Foraging and nesting behavior of Osmia lignaria (Hymenoptera: Megachilidae) in the presence of fungicides: cage studies

During orchard pollination studies in California, we observed dramatic changes in nesting and foraging behavior of Osmia lignaria Say (Hymenoptera: Megachilidae) after sprays with tank mixtures containing fungicides. A characteristic pattern of postspray events observed includes erratic behavior and interrupted foraging and nesting activity for several days. In an effort to determine whether fungicidal sprays were disruptive to bee foraging and thus to pollination, we exposed O. lignaria females nesting in field cages planted with lacy scorpionweed, Phacelia tanacetifolia Benth (Hydrophyllaceae), to selected spray mixtures normally encountered in California orchard production systems: iprodione (Rovral), propiconazole (Orbit), benomyl (Benlate), and captan (Captan 50 WP); the surfactant Dyne-Amic, alone and mixed with Rovral; and the tank mixture IDB (Rovral + Dyne-Amic + the foliar fertilizer Bayfolan Plus). An additional cage sprayed with an equal volume of water acted as control, and a cage sprayed with the insecticide dimethoate as a toxic standard. For each female O. lignaria, we recorded time spent inside the nest depositing pollen-nectar loads, foraging time, cell production rate, and survival. All females in the dimethoate treatment died postspray + 1 d. Before death, some of these females behaved similarly to our previous orchard observations. A high proportion of females in the IDB cage were inactive for a few hours before resuming normal foraging and nesting activity. No lethal or behavioral effects were found for any of the other compounds or mixtures tested. Our results indicate that the fungicide applications that we tested are compatible with the use of O. lignaria as an orchard pollinator.     

Reproduction and survival of a solitary bee along native and exotic floral resource gradients

Native bee abundance has long been assumed to be limited by floral resources. This paradigm has been established in large measure because more bees are often found in areas supporting greater floral abundance. This could result from attraction to resource-rich sites as well as greater local demographic performance in sites supporting high floral abundance; however, demographic performance is usually unknown. Factors other than floral resources such as availability of nest sites, pressure from natural enemies, or whether floral resources are from a mixed native or mostly monodominant exotic assemblage might influence survival or fecundity and hence abundance. We examined how the survival and fecundity of the native solitary bee Osmia lignaria varied along a gradient in floral resource abundance. We released bees alongside a nest block at 27 grassland sites in Montana (USA) that varied in floral abundance and the extent of invasion by exotic forbs. We monitored nest construction and the fate of offspring within each nest. The number of nests established was positively related to native forb abundance and was negatively related to exotic forb species richness. Fecundity was positively related to native forb species richness; however, offspring mortality caused by the brood parasite Tricrania stansburyi was significantly greater in native-dominated sites. These results suggest that native floral resources can positively influence bee populations, but that the relationship between native floral resources and bee population performance is not straightforward. Rather, bees may face a trade-off between high offspring production and low offspring survival in native-dominated sites.     

Laboratory bioassays to evaluate fungicides for chalkbrood control in larvae of the alfalfa leafcutting bee (Hymenoptera: Megachilidae)

Chalkbrood, a fungal disease in bees, is caused by several species of Ascosphaera. A. aggregata is a major mortality factor in populations of the alfalfa leafcutting bee, Megachile rotundata (F.) (Hymenoptera: Megachilidae) used in commercial alfalfa seed production. Four formulated fungicides, Benlate 50 WP, Captan, Orbit, and Rovral 50 WP were tested in the laboratory for efficacy against hyphal growth of A. aggregata cultures. The same fungicides, with the addition of Rovral 4 F, were tested for their effects on incidence of chalkbrood disease, and toxicity to M. rotundata larvae. Benlate, Rovral 50 WP, and Rovral 4 F reduced incidence of chalkbrood with minimal mortality on larval bees. Benlate and Rovral 50 WP also reduced hyphal growth. Orbit was effective in reducing hyphal growth, but it did not reduce incidence of chalkbrood and was toxic to bee larvae. Captan was not effective in reducing hyphal growth or chalkbrood incidence, and it was toxic to bee larvae. Fungicides that reduce incidence of chalkbrood and larval mortality in this laboratory study are candidates for further study for chalkbrood control.     

Fungicide tests on adult alfalfa leafcutting bees (Hymenoptera: Megachilidae)

Chalkbrood is a serious disease of alfalfa leafcutting bee Megachile rotundata (F.) (Hymenoptera: Megachilidae) larvae, causing upward of 20% infection in the field. The causative agent is the fungus Ascosphaera aggregata. This bee is used extensively for alfalfa seed pollination in the United States. Using laboratory bioassays, we previously demonstrated that fungicides can reduce chalkbrood levels in the larvae. Here, we evaluate the toxicity of four fungicides, Benlate, Captan, Orbit, and Rovral, to adult bees by using three different bioassays. In the first test, fungicides were applied to bees' thoraces. In the second test, mimicking foliage residue, a piece of filter paper soaked in fungicide was placed on the bottom of a container of bees. The third test evaluated oral toxicity by incorporating fungicides into a sugar-water solution that was fed to the bees. The filter paper test did not discriminate among the fungicides well, and the oral test resulted in the greatest mortality. Toxicity to males was greater than to females. The use of fungicides for chalkbrood control is a logical choice, but caution should be used in how they are applied in the presence of bees.     

Comparative analysis of septic injury-inducible genes in phylogenetically distant model organisms of regeneration and stem cell research, the planarian Schmidtea mediterranea and the cnidarian Hydra vulgaris

Background

Host-parasite interactions are among the most important biotic relationships. Host species should evolve mechanisms to detect their enemies and employ appropriate counterstrategies. Parasites, in turn, should evolve mechanisms to evade detection and thus maximize their success. Females of the European beewolf (Philanthus triangulum, Hymenoptera, Crabronidae) hunt exclusively honeybee workers as food for their progeny. The brood cells containing the paralyzed bees are severely threatened by a highly specialized cuckoo wasp (Hedychrum rutilans, Hymenoptera, Chrysididae). Female cuckoo wasps enter beewolf nests to oviposit on paralyzed bees that are temporarily couched in the nest burrow. The cuckoo wasp larva kills the beewolf larva and feeds on it and the bees. Here, we investigated whether H. rutilans evades detection by its host. Since chemical senses are most important in the dark nest, we hypothesized that the cuckoo wasp might employ chemical camouflage.

Results

Field observations suggest that cuckoo wasps are attacked by beewolves in front of their nest, most probably after being recognized visually. In contrast, beewolves seem not to detect signs of the presence of these parasitoids neither when these had visited the nest nor when directly encountered in the dark nest burrow. In a recognition bioassay in observation cages, beewolf females responded significantly less frequently to filter paper discs treated with a cuticular extract from H. rutilans females, than to filter paper discs treated with an extract from another cuckoo wasp species (Chrysis viridula). The behavior to paper discs treated with a cuticular extract from H. rutilans females did not differ significantly from the behavior towards filter paper discs treated with the solvent only. We hypothesized that cuckoo wasps either mimic the chemistry of their beewolf host or their host's prey. We tested this hypothesis using GC-MS analyses of the cuticles of male and female beewolves, cuckoo wasps, and honeybee workers. Cuticle extracts of Hedychrum nobile (Hymenoptera: Chrysididae) and Cerceris arenaria (Hymenoptera: Crabronidae) were used as outgroups. There was little congruence with regard to cuticular compounds between H. rutilans females and honeybees as well as females of C. arenaria and H. nobile. However, there was a considerable similarity between beewolf females and H. rutilans females. Beewolf females show a striking dimorphism regarding their cuticular hydrocarbons with one morph having (Z)-9-C25:1 and the other morph having (Z)-9-C27:1 as the major component. H. rutilans females were more similar to the morph having (Z)-9-C27:1 as the main component.

Conclusion

We conclude that H. rutilans females closely mimic the composition of cuticular compounds of their host species P. triangulum. The occurrence of isomeric forms of certain compounds on the cuticles of the cuckoo wasps but their absence on beewolf females suggests that cuckoo wasps synthesize the cuticular compounds rather than sequester them from their host. Thus, the behavioral data and the chemical analysis provide evidence that a specialized cuckoo wasp exhibits chemical mimicry of the odor of its host. This probably allows the cuckoo wasp to enter the nest with a reduced risk of being detected by olfaction and without leaving traitorous chemical traces.     

Light intensity limits foraging activity in nocturnal and crepuscular bees

A crepuscular or nocturnal lifestyle has evolved in bees severaltimes independently, probably to explore rewarding pollen sourceswithout competition and to minimize predation and nest parasites.Despite these obvious advantages, only few bee species are nocturnal.Here we show that the sensitivity of the bee apposition eyeis a major factor limiting the ability to forage in dim light.We present data on eye size, foraging times, and light levelsfor Megalopta genalis (Augochlorini, Halictidae) in Panama,and Lasioglossum (Sphecodogastra) sp. (Halictini, Halictidae)in Utah, USA. M. genalis females forage exclusively during twilight,but as a result of dim light levels in the rain forest, theyare adapted to extremely low intensities. The likely factorlimiting their foraging activity is finding their nest entranceon return from a foraging trip. The lowest light intensity atwhich they can do this, both in the morning and the evening,is 0.0001 cd m^–2. Therefore, they leave the nest at dimmerlight levels in the morning than in the evening. Lasioglossum(Sphecodogastra) foraging is limited by light intensity in theevening, but probably by temperature in the morning in the temperateclimate of Utah. We propose that the evolution of nocturnalityin bees was favored by the large variance in the size of females.     

Visual marks learned by the solitary bee Megachile rotundata for localizing its nest

We studied homing behaviour of leaf-cutter bees, Megachile rotundata, by using artificial landmarks. We evaluated their nest-searching behaviour in different test situations to elucidate the nature of the visual marks they used in this task. When we modified or removed geometrical figures surrounding the nest, the bees searched for longer, showing that they noticed the introduced changes. However, these manipulations never prevented bees from finding their nest, suggesting that other visual cues were crucial in the task. Manipulations of the edges provided by the boundaries of the device (nest block, metal sheet on which the block was mounted) strongly impaired the homing performance. The further away the edges that were left intact, the stronger was the impairment of the homing behaviour. These results suggest that bees learn the distances of the various edges from the goal and that edges have a hierarchical significance according to their distance from the nest. The most distant edges provide vague information, which suffices to guide the insect towards the next edge in the sequence, until it recognizes the final, precise location of the nest. The results support the conclusion that information on distances is acquired using cues derived from motion parallax generated by the insect's self-motion. Recognition of edge parameters such as position and orientation might be achieved by an image-matching mechanism based on dynamic processes. Thus, in the homing task, there is no clear discrepancy between the eidetic and the parametric hypotheses of spatial representation.     

The Effect of Social Parasitism by Polyergus breviceps on the Nestmate Recognition System of Its Host,Formica altipetens

Highly social ants, bees and wasps employ sophisticated recognition systems to identify colony members and deny foreign individuals access to their nest. For ants, cuticular hydrocarbons serve as the labels used to ascertain nest membership. Social parasites, however, are capable of breaking the recognition code so that they can thrive unopposed within the colonies of their hosts. Here we examine the influence of the socially parasitic slave-making ant, Polyergus breviceps on the nestmate recognition system of its slaves, Formica altipetens. We compared the chemical, genetic, and behavioral characteristics of colonies of enslaved and free-living F. altipetens. We found that enslaved Formica colonies were more genetically and chemically diverse than their free-living counterparts. These differences are likely caused by the hallmark of slave-making ant ecology: seasonal raids in which pupa are stolen from several adjacent host colonies. The different social environments of enslaved and free-living Formica appear to affect their recognition behaviors: enslaved Formica workers were less aggressive towards non-nestmates than were free-living Formica. Our findings indicate that parasitism by P. breviceps dramatically alters both the chemical and genetic context in which their kidnapped hosts develop, leading to changes in how they recognize nestmates.     

Nest Entrance Marking with Colony Specific Odors by the Bumble Bee Bombus occidentalis (Hymenoptera: Apidae)

We investigated the ability of naturally foraging female bumble bees (Bombus occidentalis) to recognize their nest entrance in the field. In a blind binary-choice paradigm, females discriminated between nest entrances tipped with caps from their own nest entrances and entrances tipped with caps previously associated with foreign heterospecific or foreign conspecific colonies. Washing caps in organic solvents eliminated the manifestation of this nest-entrance recognition and significantly decreased the episodes of antennating caps. These results indicate that bees deposit colony-specific chemicals on nest entrances that are later perceived via contact chemoreception. Females treated foreign heterospecific and foreign conspecific caps similarly, suggesting that bees may not assess or respond to the degree of similarity between their own colony odor and a foreign colony odor.     

Can alloethism in workers of the bumblebee, Bombus terrestris, be explained in terms of foraging efficiency?

Bumblebee workers vary greatly in size, unlike workers of most other social bees. This variability has not been adequately explained. In many social insects, size variation is adaptive, with different-sized workers performing different tasks (alloethism). Here we established whether workers of the bumblebee, Bombus terrestris (L.) (Hymenoptera; Apidae), exhibit alloethism. We quantified the size of workers engaging in foraging compared to those that remain in the nest, and confirmed that it is the larger bees that tend to forage (X±SE thorax widths 4.34±0.01 mm for nest bees and 4.93±0.02 mm for foragers). We then investigated whether large bees are better suited to foraging because they are able to transport heavier loads of food back to the nest. Both pollen and nectar loads of returning foragers were measured, demonstrating that larger bees do return with a heavier mass of forage. Foraging trip times were inversely related to bee size when collecting nectar, but were unrelated to bee size for bees collecting pollen. Overall, large bees brought back more nectar per unit time than small bees, but the rate of pollen collection appeared to be unrelated to size. The smallest foragers had a nectar foraging rate close to zero, presumably explaining why foragers tend to be large. Why might larger bees be better at foraging? Various explanations are considered: larger bees are able to forage in cooler conditions, may be able to forage over larger distances, and are perhaps also less vulnerable to predation. Conversely, small workers are presumably cheaper to produce and may be more nimble at within-nest tasks. Further research is needed to assess these possibilities. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Social Interactions and Familiarity in a Communal Halictine Bee Lasioglossum (Chilalictus) hemichalceum

Social organization in Hymenoptera is quite variable, but most group-living species have a reproductive division of labor. Communal species, by definition, do not; all individuals work and all are reproductive. In order to understand the evolution and maintenance of such cooperative societies the essential first step is characterization of the behavioral interactions between individuals. In particular, it is important to investigate the influence of common group membership (familiarity) on these interactions. Circle-tube arenas were used to observe the interactions between 42 pairs of field-collected Lasioglossum (Chilalictus) hemichalceum females either from the same nest, from different nests but the same nest aggregation, or from different aggregations. Identical procedures were applied to 24 pairs of females from laboratory colonies. There was a striking lack of agonistic behavior with Lunging occurring in only 0.03% of all bouts, and C-Postures (threat behavior) in only 19% of bouts using field bees and 27 % of bouts using females held in the laboratory. Passing was virtually ubiquitous, occurring in 98 % of all bouts. None of these behavior patterns is significantly associated with the level of familiarity between females. The lack of agonistic behavior and the indiscriminate cooperative behavior (Passing) suggest that all conspecifics are accepted and are treated similarly. This is in contrast to previous studies of L. (Dialictus) zephyrum that demonstrated both kin recognition and nepotism. While Avoidance behavior was rare, significantly more was seen between bees taken from different nest aggregations. Trophallaxis Solicitation occurred in 13 bouts between field bees. While the proportion of bouts in which this occurred is not associated with familiarity, the number of solicitations per bout is greater between females from different nest aggregations. These minor but statistically significant differences suggest that L. hemichalceum females can discriminate between familiar and unfamiliar individuals. Therefore the proximate mechanism underlying universal acceptance of all conspecifics cannot be a lack of recognition cue expression or recognition cue perception. Rather, it is due to a lack of clearly preferential treatment toward familiar individuals. This suggests that some component of fitness is positively associated with group membership per se, or group size. The high levels of mutual tolerance suggest, as do other lines of evidence (Kukuk & Schwarz 1987), that communal societies are not intermediate between solitary and eusocial species.     

Stingless bees (<Emphasis Type="Italic">Scaptotrigona pectoralis</Emphasis>) learn foreign trail pheromones and use them to find food

Foragers of several species of stingless bees (Hymenoptera, Apidae and Meliponini) deposit pheromone marks in the vegetation to guide nestmates to new food sources. These pheromones are produced in the labial glands and are nest and species specific. Thus, an important question is how recruited foragers recognize their nestmates’ pheromone in the field. We tested whether naïve workers learn a specific trail pheromone composition while being recruited by nestmates inside the hive in the species Scaptotrigona pectoralis. We installed artificial scent trails branching off from trails deposited by recruiting foragers and registered whether newly recruited bees follow these trails. The artificial trails were baited with trail pheromones of workers collected from foreign S. pectoralis colonies. When the same foreign trail pheromone was presented inside the experimental hives while recruitment took place a significant higher number of bees followed the artificial trails than in experiments without intranidal presentation. Our results demonstrate that recruits of S. pectoralis can learn the composition of specific trail pheromone bouquets inside the nest and subsequently follow this pheromone in the field. We, therefore, suggest that trail pheromone recognition in S. pectoralis is based on a flexible learning process rather than being a genetically fixed behaviour.     

Colony disturbance and solitary nest initiation by workers in the obligately eusocial sweat bee, Halictus ligatus

Sweat bees are one of the most socially polymorphic lineages on the planet. In obligately eusocial species, newly enclosed females may become either queens or workers, depending on the environmental and social circumstances of the nest into which they emerge. In socially polymorphic species, females also have the option of nesting solitarily, founding a nest and raising future reproductives alone, without the help of other adult females. Halictus ligatus is a widespread Nearctic, ground-nesting sweat bee. It has been particularly well studied in Ontario, where detailed studies have described it as obligately eusocial. Here we report evidence that the flexibility of female H. ligatus actually extends to expressing behaviour more typical of socially polymorphic species, those in which some individuals reproduce solitarily. In a population in southern Ontario, black wasps (Astata sp.) emerged from the soil beneath the nesting aggregation and proceeded to excavate their own nesting tunnels, dislocating many H. ligatus nest entrances. Young workers whose natal nests were destroyed by the wasp activity constructed new nests, so under very specific circumstances, it is possible for potential altruists to nest solitarily.     

Visual and Olfactory Floral Cues of Campanula (Campanulaceae) and Their Significance for Host Recognition by an Oligolectic Bee Pollinator

Oligolectic bees collect pollen from a few plants within a genus or family to rear their offspring, and are known to rely on visual and olfactory floral cues to recognize host plants. However, studies investigating whether oligolectic bees recognize distinct host plants by using shared floral cues are scarce. In the present study, we investigated in a comparative approach the visual and olfactory floral cues of six Campanula species, of which only Campanula lactiflora has never been reported as a pollen source of the oligolectic bee Ch. rapunculi. We hypothesized that the flowers of Campanula species visited by Ch. rapunculi share visual (i.e. color) and/or olfactory cues (scents) that give them a host-specific signature. To test this hypothesis, floral color and scent were studied by spectrophotometric and chemical analyses, respectively. Additionally, we performed bioassays within a flight cage to test the innate color preference of Ch. rapunculi. Our results show that Campanula flowers reflect the light predominantly in the UV-blue/blue bee-color space and that Ch. rapunculi displays a strong innate preference for these two colors. Furthermore, we recorded spiroacetals in the floral scent of all Campanula species, but Ca. lactiflora. Spiroacetals, rarely found as floral scent constituents but quite common among Campanula species, were recently shown to play a key function for host-flower recognition by Ch. rapunculi. We conclude that Campanula species share some visual and olfactory floral cues, and that neurological adaptations (i.e. vision and olfaction) of Ch. rapunculi innately drive their foraging flights toward host flowers. The significance of our findings for the evolution of pollen diet breadth in bees is discussed.     

Role of chemical cues and natal rearing effect on host recognition by the parasitic wasp Melittobia digitata

Parasitoids are expected to have the ability to find, recognize, and perhaps to discern potential hosts that can best support the development of their progeny. Melittobia Westwood (Hymenoptera: Eulophidae) are gregarious ectoparasitoids, which primarily attack mud daubers (Hymenoptera: Sphecidae). How Melittobia females locate their host is not well known, but the process may involve host‐related chemical signals. In this study, we investigated the roles of host chemical cues and natal rearing effect in host recognition by Mel. digitata Dahms. In an olfactometer that contained prepupae of Trypoxylon politum Say (Hymenoptera: Sphecidae), Megachile rotundata (Fabricius) (Hymenoptera: Megachilidae), puparia of Neobellieria bullata (Parker) (Diptera: Sarcophagidae), empty or intact host cocoons, or nest mud, Mel. digitata females spent significantly more time in air fields that contained T. politum (prepupae + cocoon) and Meg. rotundata (prepupae + cocoon) than in N. bullata and control fields. Nest mud and natal host had no attraction for parasitoid host choice. Most first and last choices of Mel. digitata females in the olfactometer were not consistent, suggesting an initial random dispersion, although they responded positively towards hosts in cocoons.     

Queen-worker conflict and social parasitism in bumble bees (Hymenoptera: Apidae)

Psithyrus ashtoni (Hymenoptera: Apidae) is an obligate, workerless bumble, bee social parasite which invades nests of Bombus affinis. Although parasites are limited by host worker defence to invading very small colonies, there is considerable flexibility in the way parasites control host brood bionomics once they are accepted inside the nest. A study of 46 parasitized and 22 non-parasitized laboratory colonies of B. affinis showed that P. ashtoni females cohabited with host queens and workers while the worker force increased, but not to the maximum normally achieved in non-parasitized nests. While colony reproductive success was correlated with the number of workers reared, parasites risked being killed or ejected from the comb by workers, after the queen had lost dominance. Host bees usually succeeded in rearing offspring, and Psithyrus reproductive success was related to the ability of parasites to control proportional investment in the two species. In addition to displacing P. ashtoni females, host bees ate the eggs of parasites and ejected their larvae. These behaviours were also exhibited by workers in the later stages of development of non-parasitized colonies. These results indicate that social parasites are at least partially subject to the conflict of genetic self-interest between the queen and her workers which is believed to influence the control of reproductive investment in haplodiploid Bombus societies.     

Floating and fidelity in nest visitation byCrawfordapis luctuosa (Hymenoptera: Colletidae)

We studied nest visitation behavior of the ground-nesting beeCrawfordapis luctuosa (Colletidae) at the Monteverde Cloud Forest Preserve, Puntarenas Province, Costa Rica. Some females entered many nests, while others consistently entered only one nest. Multiple nest entry was exhibited by bees we called floaters, and those bees that consistently entered only one nest were provisioning bees. Provisioning bees abandoned their old nests after provisioning periods and became nonprovisioning bees that floated over the nesting area. Floating behavior was a nest-searching behavior typified by oscillating flights just above the ground (<15 cm). Floaters investigated available nests and were not observed to interact aggressively with nest owners. Upon establishing a new nest, floaters became provisioning bees.     

Review of bees as guests in termite nests,with a new record of the communal bee, <Emphasis Type="Italic">Gaesochira obscura</Emphasis> (Smith, 1879) (Hymenoptera,Apidae), in nests of <Emphasis Type="Italic">Anoplotermes banksi</Emphasis> Emerson, 1925 (Isoptera,Termitidae, Apicotermitinae)

Associations between bees and termites are documented infrequently, but records are available for bee species ranging in behavior from solitary to highly eusocial. The subtribe Meliponina (stingless bees) is the most common bee group reported in termite nests, and, for some species, the occupancy of termite nests may be obligatory. The records of solitary bees nesting within termite nests include species of the tribes Emphorini, Centridini, Megachilini, and Paracolletini. Most of these bees can probably nest in other substrates, and their relationships with termite nests are presumably opportunistic. This study provides a review of published records of bees as guests in termite nests, and also describes the aggregation of nests of Gaesochira obscura within one nest of Anoplotermes banksi in Brazilian Amazonia. One termite nest contained at least ten nest entrances of G. obscura, with burrows 4–6 mm in diameter and about 10 cm long. Each burrow ended in brood cells in different stages of food provisioning and larval development. As commonly reported for other associations of this nature, there was no connection between the tunnels of bees and those of termites. This record adds important data on the biology of A. banksi. Because this is a single record, it is impossible to classify G. obscura either as a termitophile or termitariophile; this species may be opportunistic in relation to nesting substrate.     

How Do Carpenter Bees Recognize the Entrance of their Nests?: An Experimental Investigation in a Natural Habitat1

Carpenter bee females (genus Xylocopa) enter only their own nests. They can be found in the same burrow over a period of months, regardless of whether they arc living singly or aggregated in close vicinity. The aim of this study was to investigate and to clarify which sensory systems are responsible for the recognition of the nest. Females of three different African species were tested in their natural habitat. Following alteration in the immediate visual surroundings of either the entrance or the entire dwelling stem, bees behaved as they did in undisturbed situations and entered their nests. Furthermore, a relative dislocation of the entrance within the dwelling stem or a displacement of the whole stem within a restricted range had no effect on the recognition of the burrow. When the entrance was plugged by foam rubber the bees landed only after extended searching. When the entrance was closed by a strip of scotch tape the bees searched for several minutes, departed, then returned and searched again with the same result. The bees showed no indications that they recognized their burrow. These results strongly support the conclusion that the bees recognized their burrows primarily or exclusively by olfactory cues, and, furthermore, that individual odors were involved. The possible contributions of different sensory systems for the recognition of the nest and the possible biological function of the use of olfactory cues are discussed.     

Adult pollen diet essential for egg maturation by a solitary Osmia bee

Reproduction is a nutritionally costly activity for many insects, as their eggs are rich in lipids and proteins. That cost seems especially acute for non-social bees, which for their size, lay enormous eggs. All adult female bees visit flowers, most of them to collect pollen and nectar, or sometimes oils, to feed their progeny. For adult bees, the need for pollen feeding has only been detailed for the honey bee, Apis mellifera. To experimentally test for the reproductive value of adult pollen feeding by a non-social bee, Osmia californica (Hymenoptera: Apiformes: Megachilidae), young female bees plus males were released into large glasshouse cages provided with either a male-fertile sunflower cultivar or a pollen-less one. Females regularly visited and drank nectar from flowers of both cultivars. Abundant orange pollen was seen regularly in guts of females confined with the male-fertile sunflowers, indicative of active pollen ingestion. All females’ terminal oocytes (next egg to be laid) were small at emergence. Oocytes of females confined with the pollen-less sunflowers remained small, despite frequent nectaring and exposure to other floral stimuli. In contrast, the basal oocytes of female O. californica with access to pollen had swelled to full size within ten days following emergence, enabling them to lay eggs in provided nest tubes. Adult females of this solitary bee required dietary pollen to reproduce; nitrogen stores acquired as larvae were inadequate. Early and regular pollen feeding in part paces the onset and maximum tempo of solitary bees’ lifetime reproductive output.