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Squash bee

The squash bee is a common name for a group of solitary bees belonging to the genera Peponapis and Xenoglossa in the family Apidae and tribe Eucerini, which are specialized pollinators of plants in the genus Cucurbita, including economically important crops such as squash, pumpkins, zucchini, and gourds.^[1]^[2] Native to the Americas and ranging from Canada to Mexico, these bees exhibit adaptations that make them highly efficient at pollinating cucurbit flowers, often outperforming generalist pollinators like honey bees in agricultural settings.^[1]^[3] Physically, squash bees are robust and slightly larger than honey bees, measuring 12–18 mm in length depending on the species, with a bulky build, long antennae, and a rounder face that aids in accessing the deep corollas of cucurbit flowers.^[4]^[3] Their coloration varies but typically includes a black, tan, or orange head and thorax covered in dense hairs—pale orange or yellow on the thorax—while the abdomen features black segments with white, tan, or yellowish bands; females possess extensive fuzzy hairs on their hind legs for carrying dry pollen, unlike the pollen baskets of honey bees.^[4]^[1] Males often have a distinctive yellow or white spot on the lower face (clypeus), and both sexes are active primarily in the early morning, foraging from pre-dawn until mid-morning when cucurbit flowers open.^[3]^[4] In their life cycle, squash bees are solitary and univoltine in many regions, with adults emerging in mid-summer (June–August in the eastern U.S.) to mate and forage exclusively on Cucurbita pollen and nectar, which they use to provision underground nests for their offspring.^[2]^[3] Females excavate vertical burrows 6–24 inches deep in bare or sparsely vegetated soil near host plants, creating individual cells stocked with a pollen-nectar loaf and a single egg; males may roost overnight in wilted flowers or shallow soil nests.^[1]^[3] Although non-social, they can nest gregariously in aggregations, leading to high local densities that enhance pollination efficiency.^[1] As obligate specialists, squash bees are vital for the reproduction of wild and cultivated Cucurbita species, with just 6–10 female visits sufficient to fully pollinate a single flower, contributing to about two-thirds of commercial squash pollination in the U.S..^[3] In 2025, squash bees were recognized as Pollinator of the Year by Pollinator Partnership for their vital role in cucurbit pollination.^[5] Their historical role likely supported the domestication of cucurbits by indigenous peoples across the Americas, and today they remain essential in organic and no-till farming systems where they can outpollinate introduced bees.^[1]^[2] However, populations face threats from soil disturbance via tillage, which destroys nests, and neonicotinoid pesticides like imidacloprid, which can cause up to 89% reproductive failure; conservation efforts emphasize preserving untilled soil, mulching lightly, and avoiding insecticides during bloom.^[3]

Taxonomy

Etymology and nomenclature

The common name "squash bee" derives from these bees' obligate specialization in pollinating flowers of plants in the genus Cucurbita, which encompasses squashes, pumpkins, and gourds.^[6] The genus Peponapis was established by Charles Robertson in 1902 to classify bees adapted specifically for cucurbit pollination; the name combines the Greek pepon (referring to a gourd or melon) and apis (bee).^[7]^[8] The type species, Peponapis pruinosa, was originally described by Thomas Say in 1837 as Anthophora pruinosa in the Boston Journal of Natural History. A major taxonomic revision of the squash bees occurred in 1970, when Paul D. Hurd Jr. and E. Gorton Linsley provided a comprehensive classification of the genera Peponapis and Xenoglossa, clarifying their relationships within the tribe Eucerini.^[7] Squash bees are placed in the family Apidae (true bees) and tribe Eucerini (long-horned bees), a group known for floral specialization.^[9] They should not be confused with cuckoo bees of the genus Nomada (tribe Nomadini), which are cleptoparasites that may target squash bee nests but lack the dense pubescence and cucurbit specialization of Peponapis. The genus Peponapis encompasses approximately 15 species across the Americas, with details on individual taxa provided elsewhere.^[7]^[10]

Species and genera

Squash bees are classified within two principal genera of the tribe Eucerini: Peponapis, encompassing approximately 15 species predominantly distributed across North America, and Xenoglossa, including 7 species largely restricted to the southwestern United States and northern Mexico.^[10]^[11] These genera represent the core of the squash-specialist bees, with Peponapis species exhibiting broader continental ranges and Xenoglossa showing more localized adaptations to arid environments. Peponapis includes several subgenera, such as Peponapis and Eopeponapis.^[12]^[13] Key species within Peponapis include P. pruinosa, which is the most extensively studied due to its wide distribution across much of North America from southern Canada south to central Mexico and its association with cultivated cucurbits.^[1]^[14]^[15] In western regions, P. limitaris serves as a prominent representative, occurring along the Pacific coast from California to British Columbia and demonstrating specialized foraging traits.^[16] For Xenoglossa, X. strenua stands out as the large squash bee, characterized by its robust build and prevalence in the southwestern deserts, where it contributes significantly to pollination dynamics.^[17] Phylogenetically, squash bees maintain close affinities with other long-horned bees in the Eucerini tribe, forming a specialized clade tied to Cucurbita pollination. Recent genetic analyses, including ultraconserved element sequencing in 2023, affirm the monophyly of the Peponapis and Xenoglossa group, highlighting their shared evolutionary history predating human agriculture.^[18] DNA barcoding initiatives from the 2020s have bolstered species delineation within this clade, revealing cryptic diversity and reinforcing their distinct lineage amid broader Eucerini relationships.^[19]

Physical characteristics

Morphology

Squash bees, belonging to the genera Peponapis and Xenoglossa in the tribe Eucerini, possess a robust body structure adapted to their solitary lifestyle and specialization on cucurbit pollination. Adults typically measure 11 to 18 mm in length, with females slightly larger than males at 12.5–14 mm and males at 11–13 mm. They have a rounder face compared to honey bees, aiding access to the deep corollas of cucurbit flowers. Their bodies are notably hairy, featuring dense pubescence that covers the head, thorax, and legs, which facilitates the collection and transport of pollen. The head is adorned with long, erect, light yellowish-brown hairs, while the thorax bears shorter, denser, reddish-yellow or tawny pubescence, contributing to their overall bulkier appearance compared to honey bees.^[20]^[21]^[22] A key specialized feature is the scopa on the hind legs of females, consisting of pale yellowish-brown, sparsely plumose (branched) hairs on the tibia, designed to efficiently gather and hold the large, sticky pollen grains unique to Cucurbita species. Males lack this scopa, as they do not collect pollen. The mouthparts include a protruding clypeus—black in females and mostly black with a yellow lower rim in males—and an elongated proboscis characteristic of long-tongued bees in the Eucerini, enabling access to nectar deep within the tubular corollas of squash flowers.^[20]^[23]^[24] Sensory adaptations support their floral specialization, with large compound eyes positioned close to the clypeus (leaving a very small space between them), enhancing the detection of visual cues from flowers during early morning foraging. The antennae, black and segmented into 12 parts in females (long scape, short pedicel, and 10-segment flagellum) and 13 in males, bear sensilla that detect pheromones and chemical signals essential for mate location and navigation. These features underscore the morphological convergence in squash bees for efficient interaction with their primary host plants.^[20]^[23]

Size and coloration variations

Squash bees exhibit sexual size dimorphism, with females typically larger than males to support pollen collection and nesting activities. In the common species Peponapis pruinosa, females range from 12 to 14 mm in length, comparable to or slightly larger than the average honey bee (13 mm), while males are smaller and more slender.^[4]^[20] This dimorphism follows patterns observed in many solitary bees, where female body mass can be 1.5 to 2 times that of males.^[25] Interspecific variation further diversifies sizes within the squash bee group; for example, species in the genus Xenoglossa, such as X. strenua and X. fulva, attain lengths of 14 to 18 mm for both sexes, rendering them bulkier than Peponapis congeners.^[4]^[26] Coloration in squash bees is predominantly black with dense pubescence that provides camouflage and sensory functions, varying by genus and sex. Peponapis pruinosa features a black integument overlaid with pale orange to reddish-yellow hairs on the thorax and off-white or whitish bands across the abdominal tergites, creating a frosted appearance.^[20]^[21] In contrast, Xenoglossa species display similar black bases but with more pronounced yellowish bands on the abdomen and yellow markings at the base of the mandibles, enhancing their distinction from Peponapis.^[27]^[28] Sexual dimorphism manifests in facial coloration, where males possess a yellow or cream-colored clypeus and often appear duller due to sparser or less vibrant pubescence compared to the uniformly black clypeus and denser hair coverage of females.^[3]^[27] Geographic variation in squash bee morphology, including potential size differences, has been documented through morphometric analyses of populations across North America. Studies using geometric wing morphometry on P. pruinosa specimens from sites spanning Mississippi to Ontario revealed significant differentiation in wing shape between eastern and western populations, with southern groups showing distinct traits potentially influenced by local resource availability and climatic factors.^[29] Such variations underscore adaptive responses to regional environments, though direct body size metrics remain less quantified in these comparisons.^[29]

Life history

Developmental stages

The developmental stages of squash bees, such as Peponapis pruinosa, follow a complete metamorphosis typical of Hymenoptera, encompassing egg, larva, pupa, and adult phases within a solitary, univoltine lifecycle that spans one year.^[30] Females construct underground nests consisting of multiple cells, each provisioned with a pollen loaf composed primarily of Cucurbita (squash family) pollen mixed with nectar to form a compact mass suitable for larval feeding.^[31] A single egg is laid upright on or near this provision in each waterproof cell, which the female seals before moving to the next; this solitary provisioning ensures each offspring has dedicated resources without competition from siblings.^[30] The egg is an arcuate, white, cylindrical structure with rounded ends, measuring approximately 1-2 mm in length.^[32] Upon hatching, the larva—a legless, grub-like form—consumes the entire pollen loaf over the course of its development, progressing through five instars characterized by increasing size and changes in head capsule width.^[32] Larval growth occurs rapidly during the active summer season, with the insect relying solely on the maternal provision for nutrition, as squash bees exhibit no progressive provisioning like some social bees.^[30] By the final instar, the larva has fully depleted the food source and defecates, forming a fecal pellet that is pushed aside; it then molts into a prepupal stage and enters diapause, remaining dormant through winter in the capped cell to avoid unfavorable conditions.^[31] This overwintering diapause in the final larval or prepupal form is a key adaptation, synchronizing the population with the annual Cucurbita bloom cycle.^[30] In late spring or early summer, the prepupa initiates pupation within a silken cocoon inside the cell, undergoing metamorphosis over 7-10 days into the imago (adult) stage.^[32] Adult emergence is precisely timed to coincide with the opening of squash flowers, typically from late May to August depending on latitude, with males often appearing 7-10 days before females to establish territories near blooming patches.^[30] The entire lifecycle, from egg deposition to adult eclosion, thus aligns with the seasonal availability of host plants, reinforcing the bees' obligate oligophagy on Cucurbita.^[31]

Reproduction and mating

Squash bees employ a mating system in which males actively patrol flowers of Cucurbita species in search of receptive virgin females, often aggregating at bloom sites to increase encounter rates. Males preferentially visit staminate flowers, where the probability of locating unmated females is higher, and they frequently sleep inside closed blossoms overnight to resume searching at dawn when flowers reopen.^[33]^[1] Females are typically monandrous, mating with only a single male and rarely exhibiting polyandry, while males are polygynous and attempt to mate with multiple partners. Following mating, females provision individual nest cells with pollen and nectar collected exclusively from Cucurbita flowers before laying a single egg in each; a typical female provisions around 5 to 10 cells per nest across one or more nests per season.^[30]^[34] As solitary bees, squash bees provide no biparental care, with females sealing each provisioned cell with a mud cap immediately after oviposition and abandoning the nest thereafter.^[35]^[36] Reproduction in squash bees is univoltine and tightly synchronized with the flowering phenology of Cucurbita plants, ensuring that adult emergence aligns with peak nectar and pollen availability to support foraging, mating, and nest provisioning.^[30]

Habitat and distribution

Native ranges

Squash bees of the genus Peponapis, including the common species P. pruinosa (classified as Eucera (Peponapis) pruinosa), are native across North America, with P. pruinosa having a primary range in eastern North America extending from southern Canada, including Quebec, southward through the United States (excluding the Great Plains and Great Basin regions) to central Mexico.^[15]^[1] In contrast, bees of the genus Xenoglossa are primarily distributed in the southwestern United States and northern Mexico, with some species extending into Central America.^[28]^[37] These ranges align closely with the natural distribution of wild Cucurbita species, upon which squash bees are specialized pollinators.^[14] Habitat preferences for both genera center on areas suitable for ground nesting near host plants, particularly sandy or well-drained soils adjacent to wild relatives of domesticated squash such as Cucurbita foetidissima.^[38] These bees construct solitary nests in bare or sparsely vegetated ground, often in aggregations close to Cucurbita patches, and their presence is limited to regions supporting these plants, including up to elevations of approximately 1,000 m in suitable habitats.^[39] The historical distribution of squash bees reflects their long co-evolutionary ties to Cucurbita, predating human domestication, with expansion across North America linked to the pre-Columbian spread of cultivated squash from Mesoamerican origins.^[14] Archaeological evidence from sites like Guilá Naquitz Cave in Oaxaca, Mexico, includes Cucurbita pepo remains—such as seeds, peduncles, rind fragments, and associated pollen—dated to around 10,000 years ago (approximately 8,000 BCE), indicating early squash cultivation that facilitated bee range expansion.^[40]^[41]

Expansion and introductions

The spread of squash bees, particularly the species Peponapis pruinosa (classified as Eucera (Peponapis) pruinosa), beyond their native Mexican origins has been closely tied to human agricultural practices involving Cucurbita crops. Originating in regions of Mexico where wild Cucurbita species occur, these bees expanded northward across North America as Indigenous peoples domesticated and cultivated squashes like Cucurbita pepo around 5,000–10,000 years ago, creating reliable floral resources that acted as stepping-stones for dispersal. This facilitated a rapid geographical range extension, with populations establishing in the eastern United States via the Midwest and in the western United States, including California, through independent northward migrations. Recent expansions include the first confirmed records of P. pruinosa in Oregon in 2016, highlighting ongoing facilitation by agriculture in the Pacific Northwest.^[42]^[43] In California, squash bee populations were further supported by modern crop transport and experimental introductions of pre-nesting females to enhance pollination in commercial fields, demonstrating how agricultural activities can locally bolster or relocate these specialist pollinators. While primarily a natural expansion driven by pre-Columbian agriculture, such human-mediated movements highlight the bees' reliance on cultivated Cucurbita habitats, with accidental dispersal potentially occurring through infested soil or plant material in farming operations. Established populations in non-native agricultural zones, such as parts of the Intermountain West, reflect this ongoing facilitation by intensive squash cultivation.^[44]^[43] Genetic analyses reveal low gene flow among introduced or expanded populations, with northern groups exhibiting reduced diversity compared to southern core areas—such as allelic richness as low as 1.26 and heterozygosity of 0.084 in California samples—indicative of founder effects and bottlenecks during dispersal. Recent studies from the 2020s confirm adaptation to novel climates in these expanded ranges, where populations have undergone rapid growth over the past 1,000 years alongside agricultural intensification; for instance, approximately 20% of the genome in eastern lineages shows selective sweeps affecting sensory genes, enabling persistence in human-dominated landscapes. This low connectivity persists, as western and eastern lineages show minimal admixture, underscoring the bees' dependence on fragmented agricultural niches for survival.^[43]^[45]

Behavior

Foraging patterns

Squash bees, primarily species in the genus Peponapis such as P. pruinosa, display an oligolectic foraging strategy characterized by exclusive specialization on flowers of the Cucurbitaceae family, particularly the genus Cucurbita encompassing squashes, gourds, and pumpkins. Females collect pollen solely from these sources to provision their brood, while males and females both gather nectar from the same blooms, reflecting their adaptation to this narrow resource base. To harvest the heavy, sticky pollen presented on exposed anthers, they utilize a vibratory technique known as sonication, contracting flight muscles to shake the flower and release grains, which are then gathered dry using specialized scopal hairs on the hind legs.^[46]^[47] Foraging activity aligns closely with the ephemeral nature of Cucurbita flowers, which open predawn and close by late morning. Squash bees initiate visits as early as 5 AM, often in low light conditions, peaking between sunrise and 9 AM to coincide with peak floral availability and nectar flow, ceasing as flowers wilt around midday. This dawn-focused pattern allows them to outpace generalist pollinators like honey bees, with females achieving high collection efficiency; their scopal loads typically comprise over 90% Cucurbita pollen, underscoring their dietary fidelity and rapid handling times per flower.^[3]^[48]^[49] Resource switching to non-Cucurbita flowers occurs rarely, limited mostly to occasional nectar foraging during scarcity, as their physiology is ill-suited to alternative pollens containing defensive compounds like cucurbitacins. Navigation to patches relies on visual cues from the large, bright yellow flowers serving as landmarks, complemented by olfactory detection of floral volatiles, facilitating efficient localization even across fragmented agricultural landscapes. These behaviors, supported by morphological features like the dense hind-leg scopa, optimize their role as specialized foragers.^[45]^[34] Squash bees, primarily Peponapis pruinosa, are solitary ground-nesters that excavate burrows in loose, well-drained soil, often in aggregations near patches of their host plants in the genus Cucurbita. These nest sites are typically located at the edges of squash fields or under plants, where females dig individual entrances to access foraging resources efficiently. Aggregations commonly consist of 10 to 100 nests per square meter, facilitating concentrated nesting without communal cooperation.^[3]^[34]^[50] Nest architecture features a main vertical tunnel, usually 10 to 30 cm deep, from which lateral branches lead to 4 to 6 individual brood cells per nest. Each cell is provisioned with pollen and nectar collected from squash flowers, then sealed with a waterproof lining secreted by the female's Dufour's gland to protect the developing offspring from moisture and pathogens. Soil excavated from the main tunnel is pushed to the surface, forming small mounds, while lateral tunnels are often backfilled after cell construction. Brood cells are positioned at shallower depths within this range to optimize conditions for larval development.^[51]^[34]^[52] Despite forming dense aggregations, squash bees exhibit solitary social dynamics with no division of labor, queen-worker castes, or cooperative brood care; each female independently constructs, provisions, and seals her own nest for a single brood per season. Interactions among females in aggregations are minimal and non-cooperative, though occasional kleptoparasitism by parasitic bees such as Triepeolus species can occur, where intruders steal provisions from unguarded cells. This semi-social aggregation pattern enhances local population density but does not evolve into eusocial structures.^[3]^[53]^[50]

Ecological importance

Pollination services

Squash bees (Peponapis and Xenoglossa spp.) are highly specialized pollinators of plants in the genus Cucurbita, rendering them more effective than generalist pollinators like honey bees (Apis mellifera) for achieving successful fruit and seed set in these species.^[54] Their efficiency stems from early morning foraging synchronized with Cucurbita flower anthesis, combined with behaviors that facilitate substantial pollen removal and deposition per visit—up to seven times more pollen removed than by honey bees.^[54] Additionally, squash bees employ sonication, vibrating their flight muscles to dislodge sticky pollen from the flowers' anthers, ensuring effective transfer to stigmas and higher reproductive success compared to non-sonicating visitors.^[55] This specialization results in superior pollination outcomes, with studies demonstrating greater positive effects on fruit set likelihood and seed production in Cucurbita pepo when squash bees are the primary visitors.^[54] In agricultural contexts, squash bees play an essential role in the pollination of economically vital Cucurbita crops, including pumpkins, squashes, and gourds, which collectively generate over $450 million in annual value for U.S. producers as of 2024.^[56]^[57] The U.S. pumpkin sector alone contributed more than $274 million in 2024, with squash bees often providing sufficient natural pollination to support high yields without supplemental honey bee hives.^[56] Research from 2015 to 2025 has quantified yield benefits, showing that abundant squash bee activity correlates with optimal pollen deposition and fruit production in commercial fields, reducing the need for managed pollinators and enhancing crop reliability in regions like Florida and the Northeast.^[58]^[59]^[17] However, a 2025 study found that pollination services to squash may be insufficient despite abundant squash bees, influenced by local and landscape-level land management.^[60] For instance, fields with high squash bee visitation exceed pollination thresholds for Cucurbita crops, leading to consistent fruit set and larger harvests.^[59] Beyond crops, squash bees sustain wild Cucurbita populations, such as the buffalo gourd (Cucurbita foetidissima), by facilitating pollen transfer that promotes gene flow and maintains genetic diversity across fragmented habitats.^[61]^[43] These native plants exhibit strong seasonal dependency on squash bees, as their brief flowering periods align precisely with bee emergence, ensuring reproduction in natural ecosystems from the southwestern U.S. to Mexico.^[46] This mutualism underscores the bees' broader ecological contributions, supporting biodiversity in arid and semi-arid wildlands where alternative pollinators are scarce.^[61]

Interactions with plants and other insects

Squash bees, particularly species in the genera Peponapis and Xenoglossa, exhibit a high degree of plant specificity, having coevolved with plants in the genus Cucurbita over millennia. Genomic analyses reveal that agricultural expansion of Cucurbita cultivation in North America drove rapid population growth and adaptive evolution in squash bees, with approximately 20% of the genome showing selective sweeps associated with sensory functions like olfaction, enabling specialization on cultivated crops.^[62] This oligolectic behavior restricts female squash bees to collecting pollen almost exclusively from Cucurbita flowers, while males may visit other plants for nectar, reflecting a tight mutualistic relationship shaped by domestication that has not significantly altered key floral rewards like nectar volume or pollen protein content.^[63] Floral volatiles from Cucurbita play a crucial role in attracting squash bees, with specific compounds mediating this exclusive interaction. For instance, the volatile (E)-cinnamaldehyde uniquely draws squash bees to flowers without attracting herbivorous cucumber beetles, whereas 1,2,4-trimethoxybenzene appeals to both pollinators and herbivores, highlighting evolutionary trade-offs in volatile profiles.^[64] These chemical cues ensure targeted visitation, reinforcing the bees' dependence on Cucurbita as their primary resource. At Cucurbita flowers, squash bees overlap with generalist competitors such as bumblebees (Bombus spp.) and honeybees (Apis mellifera), which also seek nectar and pollen from these plants. However, niche partitioning occurs temporally: squash bees initiate foraging at dawn when Cucurbita flowers open, ceasing activity by midday as flowers wilt, while honeybees and bumblebees typically forage later in the day, reducing direct resource contention.^[31] This temporal separation allows squash bees to dominate early pollination without substantial interference, though high densities of generalists can still limit access in intensive agricultural settings. Parasitic interactions primarily involve cuckoo bees in the genus Triepeolus, such as T. remigatus, which target squash bee nests to steal provisions. These kleptoparasites lay eggs in host nests (Peponapis pruinosa or Xenoglossa pruinosa) during the host's foraging absences, with larvae consuming the stored Cucurbita pollen; interactions are largely non-aggressive, characterized by tolerance or avoidance rather than confrontation. Predators of squash bees include ants and spiders, which exploit the bees' ground-nesting habits and floral foraging. Ants raid nests for eggs and provisions, while crab spiders (Thomisidae) ambush adults on flowers, contributing to mortality rates in vulnerable aggregations.^[65]

Conservation

Threats and declines

Squash bees, as ground-nesting solitary pollinators, face substantial population reductions due to habitat loss driven by agricultural intensification and urbanization. Agricultural practices such as tillage and monoculture farming destroy or fragment the loose, sandy soils preferred for nesting, leading to isolation from natural habitats like woodlands and reducing bee abundance by up to 3- to 6-fold in intensively farmed areas compared to organic sites with surrounding natural cover.^[66] Urbanization exacerbates this by paving over potential nesting sites, further limiting access to suitable soil and floral resources.^[67] Pesticide exposure, particularly from neonicotinoids, severely impacts squash bee foraging and reproduction. Soil applications of imidacloprid, commonly used on cucurbit crops, reduce nest establishment by 76-96% and offspring production by 89% compared to untreated controls, as females collect less pollen and initiate fewer nests after exposure.^[68] Studies from the 2020s have linked sublethal doses of insecticides like flupyradifurone (Sivanto) and fungicides like Quadris Top to decreased pollen collection efficiency by 32% per flower visit and reduced offspring per nest, with combined exposures causing hyperactivity and further reproductive impairment.^[69] These effects stem from the bees' ground-nesting behavior, which exposes them directly to contaminated soil during nesting.^[70] Climate change poses additional risks through phenological mismatches and drought effects on nesting. Warmer temperatures can shift squash bloom timing relative to bee emergence, resulting in imperfect synchrony where fewer than 50% of Peponapis pruinosa (syn. Eucera pruinosa) emerge during the crop's pollination window, potentially limiting reproductive success to 3.6 offspring per nest on average.^[30] As a northern range species, squash bees may experience worsening desynchronization with continued warming, reducing overlap between adult activity and floral availability.^[30] Droughts, increasingly frequent due to climate change, harden soils and reduce moisture needed for excavation and larval development in ground nests, contributing to broader declines in solitary bee populations.^[71]^[67]

Management and protection

Conservation efforts for squash bees emphasize agricultural practices that support nesting and foraging while minimizing exposure to pesticides. Farmers can protect nesting sites by maintaining untreated field margins and avoiding deep tillage, as squash bees nest shallowly (6-12 inches) beneath host plants like squash vines; no-till practices have been shown to support three times higher bee densities compared to tilled fields. Integrated pest management (IPM) strategies are crucial, including scouting for pests, crop rotation, and sanitation to reduce reliance on broad-spectrum insecticides; when pesticides are necessary, applications should occur in the evening when bees are inactive, and systemic neonicotinoids like imidacloprid should be avoided due to their severe impacts on reproduction. These practices not only preserve squash bee populations but also enhance pollination efficiency in cucurbit crops. Recent surveys, such as the Oregon State University Squash Bee Survey, have documented range expansion into new states like Oregon as of 2025, underscoring the importance of continued monitoring.^[72] Research initiatives focus on monitoring and understanding population dynamics to inform conservation. Programs such as the Oregon State University Squash Bee Survey engage volunteers to document sightings via mobile apps, contributing to distribution data across agricultural landscapes. Similarly, the Lopez-Uribe Lab's Squash Pollinator Search collects weekly observations of bee visits to Cucurbita flowers, aiding in assessments of local abundance and diversity. The U.S. Geological Survey's Native Bee Inventory and Monitoring Program includes surveys that capture squash bee occurrences as part of broader native bee assessments, providing baseline data for long-term trends. While captive rearing trials remain limited for this ground-nesting solitary species, ongoing field-based studies explore reproductive responses to environmental stressors. Policy measures integrate squash bees into broader pollinator protections without granting them formal endangered status. The species holds a global rank of G5 (secure) from NatureServe, indicating no widespread decline warranting listing, though some regional populations are monitored as watchlist species due to habitat pressures. Squash bees are referenced in the proposed Saving America's Pollinators Act (H.R. 4277, 118th Congress, 2023-2024), which advocated for reduced neonicotinoid use to safeguard native pollinators including Peponapis pruinosa but did not become law. U.S. Farm Bill programs, such as the Conservation Reserve Program and Environmental Quality Incentives Program (extended through at least 2026), fund habitat enhancements like pollinator-friendly cover crops and buffer strips that indirectly benefit squash bees by preserving nesting areas in agricultural settings.

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Bee body mass, pathogens and local climate influence heat tolerance
May 3, 2024 · Females are larger than males and collect pollen for offspring in the morning through midday. The females nest underground, which buffers them ...
[22]
Exploring Bee Species: 14 Types of Bees and Bee Varieties
Mar 25, 2022 · Squash Bee (Xenoglossa fulva) Size: The body length of a female is 0.55 – 0.71 inches. The body length of a male is 0.55 – 0.63 inches. ...
[23]
Xenoglossa | Minnesota Native Bees
In Minnesota, there are three described species within the genus Xenoglossa. All three species nest in the ground, have solitary nests, and are oliogolectic.
[24]
Genus Xenoglossa - Large Squash Bees - BugGuide.Net
Genus Xenoglossa - Large Squash Bees · Numbers. 5 spp. · Identification. Robust and hairy. · Range. nearctic (US to Costa Rica: map)(1) · Food. Collects pollen only ...
[25]
Differentiation and Range Expansion of North American Squash ...
Because the genus Peponapis is predominately associated with mesophytic ... The squash and gourd bees—genera Peponapis Robertson and Xenoglossa Smith ...Missing: etymology | Show results with:etymology
[26]
Phenological synchrony between the hoary squash bee (Eucera ...
Among this important group of bees, the solitary hoary squash bee (Eucera (Peponapis) pruinosa (Say, 1837)) is one of the most well understood ground-nesting ...
[27]
[PDF] Fact Sheet - MSU Vegetable Entomology
On a seasonal basis, squash bees develop from pre-pupa into active adults at about the same time as Cucurbita crops begin to bloom in Ontario. On a daily basis ...
[28]
Nest Construction and Life History of the Eastern Cucurbit Bee ... - jstor
Peponapis pruinosa. Structure of nest (diagrammatic). have never found a larva in an open cell, but have often found un hatched eggs in closed cells. In ...
[29]
The Pollination Efficiency of the Squash Bee (Peponapis pruinosa ...
The preference of male Peponapis for sta minate flowers is probably a result of the much higher probability of their encountering a receptive female there with ...
[30]
Assessment of risk to hoary squash bees (Peponapis pruinosa) and ...
Aug 14, 2019 · We provide the first probabilistic risk assessment of exposure to systemic insecticides in soil for ground-nesting bees.Missing: instars | Show results with:instars
[31]
Building a Home, Squash Bee Style - Pollinator Partnership Canada
Oct 22, 2020 · On top of those provisions, she lays an egg and seals the cell with a mud door. The egg subsequently hatches and develop into a larva which ...Missing: reproduction polyandry<|separator|>
[32]
Squash Bees; Genus Peponapis - NATIVE BEES
Squash bees' thoraxes are covered with light-gold hairs. Their legs are a dark reddish-brown. Both males and females have big noses, as shown in the photos here ...<|separator|>
[33]
Peponapis pruinosa - NatureServe Explorer
Squash Bee - Natureserve Global Rank: G5: This species is widespread, known from many localities, and does not appear to be declining.
[34]
Squash bees (Peponapis sp.) and (Xenoglossa sp.)
Jun 5, 2018 · Peponapis and Xenoglossa bees are commonly called squash bees because of their unique relationship with squash, pumpkins, and related plants.
[35]
[PDF] Squash Bees 2 Pager 2025 - Pollinator Partnership
Squash bees are solitary bees that specialize in pollinating squash and gourds, are active early, and are vital for providing pollination for these crops.Missing: taxonomy | Show results with:taxonomy
[36]
Squash Bee Insect Facts - A-Z Animals
Squash Bee Physical Characteristics. Color. Brown; Yellow; Black; Gold; Orange; Caramel; Ginger. Skin Type: Exoskeleton; Top Speed: -27 mph; Lifespan: 3 weeks - ...
[37]
https://www.backyardecology.net/squash-bees/
[38]
(PDF) The Initial Domestication of Cucurbita pepo in the Americas ...
Squash seeds, peduncles, and fruit rind fragments from Archaic period stratigraphic zones of Guilá Naquitz cave in Oaxaca, Mexico, are assigned to Cucurbita ...
[39]
Pollen Records of Guila Naquitz Cave - jstor
6000-8000 B.C. A generally cooler yet more xeric climate apparently prevailed, as had been suggested by faunal analyses at Coxcatlan Cave near Tehuacan ...Missing: BCE | Show results with:BCE
[40]
Crop domestication facilitated rapid geographical expansion of a ...
Jun 29, 2016 · 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 ...<|control11|><|separator|>
[41]
Experimental Introduction of Squash Bees (Peponapis) to Improve ...
The present article brings the story up to date, describing experimental transport of pre-nesting females within California, and from California to Hawaii.
[42]
Squash Bee Overview : USDA ARS
Jun 26, 2018 · The squash and gourd bees -- genera Peponapis Robertson and Xenoglossa Smith -- inhabiting America north of Mexico (Hymenoptera: Apoidea).
[43]
Bees Use Vibration to Aid Pollen Collection from Non-Poricidal ...
use of vibratory pollen harvesting (buzz pollination) by bees from non-poricidally dehiscent ... bee females (e.g., Peponapis pruinosa and P. timberlakei) did not ...
[44]
Squash Bees: Specialist Pollinators of Cucurbit Plant
Jun 27, 2025 · Squash bees are solitary, ground-nesting bees that often construct their nests in bare patches around the edges of cucurbit fields (Figure 2) ...Missing: lekking | Show results with:lekking
[45]
"Foraging behavior, pollinator effectiveness, and management ...
Foraging behavior, pollinator effectiveness, and management potential of the new world squash bees Peponapis pruinosa and Xenoglossa strenua (Apidae: Eucerini) ...
[46]
[PDF] Wild Bee Visitation Rates Exceed Pollination Thresholds in ...
Dec 9, 2019 · 1971. Squash and gourd bees (peponapis, xenoglossa) and the origin of the Cultivated cucurbita. Evolution. 25: 218–234.
[47]
A review of soil tillage impacts on ground-nesting wild bees
Nov 1, 2024 · pruinosa brood cell depths ranging from 9 to 69 cm, but mostly between 13 and 30 cm), or by nesting outside the fields at undisturbed sites, ...
[48]
[PDF] D. Susan Willis Chan - The Atrium - University of Guelph
My research undertaken on farms across Ontario demonstrated that hoary squash bees are the primary pollinators of Ontario's Cucurbita crops.
[49]
Host-parasite interactions between Xenoglossa pruinosa (Apidae ...
Nov 26, 2024 · In cleptoparasitic bees, host aggression and detection avoidance might be the main selective pressures shaping host-parasite interactions.
[50]
Cucurbit specialist, Peponapis pruinosa, is a more effective ...
Previous studies on cultivated squash conducted in agricultural settings have found that the effectiveness of honey bees and squash bees appears more similar ...Missing: efficiency | Show results with:efficiency
[51]
What's That Buzz? A Pollination Syndrome in Your Garden
Jun 9, 2022 · Only about half of all bee species can generate the frequencies required to release pollen ... pollen, including squashes and gourds. Next ...
[52]
Carving Out a Living: Celebrating America's Pumpkin Market
Oct 21, 2025 · In total, U.S. pumpkin farmers produced over $274 million in value in 2024. Illinois led in volume but earned just $21 million because of lower ...Missing: squash | Show results with:squash
[53]
Agriculture that Depends on Honeyees
The value of squash, pumpkin and gourds in the U.S. is estimated at $237.1 million. Honeybees are the most important pollinators for squash, pumpkin and gourds.
[54]
Pollination of Cucurbita spp. (Squash and Pumpkin) Crops in Florida
Feb 22, 2021 · All varieties of squash and pumpkin are highly dependent on insect pollination, without which crop yields are low to nonexistent.
[55]
Wild Bee Visitation Rates Exceed Pollination Thresholds in ...
Dec 9, 2019 · Wild bees supply sufficient pollination in Cucurbita agroecosystems in certain settings; however, some growers continue to stock fields with ...
[56]
Squash bees flourish in response to agricultural intensification
namely ...Missing: dispersal Europe Asia
[57]
https://www.nass.usda.gov/Publications/Todays_Reports/reports/cpvl0225.pdf
[58]
https://edis.ifas.ufl.edu/publication/IN1311
[59]
Chalkbrood - SARE - Sustainable Agriculture Research and Education
The single most destructive disease of cavity-nesting bees is the fungal pathogen chalkbrood. Several chalkbrood species exist, including Ascosphaera ...
[60]
Natural Enemies | Minnesota Native Bees
Philanthus (beewolves) are predators of bees (less often wasps). Philanthus females search for and capture bees on flowers. To avoid detection, these predatory ...
[61]
Crop pollination from native bees at risk from agricultural ...
The decline of native bee pollination services with agricultural intensification resulted from significant reductions in both diversity and total abundance of ...Crop Type And Study Sites · Pollen Deposition · Associated DataMissing: urbanization | Show results with:urbanization<|control11|><|separator|>
[62]
Farming Practices Influence Wild Pollinator Populations on Squash ...
Aug 8, 2025 · No-tillage farms hosted three times higher density of squash bees as farms employing tillage (Shuler et al., 2005) . ... Regard and protect ...Missing: load | Show results with:load
[63]
Native bee habitat restoration: key ecological considerations from ...
This trait makes them particularly vulnerable to population declines if negatively impacted by pesticides ... Climate change contributes to widespread declines ...
[64]
Population decline in a ground-nesting solitary squash bee (Eucera ...
Feb 19, 2021 · Among the most efficient and widely distributed wild pollinators of Cucurbita crops in North America is the hoary squash bee (Eucera (Peponapis) ...
[65]
Single and combined exposure to 'bee safe' pesticides alter ...
Mar 20, 2024 · This study demonstrates that 'low-toxicity' pesticides can adversely affect squash bees under field-realistic exposure, alone or in combination.
[66]
Assessment of risk to hoary squash bees (Peponapis pruinosa ... - NIH
Aug 14, 2019 · Risk assessments for ground-nesting bees should include exposure impacts from pesticides in soil and could use the hoary squash bee as an ...Missing: drought | Show results with:drought
[67]
Factors Contributing to Bee Decline | USU
Many factors have been attributed to bee declines, including habitat loss, improper apiary management, pesticide usage, climate change, pests and pathogens.Missing: vulnerable | Show results with:vulnerable