pollinator-sharing species segregated spatially. Furthermore, co-flowering pollinatorsharing species that diverged in anther position aggregated more than those that converged in anther position. Conclusions:Reproductive traits that facilitate the coexistence of co-flowering species include specialization on different pollinator guilds and a phenological displacement of the flowering time.Furthermore, in circumstances of increased competition due to phenological synchronization, pollinator sharing and spatial aggregation, the chance of an effective pollination might depend on differences in anther position, resulting in a divergent pollen placement on pollinators' body. One of the most interesting results we obtained is that the presence of one mechanism does not preclude the operation of others and each plant species can simultaneously exhibit different strategies. Although Accepted ArticleThis article is protected by copyright. All rights reserved. more studies are needed, our results can provide additional information about plant-plant interactions and add new insights into mechanisms allowing the coexistence of a high number of plant species into local communities.

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Flowers are complex structures devoted to pollinator attraction, through visual as well as chemical signals. As bees collect nectar on flowers to produce honey, some aspects of floral chemistry are transferred to honey, making chemical markers an important technique to identify the botanical and geographical origins of honey. We applied a new approach that considers the simultaneous analysis of different floral parts (petals, stamens + pistils, calyxes + nectarines, and nectar) and the corresponding unifloral honey. We collected fresh flowers of Robinia pseudoacacia L. (black locust), selected five samples of Robinia honey from different geographical origins, applied SPME-GC/MS for volatile analyses, and defined the chemical contribution added by different floral parts to the honey final bouquet. Our results show that honey blends products from nectar as well as other flower parts. Comparing honey and flower profiles, we detected compounds coming directly from flower parts but not present in the nectar, such as hotrienol and β-pinene. These may turn out to be of special interest when selecting floral markers for the botanical origin of honey.

Abstract. Acacia longifolia, a native legume from Australia, has been introduced in many European countries and elsewhere, thus becoming one of the most important global invasive species. In Europe, its flowering occurs in a period unsuitable for insect activity: nonetheless it is considered entomophilous. Floral traits of this species are puzzling: brightly coloured and scented as liked by insects, but with abundant staminate small-sized flowers and relatively small pollen grains, as it is common in anemophilous species. Invasion processes are especially favoured when reshaping local ecological networks, thus the interest in understanding pollination syndromes associated with invasive plant species that may facilitate invasiveness. Moreover, a striking difference exists between its massive flowering and relatively poor seed set. We introduced a novel approach: first, we consider the possibility that a part of the pollination success is carried on by wind and, second, we weighted the ethological perspective of the main pollinator. During the flowering season of A. longifolia (February-April 2016), we carried on exclusion experiments to detect the relative contribution of insects and wind. While the exclusion experiments corroborated the need for pollen vectors, we actually recorded a low abundance of insects. The honeybee, known pollinator of acacias, was relatively rare and not always productive in terms of successful visits. While wind contributed to seed set, focal observations confirmed that honeybees transfer pollen when visiting both the inflorescences to collect pollen and the extrafloral nectaries to collect nectar. The mixed pollination strategy of A. longifolia may then be the basis of its success in invading Portugal's windy coasts.

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