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The alfalfa leafcutting bee (Megachile rotundata (Fabricius)), a commercial pollinator used for alfalfa seed production, is susceptible to chalkbrood disease via ingested fungal spores. Diseases of insects can elicit behavioral changes in their hosts, but there are no recorded behaviors of alfalfa leafcutting bees in response to this fungal exposure. We conducted field studies to determine whether bees in pathogen-dense environments altered their nesting patterns, specifically if bees exposed to fungal spores produced higher numbers of nest cells and whether the proportions of nest cells that failed as eggs or small larvae (a state known as ‘pollen ball’) were greater. We found that our control bees, nontreated bees which were not exposed to chalkbrood spores other than those in the natural environment, had the highest proportion of pollen ball cells. Bees experimentally exposed to infective spores created the lowest number of nests and the fewest cells. Bees experimentally exposed to heat killed noninfective spores produced the greatest number of nests and cells overall and the greatest number of healthy progeny. We conclude that there are underlying behaviors that are elicited in response to the presence of chalkbrood spores that reduce the proportion of failed nest cells (grooming) and increase retention of bees at nesting sites (delay of bee emergence). Through further study of these behaviors, bee managers can potentially increase the productivity of their bee populations.

Interrupting the spring incubation of Megachile rotundata (F.) with a period of low-temperature storage for synchronizing the bees’ emergence with crop bloom is an essential part of M. rotundata management. Previously, we demonstrated that bees exposed to thermoperiods (TPs) during low-temperature storage have higher survival rates than bees exposed to constant temperatures. But changing the temperature in the large mass of bees commonly found in most commercial settings would place considerable stress on the chambers’ refrigeration system. Reducing the difference between a TP’s cryophase and thermophase would decrease the stress on the refrigeration system. Therefore, we investigated a range of TPs with cryophases (12 h) of 6, 12, or 15°C and thermophases (12 h) of 15 or 18°C and compared the survival rates of these bees against bees exposed to constant temperatures of 12, 15, or 18°C. For eye-pigmented pupae, the TP 6–18°C and the control fluctuating thermal regime (FTR; 6°C with a daily 1-h pulse at 20°C) had the highest survival rates for the 2 yr tested. For the constant-temperature storage protocols, constant 15 and 18°C were either equivalent or lower survival than the control FTR. For emergence-ready adults, the 6–18°C TP had the highest survival rates. The constant 15°C and the control FTR had equivalent survival rates. Under the current constraints imposed by a commercial chamber’s refrigeration system, interrupting M. rotundata spring incubation by exposing the developing bees to constant temperatures of 15–18°C is currently the best option for commercial operations.

Bees are economically critical pollinators, but are declining broadly due to several stressors, including nontarget exposure to insecticides and deficiencies in nutrition. Understanding the simultaneous impact of stressors, particularly interactions between them, is critical to effectively conserving bees. Although behavioral effects of pesticides like neonicotinoids have received some attention in solitary bees, our understanding of how they are modulated by diet quality is limited. Furthermore, scarce data exist on what concentrations of orally ingested neonicotinoids elicit mortality in solitary bees. In a controlled exposure laboratory experiment, we investigated how diet quality, as sugar concentration, and chronic oral exposure to imidacloprid affect adult alfalfa leafcutting bees, Megachile rotundata (Fabricius). We provided individuals ad libitum with either 20 or 50% (m/m) sucrose syrups containing either 0, 30, or 300 ppb imidacloprid (measuring 0, 27, and 209 ppb via an ELISA assay). Over 5 wk, we tracked behavior and survivorship of individuals. Imidacloprid decreased survivorship in a dose-dependent fashion, but sucrose content did not affect survivorship, even in bees not fed imidacloprid. In the high imidacloprid treatment, 45% of bees were observed in a motionless supine position while still alive, with this effect appearing to be buffered against by the higher sucrose diet. Our results suggest that diets higher in sugar concentration may prevent an intermediate stage of poisoning, but do not ultimately extend longevity. In devising risk assessments for bees, it is important to consider that interactions between stressors may occur in the stages leading up to death even if survivorship is unaffected.

Hybrid canola seed production is an important pollination market in Canada; typically both honey bees (Apis mellifera L. (Hymenoptera: Apidae)) and Alfalfa Leafcutting bees (Megachile rotundata Fab. (Hymenoptera: Megachilidae)) are concurrently managed to ensure pollination in this high-value crop. Beekeepers are paid to provide pollination services, and the colonies also produce a honey crop from the canola. Pollen availability from male-fertile plants is carefully managed in this crop to provide an abundance of pollen to fertilize male-sterile (‘female’) plants. This abundance of pollen represents an underutilized resource for beekeepers, and an opportunity to diversify the hive-products produced for market in this management system. We used a commercial-style pollen trap to collect pollen from colonies twice weekly for the duration of canola pollination, and compared the honey production and amount of sealed brood in colonies with pollen traps to those without pollen traps. We found that while pollen trapping reduced honey production, there was no negative impact on brood production, and at current market prices, the per-hive revenue was higher in colonies from which pollen was trapped. Pollen trapping honey bee colonies in the context of hybrid canola pollination, therefore, offers beekeepers an opportunity to diversify their products and increase their revenue.

The alfalfa leafcutting bee, Megachile rotundata (F.), is the primary pollinator for alfalfa seed production in North America. Under current management practice, developing pupae are incubated at 29-30°C until the adults emerge for pollination. If unfavorable spring weather delays peak alfalfa bloom, managers will cool pupae to slow development, which can increase mortality and causes sublethal effects. Previously, we demonstrated that exposure to a fluctuating thermal regime (FTR) increases survival and extends the viable storage period. To determine the optimal conditions for FTR during storage of developing M. rotundata , we examined four variables: temperature of the daily warm pulse, duration of the warm pulse, number of weeks exposed to the FTR treatment, and developmental stage of the bee. Survival was measured by successful eclosion to the adult stage. Under all conditions, exposure to FTR increased survival compared with exposure to a constant 6°C. When the temperature of the daily warm pulse was 20-25°C from a base temperature of 6°C, and the pulse duration was extended to 3 h, survival rates were as high as those observed under standard storage conditions (29°C). Under this FTR storage protocol, bee managers can delay emergence for ∼8 wk without significant decreases in survival. Our findings have substantial economic implications for bee management and alfalfa seed production by increasing the flexibility and efficiency of M. rotundata adult emergence.

The alfalfa leafcutting bee, Megachile rotundata F. (Hymenoptera: Megachilidae), is one of the most intensively managed solitary bees and greatly contributes to alfalfa production in both the United States and Canada. Although production of certain commodities, especially alfalfa seed, has become increasingly dependent on this species' pollination proficiency, little information is known about how M. rotundata is affected by insecticide exposure. To better understand the risk posed to M. rotundata by the increasing use of insecticides to manage mosquitoes, we conducted field experiments that directly exposed M. rotundata nests, adults, and larvae to a pyrethroid insecticide via a ground-based ultra-low-volume (ULV) aerosol generator. We directly targeted nest shelters with Zenivex® E20 (etofenprox) at a half-maximum rate of 0.0032 kg/ha at dusk and then observed larval mortality, adult mortality, and the total number of completed nests for both the treated and control groups. There was no significant difference in the proportion of dead (P = 0.99) and alive (P = 0.23) larvae when the control group was compared with the treated group. We also did not observe a significant difference in the number of emerged adults reared from the treated shelters (P = 0.22 and 0.50 for females and males, respectively), and the number of completed cells after exposure to the insecticides continued to increase throughout the summer, indicating that provisioning adults were not affected by the insecticide treatment. The results from this study suggest that the amount of insecticide reaching nest shelters may not be sufficient to cause significant mortality.

Current regulatory requirements for insecticide toxicity to nontarget insects focus on the honey bee, Apis mellifera (L.; Hymenoptera: Apidae), but this species cannot represent all insect pollinator species in terms of response to insecticides. Therefore, we characterized the toxicity of pyrethroid insecticides used for adult mosquito management (permethrin, deltamethrin, and etofenprox) on a nontarget insect, the adult alfalfa leafcutting bee, Megachile rotundata (F.; Hymenoptera: Megachilidae) in two separate studies. In the first study, the doses causing 50 and 90% mortality (LD50 and LD90, respectively) were used as endpoints and 2-d-old adult females were exposed to eight concentrations ranging from 0.0075 to 0.076 μg/bee for permethrin and etofenprox, and 0.0013-0.0075 μg/bee for deltamethrin. For the second study, respiration rates of female M. rotundata were also recorded for 2 h after bees were dosed at the LD50 values to give an indication of stress response. Results indicated a relatively similar LD50 for permethrin and etofenprox, 0.057 and 0.051 μg/bee, respectively, and a more toxic response, 0.0016 μg/bee for deltamethrin. Comparatively, female A. mellifera workers have a LD50 value of 0.024 μg/bee for permethrin and 0.015 μg/bee for etofenprox indicating that female M. rotundata are less susceptible to topical doses of these insecticides, except for deltamethrin, where both A. mellifera and M. rotundata have an identical LD50 of 0.0016 μg/bee. Respiration rates comparing each active ingredient to control groups, as well as rates between each active ingredient, were statistically different (P < 0.0001). The addition of these results to existing information on A. mellifera may provide more insights on how other economically beneficial and nontarget bees respond to pyrethroids.

The alfalfa leafcutting bee Megachile rotundata (F.) is the primary pollinator for alfalfa seed production. Under standard management conditions, the alfalfa leafcutting bee develops to the diapausing prepupal stage under field conditions, after which they are cold-stored at a static temperature until the following spring, when temperatures are raised and development resumes. We have assessed the effects of a fluctuating thermal regime (FTR) during overwintering cold storage, where bees were exposed to a daily 1 h pulse of 20°C, and compared viability and insect quality to bees stored under a static thermal regime. Our results demonstrate that implementing an FTR protocol dramatically increases the survival of cold-stored alfalfa leafcutting bees, effectively extending their shelf-life into the subsequent growing season. These findings could substantially ameliorate significant obstacles that restrict the more widespread use of this important pollinator, such as the biological constraints that restrict its use in early blooming crops, and yearly fluctuations in bee prices that add significant financial uncertainty to end users. This study also strengthens a growing body of evidence that indicates FTR protocols are superior to static thermal regime protocols for insect cold storage.

The alfalfa leafcutting bee, Megachile rotundata F., is used extensively to pollinate alfalfa for seed production in western North America. However, it usually is not possible to sustain bee populations in the United States. In the managed pollination system, variable microenvironments are experienced by developing M. rotundata during the nesting season that may influence bee reproductive success. In this study, bee cells were produced in a small alfalfa field and collected from domiciles according to nesting boards and different portions of boards. Examination of cells showed that the production of live overwintering prepupae, diseased larvae, other dead eggs and larvae, and cells attacked by natural enemies varied according to board orientations and positions, and portions of the boards. Northeast-facing board samples were significantly heavier and had more pollen balls and chalkbrood compared with southwest-facing boards. Lower boards in stacks had significantly more chalkbrood than upper boards. Outer portions of boards had significantly more live prepupae and less chalkbrood than middle and inner portions, and outer and middle portions had less summer-emerging bees. These results suggest that reproductive success could be altered by changing the orientation of boards within domiciles or changing the designs of domiciles. To accurately assess the quality of bee populations managed in the commercial loose cell system, a sample of bees should be taken from a pooled sample from all boards in each domicile. Each domicile sample should then be evaluated according to its proportional contribution to the whole field's progeny production (e.g., by weight or volume).

Japanese hornfaced bees Osmia cornifrons (Radoszkowski) (Hymenoptera: Megachilidae) are used for pollination of spring blooming fruit crops such as apple, pear, and blueberry. Because O. cornifrons has a short adult life span, synchronization of bee emergence with bloom is critical to maximize crop pollination. This study was conducted to determine lower temperature thresholds (LTDs), optimum temperatures, and required degree-day accumulation for emergence of O. cornifrons adults. Patterns of temperature-dependent emergence of O. cornifrons adults at seven temperatures (3.9, 12.0, 18.6, 26.6, 30.3, 35.6, and 42.5°C) were modeled and simulated with linear and nonlinear regression analyses. Results of this study showed that required degree-days (DD) for emergence of male and female O. cornifrons adults were 125.2 DD, with LTD of 8.9°C and 179.8 DD, with LTD of 8.6°C, respectively. The optimum temperatures for emergence were 36.5, 30.2, and 35.7°C for male, female, and both sexes combined, respectively. This study indicated that emergence of O. cornifrons adults could be manipulated to synchronize with pollination periods of target fruit crops.

With the recent decline of honey bees, Apis mellifera (L.) (Hymenoptera: Apidae), there is a need for alternative or supplemental crop pollinators, such as Osmia cornifrons (Radoszkowski) (Hymenoptera: Megachilidae). However, O. cornifrons propagation could be impeded by a cleptoparasitic mite, Chaetodactylus krombeini Baker. We investigated the effects of formic acid and wintergreen oil on mortality of C. krombeini hypopi and O. cornifrons adults by determining the lethal concentration of each compound on each species. On average, >4.8 and >1.8 h were required to cause mortality in O. cornifrons adults when 176.7 ppm of formic acid and wintergreen oil, respectively. Estimates of LC50 for C. krombeini hypopi treated with formic acid and wintergreen oil were 54.3 and 271.3 ppm, respectively. This study showed that C. krombeini could be controlled effectively without inducing O. cornifrons adult mortality based on concentration and duration of fumigation.

The alfalfa leafcutting bee, Megachile rotundata F. (Hymenoptera: Megachilidae), is widely used as a pollinator for production of alfalfa, Medicago sativa L., seed, and populations of these bees can be maintained by alfalfa seed growers or can be purchased from mostly Canadian bee providers. M. rotundata raised in Canada have higher survival rates during the incubation that occurs after winter storage than do bees produced in the northwestern United States, but no reason has been found for this difference. We investigated whether storing immature M. rotundata for various time periods at a warm temperature (16°C) before winter or allowing them to remain unmanaged at ambient temperatures affects physiological aspects of prepupae during the winter as well as the survival and longevity of adult bees after spring or summer incubation. Our results show that the timing of the onset of winter storage and incubation does affect prepupal weights, prepupal lipid and water contents, adult emergence, and adult female longevity. Winter storage of prepupae in November or December with a late June incubation resulted in heavier adults that emerged more readily than bees incubated in late May. However, adult females incubated in May thrived longer than June-incubated bees if fed a honey-water diet. Thus, some prewinter management regimes for M. rotundata commercial stocks may be more effective than others for achieving optimal adult emergence synchrony, as well as adult survival and longevity for pollination of a summer crop.

Pest management practices may be contributing to a decline in wild bee populations in or near canola (Brassica napus L.) agroecosystems. The objective of this study was to investigate the direct contact toxicity of five technical grade insecticides—imidacloprid, clothianidin, deltamethrin, spinosad, and novaluron—currently used, or with potential for use in canola integrated pest management on bees that may forage in canola: common eastern bumble bees [Bombus impatiens (Cresson); hereafter bumble bees], alfalfa leafcutting bees [Megachile rotundata (F.)], and Osmia lignaria Cresson. Clothianidin and to a lesser extent imidacloprid were highly toxic to all three species, deltamethrin and spinosad were intermediate in toxicity, and novaluron was nontoxic. Bumble bees were generally more tolerant to the direct contact applications > O. lignaria > leafcutting bees. However, differences in relative toxicities between the three species were not consistent, e.g., whereas clothianidin was only 4.9 and 1....

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.

We examined lethal and sublethal effects of imidacloprid on Osmia lignaria (Cresson) and clothianidin on Megachile rotundata (F.) (Hymenoptera: Megachilidae). We also made progress toward developing reliable methodology for testing pesticides on wild bees for use in pesticide registration by using field and laboratory experiments. Bee larvae were exposed to control, low (3 or 6 ppb), intermediate (30 ppb), or high (300 ppb) doses of either imidacloprid or clothianidin in pollen. Field experiments on both bee species involved injecting the pollen provisions with the corresponding pesticide. Only O. lignaria was used for the laboratory experiments, which entailed both injecting the bee’s own pollen provisions and replacing the pollen provision with a preblended pollen mixture containing imidacloprid. Larval development, emergence, weight, and mortality were monitored and analyzed. There were no lethal effects found for either imidacloprid or clothianidin on O. lignaria and M. rotundata. Minor suble...

We investigated fitness in natural populations of a managed solitary bee Osmia ribifloris Cockerell (Hymenoptera: Megachilidae) from sites separated from 400 to 2,700 km. Parental wild bees originated in central Texas (TX), central-northern Utah (UT), and central California (CA). They were then intercrossed and raised inside a mesh enclosure in southern Mississippi (MS). Females from all possible mated pairs of O. ribifloris produced F1 broods with 30–40% female cocoons and outcrossed progeny were 30% heavier. Mitochondrial (COI) genomes of the four populations revealed three distinct clades, a TX-CA clade, a UT clade, and an MS clade, the latter (MS) representing captive progeny of CA and UT bees. Although classified as separate subspecies, TX and CA populations from 30° N to 38° N latitude shared 98% similarity in COI genomes and the greatest brood biomass per nest straw (600- to 700-mg brood). Thus, TX and CA bees show greater adaptation for southern U.S. sites. In contrast, UT-sourced bees were more distantly related to TX and CA bees and also produced ∼50% fewer brood. These results, taken together, confirm that adult O. ribifloris from all trap-nest sites are genetically compatible, but some phenotypic variation exists that could affect this species performance as a commercial blueberry pollinator. Males, their sperm, or perhaps a substance in their sperm helped stabilize our captive bee population by promoting legitimate nesting over nest usurpation. Otherwise, without insemination, 50% fewer females nested (they nested 14 d late) and 20% usurped nests, killing 33–67% of brood in affected nests.

During 1998-2003, we used populations of the solitary bee Osmia lignaria Say to pollinate a commercial sweet cherry orchard in northern Utah. Bee densities released each year ranged from 1290 to 1857 females/ha, with approximately twice as many males. Female progeny produced each year were greater than parental populations released, except in 2003, when nesting was poor due to bird predation. Despite poor weather during bloom, and in contrast to most other local producers, the study orchard produced harvestable crops in 1999 (2,964 kg/ha) and 2001 (3,154 kg/ha). In 1998 and 2000, record yields were obtained (10,625 and 12,096 kg/ha, respectively). Including only those years with harvestable crops, average production was 2.2 times higher in 1998-2003 (when O. lignaria populations were used) compared with 1992-1997 (when 10 Apis mellifera hives were used). This is the first study reporting multiyear cherry yields in an orchard pollinated with O. lignaria in North America.

The solitary bee Osmia lignaria Say continues to be developed as an orchard pollinator in the United States and southern Canada. Female bees are active during the early spring and construct nests consisting of a linear series of unlined cells delimited by mud partitions. Cells are provisioned with a pollen/nectar mass on which an egg is deposited, and nests are sealed with a mud plug. In 1997, we initiated two experiments on the development, mortality, and emergence of O. lignaria at selected laboratory temperature regimes and outdoors. In the first experiment (published previously), we compared temperature treatments for their adequacy in maintaining healthy O. lignaria populations. In a second experiment (reported here), we determined the relationship between rearing temperatures and prepupa–adult development rates as well as emergence time and longevity after wintering and incubation the following spring. We observed important differences in O. lignaria prepupa versus pupa responses to selected temperature treatments. The relationship between temperature and days to pupa was U-shaped, with additional time to transit the prepupa–pupa interval at temperatures above and below 26°C. The negative relationship between temperature and the length of the pupa to adult interval contrasts with the U-shaped thermal response observed for prepupae. Thus, with each increase in thermal heat units over the range of temperature treatments tested, we observed an additional reduction in the pupa–adult interval. Individuals reared at constant 18°C required 2.4 times as many days to transit the pupa–adult interval compared with those at constant 32°C. Our results suggest that there is a need for the development of regionally adapted, latitude-specific breeding populations of this outstanding orchard crop pollinator.

The alfalfa leafcutting bee, Megachile rotundata (F.), is a solitary, cavity-nesting bee and is the primary pollinator for alfalfa seed production. Bee management practices include cold storage during the prepupal stage. Fluctuating thermal regimes during cold storage increases survival of cold storage and allows a doubling of the cold storage period with no decrease in survival. However, survival, characterized as successful adult emergence, is not qualitative. In this study, we determined whether extended storage affects adult bee respiration or flight physiology. We overwintered bees for a single winter (current management protocol) or for 12 mo longer (extended storage). We used resting and tethered flight metabolic rates and resting critical PO2 (the oxygen partial pressure below which metabolism can no longer be sustained) as indices of adult bee quality. We found no significant differences in body mass, resting or flight metabolic rates, or critical PO2 between the two groups. Together these data indicate that extended storage of M. rotundata produces bees of similar respiratory capacity and flight ability. These findings could increase the use of M. rotundata as an alternative pollinator, allowing for extended storage to time adult emergence with early blooming crops.

Low, medium, and high stocking densities (15,000; 30,000; and 45,000–50,000 bees per acre, respectively) of Megachile rotundata (F.) (Hymenoptera: Megachilidae), the alfalfa leafcutting bee, were released over 4 yr in three research plots of Utah alfalfa, Medicago sativa L. (Fabaceae), planted at seed-production rates. A low percentage of bees (46–79% of released) survived the incubation and field-emergence processes; of those bees, the number of females that established at the nesting sites was 25–100%. Of the three field sites, one site consistently retained more females and produced more completed nests than the other sites, all of which usually had poor female establishment and progeny production. In addition, floral resources were depleted over time, but many flowers remained unvisited over the season. Nest production decreased over time, as numbers of flowers and female bees declined. Significant positive relationships were found between the intended stocking densities and 1) the number of females that actually survived incubation and field emergence and 2) the number of females that established nests. The number of females that established nests was positively affected by the number of females that survived to emerge in the field. The percentage of females that established was not significantly affected by the intended number of released bees, countering a prediction that the release of fewer bees would allow more females to establish nests and achieve high reproductive success. For growers, it may be more frugal to use modest numbers of M. rotundata for pollination, because many of the bees at medium and high stocking densities do not nest in grower-provided bee boards.

The effect of laboratory rearing temperature was related to chalkbrood susceptibility among larvae of the alfalfa leafcutting bee, Megachile rotundata (F.). Bees were reared at several temperature regimes. Mortality was inversely correlated with warmer rearing temperatures. LC 50 estimates (spores per larva) for the 6 temperature treatments ranged from 34 at constant 23°C to 8,511 at constant 37°C. Slopes of probit regression lines ranged from 0.77 to 1.29. Larvae died more quickly with increasing dose for all temperature treatments except for constant 37°C. Proportionally more cadavers sporulated with increasing dose for all treatments, but this proportion was lower on average for the warmer temperature treatments. Cadaver weight decreased with increasing dose, but was higher for bees reared at higher temperatures. Rearing temperature did not affect weight of adult females. Adult males reared at constant 37°C were significantly smaller than those reared at other temperatures. Spores of Ascosphaera aggregata Skou were tested for germinability in vitro using each of the temperature regimes. Fungus spores germinated equally well at all of the temperature regimes tested, but germination was slower at cooler temperatures.

A series of cage studies was used to investigate nesting preferences of Osmia sanrafaelae Parker. Bees preferred nest holes of smaller diameter in hlocks in the southwest corner of cages. Within multiple-block arrays, nests were more frequent in the lower left corner. The number of cells per nest was lowest in cages with the fewest nests in some experiments, but did not vary in others. Nest diameter affected neither the proportion of females produced per nest, nor the size of offspring of either sex. Rather, nesting females constructed cells of varying orientations within nests of different diameters, a process that resulted in approximately equal cell volumes. In one experiment, the sex ratio of bees released within cages was varied and influenced the sex ratio of offspring. Nests within cages in which a higher ratio of males to females was released contained a higher proportion of female offspring.

Temperatures within polystyrene, paper, and wooden nests of the alfalfa leafcutting bee, Megachile rotundata (F.), were monitored during incubation at 30.0 °C. In the polystyrene and paper nests, temperatures began to increase 10 d after the start of incubation and continued to rise until adults emerged. Temperatures reached 35.1 °C In the polystyrene and 37.2 °C in the paper, while the wood nests never exceeded 30.6 °C. Compared with air temperature degree-days, cells in the center of the polystyrene nests accumulated up to 28 more degree-days after 20 d, whereas those in the paper accumulated 32 more degree-days. In polystyrene nests, heat units increased toward the center holes, reaching a maximum at the fifth row toward the center. Emergence was also monitored from polystyrene blocks, wooden boards, and loose cells. Females emerged 2.1 d later from polystyrene blocks compared with wooden boards and the emergence period was lengthened by 2.5 d. Because of the increased temperatures and the prolonged emergence period experienced with these materials, we recommend that filled polystyrene and paper nests not be incubated but be stripped of their cocoons as they were designed.

Bioassays were conducted to determine effects of diet on chalkbrood, a devastating mycosis of larvae of the alfalfa leafcutting bee, Megachile rotundata (F.). Bees reared in the laboratory on four sterilized diets were assayed for chalkbrood susceptibility, bee development, and survival. Bees reared on a pollen + pollen substitute diet had an LC50 of 138 spores per larva, the shortest time to death, the smallest cadaver size, and the most frequent sporulation. Uninfected larvae developed into normal adults. Bees reared on a pollen + sugars diet had the lowest LC50 (79 spores per larva), the longest time to death, and the lowest sporulation rate. On this diet, the development and survival of uninfected larvae were delayed, and only a few small adults emerged. Bees reared on a pollen + sugars diet with a higher pollen concentration had the highest LC50 (1,549 spores per larva). These bees also had delayed development and reduced survival to the adult stage, but adults were of normal size. Bees reared on sterilized natural provisions had an intermediate LC50 (219 spores per larva), time to death, and sporulation rate, and the largest cadaver size. Development and survival rates for bees reared on this diet were similar to those obtained with bees reared on the pollen + pollen substitute diet, but adults were much larger than those reared on any other diet. Thus, diet factors that promote bee survival and growth are different from those that determine chalkbrood susceptibility.

The alfalfa leafcutting bee, Megachile rotundata (F.), was used as a representative nonsocial bee on which to test the toxicity of carbaryl bran bait, which is applied commonly to control rangeland grasshoppers. Adult M. rotundata fed a 25% honey solution containing 1 or 2 mg of carbaryl bran bait continuously for either 20 or 40 d did not experience significantly greater mortality than bees fed plain honey solution. Mortality of M. rotundata larvae fed pollen-nectar provisions contaminated with or 2 mg of carbaryl bran bait was 17–30% compared with 9–11% for bees fed uncontaminated provisions. The likelihood that foraging females could collect such a large amount of bait while provisioning their progeny is low. We conclude that carbaryl bran bait is relatively safe to both adult and larval M. rotundata at levels likely to be experienced in the field and that it is likely to be equally safe for native bee pollinators as well.

Populations of the alfalfa pollinator, Megachile rotundata (F.), nest in dense aggregations in artificial shelters in alfalfa fields. The closeness of the nest tunnels in these shelters (5–10 mm apart) fosters what appear to be numerous counterproductive interactions among females. By increasing the distance between nest tunnels and patterning the nest-block surface to help orient bees returning from foraging trips, we sought to reduce interactions and to increase both the number of progeny produced and the proportion of female progeny. Only the nest-block surface aspect affected progeny production rate; progeny were produced at a Significantly faster rate in patterned blocks than in plain blocks. Intertunnel distance did not affect progeny production rate. Nest blocks with tunnels 5 mm apart contained a significantly greater proportion of female progeny than did nest blocks with tunnels spaced farther apart. Sex ratio of progeny was unaffected by surface patterning of blocks. Females strongly preferred patterned to plain blocks and preferred 5-mm and 2-cm intertunnel distances to those of 4 cm. Beekeepers and alfalfa growers can increase bee productivity by supplying patterned nest blocks with intertunnel distances between 5 and 10 mm.

A significant concern in the commercial application of the alfalfa leafcutting bee, Megachile rotundata (F.) (Hymenoptera: Megachilidae), for pollination is synchronizing bee emergence and activity with peak crop bloom. Previous studies have demonstrated that the commercial spring incubation of this species can be successfully interrupted by low temperature incubation, thereby slowing development and giving producers flexibility in timing emergence to weather conditions or crop bloom. In this study, we demonstrate that the implementation of a fluctuating thermal regime, during which bees are given a daily one hour pulse of high temperature, markedly increases the “shelf-life” of individuals of this species. Although constant temperatures can be used to store bees for up to 1 wk with no decrease in survival, properly staged bees can be stored for up to 6 wk without a significant decrease in percentage of emergence. Hence, we expect fluctuating thermal regime protocols to become a valuable tool for M. rotundata managers, especially when timing nesting activity with peak bloom to maximize effectiveness.

Osmia lignaria Say (Hymenoptera: Megachilidae) can be used to pollinate fruit trees. Populations are sometimes difficult to sustain because some female bees fail to establish at provided nesting sites. We address the hypothesis that rough handling of overwintered O. lignaria results in decreased establishment. We tested this by shaking (200 rpm for 2 min) overwintering bees as a proxy for rough handling. Bees were then released in an orchard, and nest establishment of shaken and unshaken bees was recorded. There was no significant difference in the proportion of shaken and unshaken females that nested, indicating that rough handling of overwintering bees does not discourage nest establishment.

The relative efficiency of two groups of insects that pollinate strawberries, Fragaria × ananassa Duch., was established by comparing the relative influence of the number and length of their visits to flowers, their foraging behavior, and the fertilization rates of the achenes during these visits. Honey bees, Apis mellifera L., were more efficient than indigenous pollinators (Hymenoptera: Andrenidae, Halictidae, and Megachilidae) when the frequency of visits was low. Average-sized apoids such as honey bees pivot at the top of the receptacle and pollinate the apical stigmata, whereas small apoids circle on the stamens and around the receptacle, pollinating mainly the basal stigmata. Data collected revealed that these two groups of pollinators playa complementary role in strawberry pollination.

A useful technique for synchronizing pollinators with the alfalfa, Medicago savita L. (Fabaceae), bloom is to interrupt the late spring incubation of developing bee pupae and pharate adults of Megachile rotundata (F.) (Hymenoptera: Megachilidae) with short-term low-temperature storage. However, low-temperature exposure can be stressful depending on the temperature, duration of exposure, and the developmental stage exposed. To evaluate the effect of low-temperature storage after development had already been initiated by exposure to 29 degrees C, three developmental stages (eye pigment pupae, body pigment pupae, and pharate adults ready to emerge) were exposed to 6, 12, or 18 degrees C for durations up to 28 d. The effect of delaying the termination of overwintering storage (6 degrees C) from April to July ("April bees" and "July bees," respectively) also was examined. The following observations were made: (1) All developmental stages of the April bees and July bees examined can be stored without harmful effects, as measured by percentage of survival, for 14 d at 12 degrees C and above. (2) Postponing the termination of overwintering storage at 6 degrees C from April until July significantly decreased the mean number of days to 95% adult emergence in the July ready-to-emerge bees across the three temperatures tested. (3) The increase of overwintering duration also caused a decrease in the postemergence longevity of the ready-to-emerge adults stored at 6 degrees C for 14 d. (4) Of the three storage temperatures examined, 18 degrees C seems to be the optimal storage temperature for short-term storage of developing bees because of their slow but continuing development without increased mortality either during storage or after emergence.

Cavity-nesting alfalfa leafcutting bees, Megachile rotundata (F.) (Hymenoptera: Megachilidae), are excellent pollinators of alfalfa, Medicago savita L., for seed production. In commercial settings, artificial cavities are placed in field domiciles for nesting and, thereby, bee populations are sustained for future use. For this study, cells from leafcutting bee nests were collected in late summer from commercial seed fields. Over 3 yr (2003-2005), 39 samples in total of approximately equal to 1,000 cells each were taken from several northwestern U.S. states and from Manitoba, Canada. X-radiography of 500 cells from each sample was used to identify "pollen balls" (i.e., cells in which the pollen-nectar provision remained, but the egg or larva, if present, was not detectable on an x-radiograph). Most U.S. samples seemed to have higher proportions of pollen ball cells than Manitoba samples. Pollen ball cells were dissected to determine the moisture condition of the mass provision and true contents of each cell. Most pollen ball cells from Manitoba samples contained fungus, the frequency of which was positively correlated with cool, wet weather. In the United States, most pollen ball cells had moist provisions, and many of them lacked young brood. Correlation analysis revealed that pollen ball cells occurred in greater proportions in fields with more hot days (above 38 degrees C). Broodless pollen ball cells occurred in greater proportions under cool conditions, but dead small larvae (second-third instars) seemed to occur in greater proportions under hot conditions. Pollen ball cells with unhatched eggs and first instars (in the chorion) occurred in lesser proportions under hot conditions.

The alfalfa leafcutting bee, Megachile rotundata (F.), is a solitary, cavity-nesting bee that has been managed in large numbers to pollinate alfalfa, Medicago spp., seed crops since the 1960s. Propagation of these bees from 1 yr to the next has been seriously hampered by chalkbrood, a larval disease caused by the fungus Ascosphaera aggregata Skou. In the United States, attempts to control the disease have been fairly unsuccessful, but include removing nests from the nesting boards and then disinfecting the boards with heat treatments or a fumigant. The problem is that many boards are made of polystyrene (so heat cannot be used), and very few fumigants are registered for this use. In this study, ozone was tested as a fumigant and compared with heat treatments and methyl bromide fumigation. Ozone was found to be inadequate for killing A. aggregata spores and for reducing chalkbrood levels in the field. Methyl bromide and heat treatments did greatly reduce spore viability in the boards, but did not reduce chalkbrood levels in the field. Surprisingly, larvae in new nesting boards (boards free of contamination) and chalkbrood infection levels were similar to those from nests in contaminated, used boards. Disinfecting nesting boards may be necessary for controlling chalkbrood, but the results reported here indicate that it is not sufficient in and of itself. Some other source of spores was present in the field that was greater than the effect of contamination from the boards, but the source still needs to be determined.

Since the 1970s, it has become increasingly difficult for U.S. alfalfa seed producers to maintain Megachile rotundata (F.) populations used for alfalfa, Medicago sativa L., pollination. In 1998, we monitored M. rotundata population dynamics and foraging behavior, as well as alfalfa bloom and pollination rates in two fields in eastern Oregon. Despite marked differences in bee management, establishment was very similar in the two fields (approximately 0.5 females per nesting cavity) and lagged peak bloom by approximately 2 wk. Pollination rates increased from 0-10% in the first 3 wk to 80-90% in week 4-5. By then, M. rotundata females had difficulty finding untripped (nonpollinated) flowers and visited large numbers of already tripped or not fully matured flowers. M. rotundata progeny mortality was very high (54-78%). Estimated seed yields were similar in both fields. We contend similar seed yields, and improved bee production, could be accomplished with smaller bee populations, better timed with alfalfa bloom.

Postdiapause prepupae of Megachile rotundata (F.) were subjected to a number of temperature-time incubation regimes, A drastic reduction in percent emergence was observed for all incubation regimes when the final incubation temperature was 20°C. A temperature of 25 or 30°C resulted in high percent emergence for all incubation regimes, including treatments where development was retarded for up to 36 days. We recommend that postdiapause prepupae be incubated at 30°C. If field conditions just before field release are unfavorable, we recommend retardation of development at 15 to 20°C. Unfavorable conditions after field placement could be mitigated by a second incubation at 30°C.

Uninfested cocoons of the alfalfa leafcutter bee, Megachile rotundata (F.), bare prepupae, and cocoons infested with the chalcidoid, Pteromalus venustus Walker, were exposed to vapor from dichlorvos resin strips at rates of 0.16, 0.37, 0.71, and 0.99 strip per 28.3 m3 during 4 days of incubation. For uninfested cocoons, there was no significant reduction in leafcutter bee survival at the dichlorvos rates tested. For bare prepupae, there was a significant linear trend of reduced survival with increased dichlorvos rate. Cocoons apparently protect the bees from dichlorvos vapor. For infested cocoons, 0.71 strip per 28.3 m3 reduced the parasitism from 40 to 13% which significantly increased bee survival. Adult parasites were placed in different housing environments and exposed to the 0.71 strip per 28.3 m3 rate. When parasites were directly exposed to dichlorvos vapor, mortality occurred within 2 h. When parasites were shielded by layers of cocoons, mortality was delayed 7 to 10 h. Because 25% parasitism can occur within 4 h of P. venustus emergence, it is unlikely that dichlorvos resin strips alone will totally control parasitism in a commercial operation.

Field trials of two fungicides, Captan and Thiabendazole, were conducted to determine their influence on chalkbrood disease of the alfalfa leafcutting bee ( Megachile rotundata [F.L]). Captan effectively controlled the disease when applied triweekly; similar applications of Thiabendazole were ineffective in controlling this disease. Some disruption in nesting behavior was observed, but total cell production in all treatments was similar. No interactions were evident between mortalities of immature stages from any cause other than chalkbrood. Chalkbrood disease appears to be density dependent and shelter size may be a factor in disease control.

A fungicide (Vitavax, carboxin with captan) was field tested to determine its effect on chalk brood disease of Megachile rotundata (F.). The fungicide was blown into the holes of nesting materials. Disease incidence was significantly reduced when nesting units were treated once a week. Fungicide application had no effect on nesting behavior (number of cells per nest) or immature mortality (from causes other than chalkbrood) when treated and untreated nests were compared. Thus, fungicide applications reduced fungal growth but did not disturb nesting of the bees.