The Extraordinary Alkali Bee, Nomia melanderi (Halictidae), the World's Only Intensively Managed Ground-Nesting Bee.

Among the ground-nesting bees are several proven crop pollinators, but only the alkali bee (Nomia melanderi) has been successfully managed. In <80 years, it has become the world's most intensely studied ground-nesting solitary bee. In many ways, the bee seems paradoxical. It nests during the torrid, parched midsummer amid arid valleys and basins of the western United States, yet it wants damp nesting soil. In these basins, extensive monocultures of an irrigated Eurasian crop plant, alfalfa (lucerne), subsidize millions of alkali bees. Elsewhere, its polylectic habits and long foraging range allow it to stray into neighboring crops contaminated with insecticides. Primary wild floral hosts are either non-native or poorly known. Kleptoparasitic bees plague most ground nesters, but not alkali bees, which do, however, host other well-studied parasitoids. Building effective nesting beds requires understanding the hydraulic conductivity of silty nesting soils and its important interplay with specific soil mineral salts. Surprisingly, some isolated populations endure inhospitably cold climates by nesting amid hot springs. Despite the peculiarities and challenges associated with its management, the alkali bee remains the second most valuable managed solitary bee for US agriculture and perhaps the world.

Wildfire severity influences offspring sex ratio in a native solitary bee.

Although ecological disturbances can have a strong influence on pollinators through changes in habitat, virtually no studies have quantified how characteristics of wildfire influence the demography of essential pollinators. Nevertheless, evaluating this topic is critical for understanding how wildfire is linked to pollinator population dynamics, particularly given recent changes in wildfire frequency and severity in many regions of the world. In this study, we measured the demographic response of the blue orchard bee (Osmia lignaria) across a natural gradient of wildfire severity to assess how variation in wildfire characteristics influenced reproductive output, offspring sex ratio, and offspring mass. We placed nest blocks with a standardized number and sex ratio of pre-emergent adult bees across the wildfire gradient, finding some evidence for a positive but highly variable relationship between reproductive output and fire severity surrounding the nest site at both local (100 m) and landscape (750 m) scales. In addition, the production of female offspring was > 10% greater at nest sites experiencing the greatest landscape-scale fire severity relative to the lowest-severity areas. The finding that blue orchard bees biased offspring production towards the more expensive offspring sex with increasing fire severity shows a functional response to changes in habitat quality through increased density of flowering plants. Our findings indicate that burned mixed-conifer forest provides forage for the blue orchard bee across a severity gradient, and that the increase in floral resources that follows high-severity fire leads females to shift resource allocation to the more costly sex when nesting.

Comparative Pollination Efficacies of Five Bee Species on Raspberry.

Unlike many other rosaceous fruit crops, commercial raspberry cultivars are largely self-fertile and mostly self-pollinate autogamously. However, their floral morphology does not allow for complete autopollination, which often yields unmarketable small or crumbly fruits. Insect visitation is therefore essential to maximize raspberry production. Honey bees are typically used to pollinate commercial raspberries, but escalating prices for hive rentals coupled with increasing acreage encourages evaluation of other manageable pollinators. Four other manageable bee taxa-various Bombus spp., Osmia lignaria Say, Osmia aglaia Sandhouse, and Osmia bruneri Cockerell-are all promising raspberry pollinators. Because honey bees remain the least expensive option on a per forager basis, adoption of an alternative pollinator should entail some other advantage, such as superior pollination efficacy. In this study, we compared honey bees with these other bee species for their pollination efficacies at red raspberries, measured as the number of drupelets resulting from a single visit to a virgin flower. Each species' single-visit pollination efficacy was also compared with drupelet set from both unvisited and hand-pollinated flowers, and their pollination effectiveness scores were calculated. All five bee species were equally effective raspberry pollinators; therefore, honey bees remain the most cost effective option for open field pollination of raspberry. Mason bees and bumble bees may have greater utility during cool weather or for protected cultivation systems, contexts unfavorable to honey bee foraging.

Survey of Hatching Spines of Bee Larvae Including Those of Apis mellifera (Hymenoptera: Apoidea).

This article explores the occurrence of hatching spines among bee taxa and how these structures enable a larva on hatching to extricate itself from the egg chorion. These spines, arranged in a linear sequence along the sides of the first instar just dorsal to the spiracles, have been observed and recorded in certain groups of solitary and cleptoparasitic bee taxa. After eclosion, the first instar remains loosely covered by the egg chorion. The fact that this form of eclosion has been detected in five families (Table 1 identifies four of the families. The fifth family is the Andrenidae for which the presence of hatching spines in the Oxaeinae will soon be announced.) of bees invites speculation as to whether it is a fundamental characteristic of bees, or at least of solitary and some cleptoparasitic bees. The wide occurrence of these spines has prompted the authors to explore and discover their presence in the highly eusocial Apis mellifera L. Hatching spines were indeed discovered on first instar A. mellifera. The honey bee hatching process appears to differ in that the spines are displayed somewhat differently though still along the sides of the body, and the chorion, instead of splitting along the sides of the elongate egg, seems to quickly disintegrate from the emerging first instar in association with the nearly simultaneous removal of the serosa that covers and separates the first instar from the chorion. Unexpected observations of spherical bodies of various sizes perhaps containing dissolving enzymes being discharged from spiracular openings during hatching may shed future light on the process of how A. mellifera effects chorion removal during eclosion. Whereas hatching spines occur among many groups of bees, they appear to be entirely absent in the Nomadinae and parasitic Apinae, an indication of a different eclosion process.

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.

Crop domestication facilitated rapid geographical expansion of a specialist pollinator, the squash bee Peponapis pruinosa.

Squash was first domesticated in Mexico and is now found throughout North America (NA) along with Peponapis pruinosa, a pollen specialist bee species of the squash genus Cucurbita The origin and spread of squash cultivation is well-studied archaeologically and phylogenetically; however, no study has documented how cultivation of this or any other crop has influenced species in mutualistic interactions. We used molecular markers to reconstruct the demographic range expansion and colonization routes of P. pruinosa from its native range into temperate NA. Populations east of the Rocky Mountains expanded from the wild host plant's range in Mexico and were established by a series of founder events. Eastern North America was most likely colonized from squash bee populations in the present-day continental Midwest USA and not from routes that followed the Gulf and Atlantic coasts from Mexico. Populations of P. pruinosa west of the Rockies spread north from the warm deserts much more recently, showing two genetically differentiated populations with no admixture: one in California and the other one in eastern Great Basin. These bees have repeatedly endured severe bottlenecks as they colonized NA, following human spread of their Cucurbita pollen hosts during the Holocene.

Progeny of Osmia lignaria from distinct regions differ in developmental phenology and survival under a common thermal regime.

Many insects, including some bees, have extensive subcontinental distributions that can differ in climatic conditions. Within and beyond these distributions, humans intentionally transport beneficial insects, including bees, to non-natal geographic locations. Insects also are experiencing unprecedented climatic change in their resident localities. For solitary bees, we know very little about the adaptive plasticity and geographic variation in developmental physiology that accommodates the different climates experienced within distributional ranges. Osmia lignaria Say (Hymenoptera: Megachilidae) is a widely distributed North American spring-emerging bee being developed as a managed pollinator for tree fruit crops, including almonds. We examined the development and survival of O. lignaria progeny that were descended from populations sourced from southern California, western Washington, and northern Utah, and then were reared together under an hourly and weekly temperature regime simulating those of a California almond-growing region. We found that developmental physiologies of Washington and Utah progeny were generally similar. However, California progeny developed slower, were more metabolically active, and survived better under California conditions than did populations native to regions at higher latitudes. Regardless of geographic origin, cocooned adults managed under prescribed thermal regimes emerged faster and lived longer after wintering. Progeny of parents from different regions exhibited some acclimatory plasticity in developmental phenologies to a novel climatic regime, but overall their responses reflected their geographic origins. This outcome is consistent with their developmental phenologies being largely heritable adaptations to regional climates.

Specialist Osmia bees forage indiscriminately among hybridizing Balsamorhiza floral hosts.

Pollinators, even floral generalists (=polyleges), typically specialize during individual foraging bouts, infrequently switching between floral hosts. Such transient floral constancy restricts pollen flow, and thereby gene flow, to conspecific flowers in mixed plant communities. Where incipient flowering species meet, however, weak cross-fertility and often similar floral traits can yield mixed reproductive outcomes among pollinator-dependent species. In these cases, floral constancy by polyleges sometimes serves as an ethological mating barrier. More often, their foraging infidelities instead facilitate host introgression and hybridization. Many other bee species are oligolectic (taxonomic specialists for pollen). Oligoleges could be more discriminating connoisseurs than polyleges when foraging among their limited set of related floral hosts. If true, greater foraging constancy might ensue, contributing to positive assortative mating and disruptive selection, thereby facilitating speciation among their interfertile floral hosts. To test this Connoisseur Hypothesis, nesting females of two species of oligolectic Osmia bees were presented with randomized mixed arrays of flowers of two sympatric species of their pollen host, Balsamorhiza, a genus known for hybridization. In a closely spaced grid, the females of both species preferred the larger flowered B. macrophylla, evidence for discrimination. However, both species' females showed no floral constancy whatsoever during their individual foraging bouts, switching randomly between species proportional to their floral preference. In a wider spaced array in which the bouquets reflected natural plant spacing, foraging oligolectic bees often transferred pollen surrogates (fluorescent powders) both between conspecific flowers (geitonogamy and xenogamy) and between the two Balsamorhiza species. The Connoisseur Hypothesis was therefore rejected. Foraging infidelity by these oligolectic Osmia bees will contribute to introgression and hybridization where interfertile species of Balsamorhiza meet and flower together. A literature review reveals that other plant genera whose species hybridize also attract numerous oligolectic bees, providing independent opportunities to test the generality of this conclusion.

The alfalfa leafcutting bee, Megachile rotundata: the world's most intensively managed solitary bee.

The alfalfa leafcutting bee (ALCB), Megachile rotundata F. (Megachildae), was accidentally introduced into the United States by the 1940s. Nest management of this Eurasian nonsocial pollinator transformed the alfalfa seed industry in North America, tripling seed production. The most common ALCB management practice is the loose cell system, in which cocooned bees are removed from nesting cavities for cleaning and storage. Traits of ALCBs that favored their commercialization include gregarious nesting; use of leaves for lining nests; ready acceptance of affordable, mass-produced nesting materials; alfalfa pollination efficacy; and emergence synchrony with alfalfa bloom. The ALCB became a commercial success because much of its natural history was understood, targeted research was pursued, and producer ingenuity was encouraged. The ALCB presents a model system for commercializing other solitary bees and for advancing new testable hypotheses in diverse biological disciplines.

Pollination value of male bees: the specialist bee Peponapis pruinosa (Apidae) at summer squash (Cucurbita pepo).

Male bees can be abundant at flowers, particularly floral hosts of those bee species whose females are taxonomic pollen specialists (oligolecty). Contributions of male bees to host pollination are rarely studied directly despite their prevalence in a number of pollination guilds, including those of some crop plants. In this study, males of the oligolectic bee, Peponapis pruinosa Say, were shown to be effective pollinators of summer squash, Cucurbita pepo L. Seven sequential visits from male P. pruinosa maximized squash fruit set and growth. This number of male visits accumulated during the first hour of their foraging and mate searching at flowers soon after sunrise. Pollination efficacy of male P. pruinosa and their abundances at squash flowers were sufficient to account for most summer squash production at our study sites, and by extrapolation, to two-thirds of all 87 North American farms and market gardens growing squashes that were surveyed for pollinators by collaborators in the Squash Pollinators of the Americas Survey. We posit that the substantial pollination value of male Peponapis bees is a consequence of their species' oligolecty, their mate seeking strategy, and some extreme traits of Cucurbita flowers (massive rewards, flower size, phenology).

Importance of pollinators in changing landscapes for world crops.

The extent of our reliance on animal pollination for world crop production for human food has not previously been evaluated and the previous estimates for countries or continents have seldom used primary data. In this review, we expand the previous estimates using novel primary data from 200 countries and found that fruit, vegetable or seed production from 87 of the leading global food crops is dependent upon animal pollination, while 28 crops do not rely upon animal pollination. However, global production volumes give a contrasting perspective, since 60% of global production comes from crops that do not depend on animal pollination, 35% from crops that depend on pollinators, and 5% are unevaluated. Using all crops traded on the world market and setting aside crops that are solely passively self-pollinated, wind-pollinated or parthenocarpic, we then evaluated the level of dependence on animal-mediated pollination for crops that are directly consumed by humans. We found that pollinators are essential for 13 crops, production is highly pollinator dependent for 30, moderately for 27, slightly for 21, unimportant for 7, and is of unknown significance for the remaining 9. We further evaluated whether local and landscape-wide management for natural pollination services could help to sustain crop diversity and production. Case studies for nine crops on four continents revealed that agricultural intensification jeopardizes wild bee communities and their stabilizing effect on pollination services at the landscape scale.

Complex responses within a desert bee guild (Hymenoptera: Apiformes) to urban habitat fragmentation.

Urbanization within the Tucson Basin of Arizona during the past 50+ years has fragmented the original desert scrub into patches of different sizes and ages. These remnant patches and the surrounding desert are dominated by Larrea tridentata (creosote bush), a long-lived shrub whose flowers are visited by > 120 native bee species across its range. Twenty-one of these bee species restrict their pollen foraging to L. tridentata. To evaluate the response of this bee fauna to fragmentation, we compared species incidence and abundance patterns for the bee guild visiting L. tridentata at 59 habitat fragments of known size (0.002-5 ha) and age (up to 70 years), and in adjacent desert. The 62 bee species caught during this study responded to fragmentation heterogeneously and not in direct relation to their abundance or incidence in undisturbed desert. Few species found outside the city were entirely absent from urban fragments. Species of ground-nesting L. tridentata specialists were underrepresented in smaller fragments and less abundant in the smaller and older fragments. In contrast, cavity-nesting bees (including one L. tridentata specialist) were overrepresented in the habitat fragments, probably due to enhanced nesting opportunities available in the urban matrix. Small-bodied bee species were no more likely than larger bodied species to be absent from the smaller fragments. The introduced European honey bee, Apis mellifera, was a minor faunal element at > 90% of the fragments and exerted little if any influence on the response of native bee species to fragmentation. Overall, bee response to urban habitat fragmentation was best predicted by ecological traits associated with nesting and dietary breadth. Had species been treated as individual units in the analyses, or pooled together into one analysis, these response patterns may not have been apparent. Pollination interactions with this floral host are probably not adversely affected in this system because of its longevity and ability to attract diverse pollinators but will demand careful further study to understand.

Dose-response relationships between pollination and fruiting refine pollinator comparisons for cranberry (Vaccinium macrocarpon [Ericaceae]).

Comparisons of pollinator efficacy using pollen received on stigmas can be refined by incorporating experimental dose-response relationships for pollen deposition and fruiting responses. A range of discrete pollen doses applied to cranberry stigmas resulted in decelerating curvilinear responses for fruiting, berry size, and seed set. Minimum thresholds and maximum asymptotes bounded reproductive responses to incremental stigmatic pollen loads. Four bee species were compared for their pollination efficacies on commercial cranberries, using counts of pollen received by stigmas during single bee visits to previously virgin flowers. Differences between these bee species were found to be exaggerated when raw pollen counts were used for comparison because foragers of some species often delivered pollen in excess of that needed to maximize fruit and seed production. Sixfold differences between species in mean pollen deposition translated into 1.5-2-fold differences in predicted cranberry fruit set and size. Implications for pollen tube competition and agricultural production are discussed.

Pollinating bees (Hymenoptera: Apiformes) of U.S. alfalfa compared for rates of pod and seed set.

Alfalfa (=lucerne) flowers require visiting bees to trip the sexual column, thereby providing pollination and subsequent pod and seed set. Previous studies have compared the pollination values of different bee species solely by the speed with which they handle flowers and the proportion of visited flowers tripped. In this greenhouse study, five species of bees, including the three commercially managed U.S alfalfa pollinators, are likewise compared for their floral tripping frequencies. These bee species are also compared for the pod set and mature seed that results from their single visits to virgin flowers. Regardless of the identity of the pollinating bee, tripped flowers had the same probabilities of pod set and seed set. Thus, differences in the single-visit pollination efficiencies of the various bee species are entirely attributable to the proportion of visited flowers that they trip. Females of the alkali bee, Nomia melanderi Cockerell, and the alfalfa leafcutting bee, Megachile rotundata F, tripped 81 and 78% of visited flowers, respectively. Males of these species are significantly less effective (61 and 51%, respectively), but still significantly superior to the honey bee, Apis mellifera L. (22% of visited flowers tripped). These relationships are supported by field data for tripping frequencies. One candidate pollinator, Osmia sanrafaelae Parker, shows promise (44% tripped), but not the congeneric O. aglaia Sandhouse (13% tripped).

Bees assess pollen returns while sonicating Solanum flowers.

Can bees accurately gauge accumulating bodily pollen as they harvest pollen from flowers? Several recent reports conclude that bees fail to assess pollen harvest rates when foraging for nectar and pollen. A native nightshade (Solanum elaeagnifolium Cavanilles) that is visited exclusively for pollen by both solitary and social bees (eg. Ptiloglossa and Bombus) was studied in SE Arizona and SW New Mexico. The flowers have no nectaries. Two experiments were deployed that eliminated "pollen feedback" to the bees by experimentally manipulating flowers prior to bee visits. The two methods were 1) plugging poricidal anthers with glue and 2) emptying anthers of pollen by vibration prior to bee visitation. Both experiments demonstrated that bees directly assess pollen harvest on a flower-by-flower basis, and significantly tailor their handling times, number of vibratile buzzes per flower and grooming bouts according to the ongoing harvest on a given flower. In comparison to experimental flowers, floral handling times were extended for both Bombus and Ptiloglossa on virgin flowers. Greater numbers of intrafloral buzzes and numbers of times bees groomed pollen and packed it into their scopae while still on the flower were also more frequent at virgin versus experimental flowers. Flowers with glued andreocia received uniformly brief visits from Bombus and Ptiloglossa with fewer sonications and virtually no bouts of grooming. Curtailed handling with few buzzes and grooms also characterized visits to our manually harvested flowers wherein pollen was artificially depleted. Sonicating bees respond positively to pollen-feedback while harvesting from individual flowers, and therefore we expect them to adjust their harvesting tempo according to the currency of available pollen (standing crop) within Solanum floral patches.