Why do leafcutter bees cut leaves? New insights into the early evolution of bees

Stark contrasts in clade species diversity are reported across the tree of life and are especially conspicuous when observed in closely related lineages. The explanation for such disparity has often been attributed to the evolution of key innovations that facilitate colonization of new ecological niches. The factors underlying diversification in bees remain poorly explored. Bees are thought to have originated from apoid wasps during the Mid-Cretaceous, a period that coincides with the appearance of angiosperm eudicot pollen grains in the fossil record. The reliance of bees on angiosperm pollen and their fundamental role as angiosperm pollinators have contributed to the idea that both groups may have undergone simultaneous radiations. We demonstrate that one key innovation--the inclusion of foreign material in nest construction--underlies both a massive range expansion and a significant increase in the rate of diversification within the second largest bee family, Megachilidae. Basal clades within the family are restricted to deserts and exhibit plesiomorphic features rarely observed among modern bees, but prevalent among apoid wasps. Our results suggest that early bees inherited a suite of behavioural traits that acted as powerful evolutionary constraints. While the transition to pollen as a larval food source opened an enormous ecological niche for the early bees, the exploitation of this niche and the subsequent diversification of bees only became possible after bees had evolved adaptations to overcome these constraints.     

Orchard pollination in Capitol Reef National Park,Utah, USA. Honey bees or native bees?

Capitol Reef National Park in central Utah, USA surrounds 22 managed fruit orchards started over a century ago by Mormon pioneers. Honey bees are imported for pollination, although the area in which the Park is embedded has over 700 species of native bees, many of which are potential orchard pollinators. We studied the visitation of native bees to apple, pear, apricot, and sweet cherry over 2 years. Thirty species of bees visited the flowers but, except for pear flowers, most were uncommon compared to honey bees. Evidence that honey bees prevented native bees from foraging on orchard crop flowers was equivocal: generally, honey bee and native bee visitation rates to the flowers were not negatively correlated, nor were native bee visitation rates positively correlated with distance of orchards from honey bee hives. Conversely, competition was tentatively suggested by much larger numbers of honey bees than natives on the flowers of apples, apricots and cherry; and by the large increase of native bees on pears, where honey bee numbers were low. At least one-third of the native bee species visiting the flowers are potential pollinators, including cavity-nesting species such as Osmia lignaria propinqua, currently managed for small orchard pollination in the US, plus several fossorial species, including one rosaceous flower specialist (Andrena milwaukiensis). We suggest that gradual withdrawal of honey bees from the Park would help conserve native bee populations without decreasing orchard crop productivity, and would serve as a demonstration of the commercial value of native pollinators.     

Why do bees turn back and look?

The timing of learning of colour and shape of the food source, as well as of near-by landmarks, was examined exploiting a behaviour described recently, the Turn Back and Look behaviour (TBL): Bees departing from a novel food source after feeding turn around to view it at a short distance (Figs. 2, 3) before departing for the hive. They repeat this behaviour on several successive visits, termed the TBL phase (Fig. 5). To examine the function of the TBL, I trained individual bees in 4 different modes. In the first 3 they could view a food source or a landmark of a particular colour or shape during (i) arrival as well as departure, (ii) only arrival, and (iii) only departure; in the final mode (iv) the bees viewed one colour (or shape) on arrival, and another on departure. At the end of the TBL phase, the bees were tested by offering them a choice between the visual stimulus to which they were trained (modes i–iii) and a different (novel) one, or between the stimulus viewed on arrival and that viewed on departure (mode iv). The test results show that learning after feeding (while performing the TBL), i.e. backward conditioning, occurs regardless of whether the colour (Fig. 6, Fig. 10a) or shape (Fig. 7) of the food source, or the colour (Fig. 10b), shape (Fig. 11), and position (Fig. 12) of a near-by landmark is considered. Bees trained in mode (iv) preferred the stimulus learned on arrival over that learned on departure in almost all cases. However, a stimulus viewed exclusively on departure (mode iii) was often learned as well as when it was viewed exclusively on arrival (mode ii) (Figs. 10a, 11, 12), or both on arrival and departure (mode i) (Fig. 6). The finding that the timing of learning can be manipulated suggests that it is not based on hard wired predispositions to learn particular visual cues on arrival, and others on departure.     

How do bees learn and recognize visual patterns?

The basic task of perceptual systems is the recognition and localization of objects. The central nervous system has to solve these problems on the basis of the excitation patterns of sensory nerves, in spite of the fact that these provide only ambiguous information about objects. Two processing principles seem to be fundamental for an efficient formation of object representations: the extraction of characteristic features and the ability to assess similarities between different objects. This article reviews investigations in which different training paradigms were applied in order to explore the honeybee's capacities to learn and recognize visual patterns. One aim of these experiments is to assess whether insects use similar processing mechanisms as vertebrates, for instance human beings. By comparing the computational performance of perceptual systems in animals with different evolutionary history we can hope to learn more about the operation of basic rules in nervous systems. “The messages which the brain receives have not the least similarity with the stimuli. They consist in pulses of given intensities and frequencies, characteristic for the transmitting nerve-fiber, which ends at a definite place of the cortex. ... From this information it produces the image of the world by a process which can metaphorically be called a consummate piece of combinatorial mathematics: it sorts out of the maze of indifferent and varying signals, invariant shapes and relations which form the world of ordinary experience.” Max Born (1949) Received: 4 October 1997 / Accepted in revised form: 26 August 1998     

Bacillus DNA in fossil bees: an ancient symbiosis?

We report here the isolation of DNA from abdominal tissue of four extinct stingless bees (Proplebeia dominicana) in Dominican amber, PCR amplification of a 546-bp fragment of the 16S rRNA gene from Bacillus spp., and their corresponding nucleotide sequences. These sequences were used in basic local alignment search tool searches of nonredundant nucleic acid data bases, and the highest scores were obtained with 16S rRNA sequences from Bacillus spp. Phylogenetic inference analysis by the maximum-likelihood method revealed close phylogenetic relationships of the four presumed ancient Bacillus sequences with Bacillus pumilus, B. firmus, B. subtilis, and B. circulans. These four extant Bacillus spp. are commonly isolated from abdominal tissue of stingless bees. The close phylogenetic association of the extracted DNA sequences with these bee colonizers suggests that a similar bee-Bacillus association existed in the extinct species P. dominicana.     

Environment or kin: whence do bees obtain acidophilic bacteria?

As honey bee populations decline, interest in pathogenic and mutualistic relationships between bees and microorganisms has increased. Honey bees and bumble bees appear to have a simple intestinal bacterial fauna that includes acidophilic bacteria. Here, we explore the hypothesis that sweat bees can acquire acidophilic bacteria from the environment. To quantify bacterial communities associated with two species of North American and one species of Neotropical sweat bees, we conducted 16S rDNA amplicon 454 pyrosequencing of bacteria associated with the bees, their brood cells and their nests. Lactobacillus spp. were the most abundant bacteria in many, but not all, of the samples. To determine whether bee-associated lactobacilli can also be found in the environment, we reconstructed the phylogenetic relationships of the genus Lactobacillus. Previously described groups that associate with Bombus and Apis appeared relatively specific to these genera. Close relatives of several bacteria that have been isolated from flowers, however, were isolated from bees. Additionally, all three sweat bee species associated with lactobacilli related to flower-associated lactobacilli. These data suggest that there may be at least two different means by which bees acquire putative probiotics. Some lactobacilli appear specific to corbiculate apids, possibly because they are largely maternally inherited (vertically transmitted). Other lactobacilli, however, may be regularly acquired from environmental sources such as flowers. Sweat bee-associated lactobacilli were found to be abundant in the pollen and frass inside the nests of halictids, suggesting that they could play a role in suppressing the growth of moulds and other spoilage organisms.     

Why do nectar-foraging bees and wasps work upwards on inflorescences?

Summary Wasps (Dolichovespula and Vespula spp.) worked predominantly upwards when foraging for nectar on inflorescences of the protogynous Scrophularia aquatica, in which the standing crop of nectar sugar per flower showed no clear pattern of vertical distribution up an inflorescence. Bumblebees taking nectar (Bombus hortorum visiting legally, and certain individuals of B. terrestris which positioned themselves head-upwards while taking nectar through holes bitten in the corolla) worked predominantly upwards on the racemose inflorescences of Linaria vulgaris, although the standing crop of nectar sugar per open flower increased up the raceme. Individuals of B. terrestris which robbed Linaria flowers in a head-down position worked predominantly downwards on inflorescences. The upward or downward directionality of intra-inflorescence movements by foraging insects may depend in part on the position these adopt during their flower visits.     

Are body size and volatile blends honest signals in orchid bees?

Secondary sexual traits may convey reliable information about males’ ability to resist pathogens and that females may prefer those traits because their genes for resistance would be passed on to their offspring. In many insect species, large males have high mating success and can canalize more resources to the immune function than smaller males. In other species, males use pheromones to identify and attract conspecific mates, and thus, they might function as an honest indicator of a male's condition. The males of orchid bees do not produce pheromones. They collect and store flower volatiles, which are mixed with the volatile blends from other sources, like fungi, sap and resins. These blends are displayed as perfumes during the courtship. In this study, we explored the relationship between inter‐individual variation in body size and blend composition with the males’ phenoloxidase (PO) content in Euglossa imperialis. PO content is a common measure of insect immune response because melanine, its derived molecule, encapsulates parasites and pathogens. Body size and blend composition were related to bees’ phenolic PO content. The inter‐individual variation in body size and tibial contents could indicate differences among males in their skills to gain access to some compounds. The females may evaluate their potential mates through these compounds because some of them are reliable indicators of the males’ capacity to resist infections and parasites.     

Spider signals: are web decorations visible to birds and bees?

We are becoming increasingly aware of animal communication outside the range of human sensitivity. Web decorations are silk structures used by orb-web spiders to deceive prey and predators. However, despite the level of interest in these structures, their visibility to prey and predators has never, to our knowledge, been objectively assessed. Here, we use spectrophotometric analyses to show that the decorations of all five tested spider species are visible to honey bees and birds over short and long distances. Furthermore, the discoid decorations of one species may provide some protection against arthropod predators. However, these decorations are inefficient at camouflaging the spider against birds, despite the overlap between the spider's body and web decoration.     

Spillover of trap-nesting bees and wasps in an urban–rural interface

A mismatch of resource availability in certain periods can lead to spillover of insects between habitats, resulting in temporal differences in insect diversity. Urban gardens are important anthropogenic habitats but it is unknown whether, when and why spillover of beneficial insects occurs between gardens and agricultural habitats. We used trap nests for Hymenoptera to monthly monitor bee and wasp abundance and species richness in 12 gardens and 12 rapeseed fields. Half of the gardens and rapeseed fields were located in the urban–rural interface and bordered each other (a garden paired with a rapeseed field) and the other half were isolated in the rural landscape (isolated rapeseed fields) and in the urban city centre (isolated gardens). In general, gardens in the urban–rural interface comprised the highest richness of bees and wasps. The abundance of bees but not of wasps was highest in paired habitats and peaked at full rapeseed blooming, indicating that mass-flowering rapeseed offers foraging resources for bees nesting in adjacent gardens. Thus, bees nest and increase their populations in both areas, benefiting from the mass-flowering resource in the agricultural habitat as well as the nesting resources from gardens, suggesting spillover of bees but not of wasps between paired gardens and rapeseed fields. Our study highlights the value of gardens in the urban–rural interface for the biodiversity of functionally important insects. Implementing urban gardening and small-scale agriculture in cities and suburban habitats can promote local pollinator populations and benefit adjacent croplands.     

Why are there so many species of bumble bees at Dungeness?

WILLIAMS, P. H., 1989. Why are there so many species of bumble bees at Dungeness? Dungeness is unique in the British Isles in that it has more species of bumble bees than any other locality. Three ideas about what governs the number of species at a locality are examined by locking at patterns of flower visits at Dungeness in comparison with those at Shoreham, a species-poor locality also in Kent. The species of bumble bees that are present at Dungeness but absent from Shoreham show no association in their distributions among 2 km grid-squares in Kent with the species of food-plants that they prefer at Dungeness, nor is there any correlation between the diversity of bees and diversity of food-plants at Dungeness and Shoreham. From the information available, Dungeness is most likely to have more species of bumble bees because it has a particularly high density of the more nectar-rich flowers that bumble bees can use. Bumble bees feed most profitably from deep flowers because these contain more nectar than shallow flowers, although direct access to deeper flowers is ultimately limited by the length of each bee's proboscis. The distribution of worker proboscis lengths among species in the species-pool in Kent is clumped about a median of 7.9 mm. The best foraging conditions for the maximum number of species should be provided when flowers of similar depths are present in sufficiently large numbers for all foragers to make near-optimal flower choices. Although there is no difference in median between the distributions of the bees' proboscis lengths and the depths of the flowers they use at Dungeness, at Shoreham the flower depths used are shorter than the proboscis lengths. Among the food-plants at Dungeness, high densities of Teucrium scorodonia and Echium vulgare are likely to be especially important.     

Floral constancy in bumble bees: handling efficiency or perceptual conditioning?

Individual bees often prefer flowers of the same species that they are already foraging on, and other individual bees prefer other flowers. This floral constancy has classically been explained as a learned behavior by which bees avoid wasting time switching between handling techniques. Choice trails were given to Bombus vagans workers that were freely foraging in mixed and pure fields of Trifolium pratense, T. repens, Viccia cracca, and Prunella vulgaris. Contrary to expectation, (1) bees showed if anything a stronger preference for their flower type in pure fields where they lacked experience than in a mixed field where they had had the opportunity to learn, (2) there was greater constancy in a mixed field of the two morphologically similar Trifolium species than in a mixed field of the morphologically disparate T. pratense and P. vulgaris, and (3) bees were more willing to switch between flowers of distinct morphologies when the colors were similar than between flowers of distinct colors when the morphologies were similar. We suggest that constancy is due to some form of perceptual conditioning whereby individual bees become temporarily sensitized to one or a few floral cues.     

Exocrine secretions of bees—VIII: 8-acetoxy-2,6-dimethyl-2-octenal: A novel mandibular gland component from Panurginus bees (Hymenoptera: Andrenidae)

A terpenoid aldehyde, 8-acetoxy-2,6-dimethyl-2-octenal, has been identified in the mandibular gland secretions of male and female Panurginus atramontensis. This is the first report of such a compound from bees. In addition, geranial and neral were shown to be present in female extracts.     

Longer tongues and swifter handling: why do more bumble bees (Bombus spp.) than honey bees (Apis mellifera) forage on lavender (Lavandula spp.)?

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Does time spent on adult bees affect reproductive success of Varroa mites?

Reproduction ofVarroa jacobsoni Oudemans (Acari: Varroidae) and the number ofVarroa mites that were found dead on the bottom board of the hive, were studied in relation to the period the mites spent on adult honey bees,Apis mellifera L. (Hymenoptera: Apidae), prior to invasion into brood cells. The maximum period on adult bees was 23 days. To introduce mites, combs with emerging worker brood, heavily infested with mites, were placed into a colony and removed the next day. At the beginning of the first day following emergence from brood cells, 18% of the mites introduced into the colony was found on the bottom of the hive. Part of these mites may already have died inside the capped brood cells, and then fallen down after cleaning of cells by the bees. At the second and third day following emergence, respectively 4% and 2% of the mites on adult bees at the previous day was recovered on the bottom, whereas from the fourth day on only 0.6% of the mites on adult bees was recovered on the bottom per day. After invasion into brood cells, 8–12% of the mites did not produce any offspring. Of the mites that did reproduce, the total number of offspring was 4.0–4.4 per mite during one reproductive cycle, part of which may reach maturity resulting in 1.2–1.3 viable daughters, and 8–10% of the mites produced only male offspring. Reproduction was independent of the period the mites had spent on adult bees prior to invasion into brood cells.     

Solitary bees – Potential vectors for gene flow from cultivated to wild strawberries

The genus Fragaria (Rosaceae) contains 24 plant species, including hybrid species such as the octoploid garden strawberry (F. × ananassa). Natural hybridization between Fragaria species has repeatedly been reported, and the potential future cultivation of genetically modified strawberries has made the study of hybridization potential between F. × ananassa and its wild relatives increasingly important. In Europe, F. × ananassa is the only octoploid species present, and the most likely candidate for hybridization is the common diploid woodland strawberry (F. vesca). To date, it is unknown whether pollinator spectra of the two Fragaria species overlap and thus might promote interspecific gene flow. We carried out a survey of flower visitors in northwestern Switzerland to identify major flower visitors of F. vesca and F. × ananassa. This survey indicated that wild bees are the most important shared flower visitors of F. × ananassa and F. vesca. Therefore, we studied flower choice behavior of the common wild bee Osmia bicornis in a greenhouse experiment. Osmia bicornis did not discriminate between F. × ananassa and F. vesca flowers. We conclude that wild bees are important shared flower visitors of both F. × ananassa and F. vesca and are potential vectors for gene flow between cultivated and wild strawberries.